Molecular phylogeny and historical biogeography of Iranian Murid rodents inferred from mitochondrial cytochrome b gene

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Here we infer the phylogenetic relationships and historical biogeography for the Iranian murid rodents, which consist in 17 species distributed in two subfamilies and six genera. Molecular dating analyses using a relaxed Bayesian molecular clock support the monophyly of Murinae and Gerbillinae and allow to set up a divergence date between them around 18.86 Ma (Million years ago). According to our results, murines may have originated approximately 13.49 Ma and the diversification of most of their evolutionary lineages may have taken place between 10–4 Ma, which is consistent with the fossil record. Our results provide strong support for the tribes Apodemyini and Rattini (Murinae) but the monophyly of the genus Meriones belonging to the Gerbillinae is questioned. Historical biogeographic analysis supports a Paleotropical origin for the Iranian murids, likely found in central and eastern Iran (Desert and Xeric Shrubland ecoregion). From there they dispersed to colonize the Afrotropical, Indomalayan and Palearctic realms. All in all, Iran seems to have acted as a corridor for faunal exchanges between the Afrotropic and Saharo-Arabian realms and the Indomalayan realm as well as between Central Asia and the Mediterranean regions. Murinae Gerbillinae Bayesian molecular clock Divergence time Biogeography Middle East Figures Figure 1 Figure 2 Figure 3 Figure 4 Figure 5 Introduction The Muridae with more than 1383 species is the largest and most evolutionarily successful family of mammals (Martin et al. 2000 ; Aghová et al. 2018 ). Its species are distributed worldwide (except for Antarctica and New Zealand), where they occupy a wide range of habitats from tropical forests to tundra and arid deserts, and have adapted a wide range of lifestyles (Steppan et al. 2004 ). Molecular data have been used for phylogenetic studies of murid rodents and other groups of small mammals with encouraging results (Michaux et al. 2001 ; Ito et al. 2010 ; Fabre et al. 2013 ; Alhajeri et al. 2015 ; Steppan and Schenk 2017 ; Aghová et al. 2018 ; Upham et al. 2019 ; Nicolas et al. 2021 ). These studies have revealed the existence of new lineages, questioned the validity of others or even described higher taxonomic ranks among groups. However, Muridae systematics is not completely understood with numerous changes occurring in the past decades (Ducroz et al. 2001 ; Jansa and Weksler 2004 ; Musser and Carleton 2005 ; Alhajeri et al. 2015 ; Aghová et al. 2018 ; Steppan and Schenk 2017 ), and the relationships within and between its lineages are still poorly known (Martin et al. 2000 ; Michaux et al. 2001 ; Jansa and Weksler 2004 ; Pagès et al. 2016 ; Ge et al. 2021 ; Nicolas et al. 2021 ). Moreover, phylogenetic knowledge of the Muridae family is critical to many areas of science because Muridae species are the most common model organisms for ecological, physiological and experimental research (Hashemian et al. 2017 ; Steppan and Schenk 2017 ; Edrisi et al. 2018 ). The Irano-Anatolian biodiversity hotspot mainly extends through the high elevations of central and eastern Turkey, Armenia, northeast Iraq and Iran. Iran covers about 54% of the surface area of this biodiversity hotspot and five biodiversity hotspots within hotspots have been identified in this country (i.e., Zagros, Azerbaijan, Alborz, Central Alborz and Kopet Dagh-Khorasan; Noroozi et al. 2018 ). Moreover, Iran is at the crossroad of the Palearctic, Oriental and Saharo-Arabian zoogeographic realms (Holt et al. 2013 ). So, the Zagros and Alborz mountains belong to the Palearctic realm, the central regions of the country are placed in the Saharo-Arabian realm, and the southeastern lowlands are situated in the Oriental realm (Zehzad et al. 2002 ; Sindaco et al. 2008 ; Djamali et al. 2011 ). In addition, Iran acts as a corridor between Central Asia and the Indian regions on one side and the Mediterranean and Arabian areas on the other side (Amir Afzali et al. 2018 ; Yousefi et al. 2019 ). To date, 27 murid rodent species are known in Iran (Yousefi et al. 2019 ; Yousefi et al. 2022 ), however, there is lack of information about their historical biogeography and phylogenetic relationships in the region. So, in this study we aim to I) obtain a robust time calibrated phylogeny for murid rodents in Iran; II) elucidate their historical biogeography; and finally, III) based on the obtained results we discuss about phylogeny and biogeography of the group. Material and methods Taxonomic Sampling To investigate the phylogenetic relationships, we have reviewed all literature concerning murid rodents in Iran, and used all available mitochondrial cytochrome b sequences ( cytb ) in the GeneBank. We have used cyt b sequences because they are relevant to the study of deep-time divergence in murid rodents, making this gene an ideal marker (Martin et al. 2000 ; Michaux et al. 2001 ; Steppan et al. 2004 ; Schenk et al. 2013; Alhajeri et al. 2015 ; Aghová et al. 2018 ; Steppan and Schenk 2017 ). Our dataset encompasses 17 out of 27 murid rodent species in Iran, which are distributed in six genera belonging to two subfamilies: The Murinae (eight species) and the Gerbillinae (nine species) (Table 1; Supplementary material). Even though our taxonomic sampling is not complete, it reflects the diversity of murid rodents in Iran (Fig. 1 ). Two cricetids, Sigmodon arizonae Mearns, 1890 and Cricetulus kamensis Satunin, 1903 have been selected as outgroups, as Cricetidae are well supported sister group of Muridae (Steppan and Schenk 2017 ). The tree has been rooted using Sicista armenica Sokolov & Baskevich, 1988 a representative of the more distant family Sminthidae. Phylogenetic analysis Sequences were aligned with the MUSCLE algorithm using the default parameters in MEGA-X (Kumar et al. 2018 ). The phylogenetic analyses were performed through Bayesian inference (BI) using BEAST 2 version 2.7.3. (Bouckaert et al. 2019 ) and Maximum Likelihood (ML) using RAxML 8.2.12 (Kozlov et al. 2019 ). Models of nucleotide substitution for analyses were selected in the jModelTest 2.1.10 (Darriba et al. 2012 ) using the Akaike Information Criterion (AIC). Four Metropolis-coupled Markov Chain Monte Carlo (MC3) were run independently, using different random starting trees, for 10 million generations, sampling every 10000 generations from the posterior distribution. Each run was checked for adequate convergence using Tracer v1.7.0, first independently and then together. All parameter’s effective sample size (ESS) values were confirmed to be higher than 200. Then, we combined the results of log and tree files of independent runs using LogCombiner v.2.7.3. The Maximum Clade Credibility tree (MCC) was obtained using TreeAnnotator 2.7.3 after removal of 20% of the trees as burn-in. To estimate the maximum likelihood tree, analyses were conducted in RAxML 8.2.12, clade support was assessed using a non-parametric bootstrap procedure with 1000 replicates. Nodes supported by bootstrap value ≥ 70 and posterior probability ≥ 0.95 were considered strongly supported (Aghová et al. 2018 ). The mean genetic divergence among evolutionary lineages derived from phylogenetic analyses calculated with MEGA-X (Kumar et al. 2018 ). Divergence time analysis To generate a time-calibrated topology, we used uncorrelated relaxed log normal clock implemented in BEAST 2 version 2.7.3. (Bouckaert et al. 2019 ). The obtained molecular dataset was reduced to include a similar number of sequences from each species level cluster identified in the phylogenetic analyses. Four calibration points were used to infer absolute divergence times (Table 2). Priors for fossil constraints were defined under lognormal and uniform distributions in two separate analyses. For each fossil calibration two independent analyses of 40 million generations were performed in BEAST 2 (one run with lognormal distribution and another run with uniform distribution). Convergence of independent runs was assessed by ESS values greater than 200 for each parameter. The software LogCombiner and TreeAnnotator (included in the BEAST 2 package) were used to generate the maximum clade credibility tree, representing the mean and 95% HPD interval for all nodal ages. Historical biogeography Bayesian Binary Markov Chain Monte Carlo (BBM) method for ancestral state has been performed on the maximum clade credibility tree in RASP 4.3 (Ronquist and Huelsenbeck 2003 ; Ali et al. 2012 ; Yu et al. 2015 ). BBM accommodates phylogenetic uncertainty by averaging the ancestral reconstructions over a sample of user-supplied trees (Sosa et al. 2016 ). The 12800 post-burn-in trees from the Bayesian Inference analyses using BEAST 2 were input into RASP to estimate the probabilities of ancestral areas at each node on the consensus tree. The maximum number of areas allowed for the ancestral ranges were constrained to six. The Markov Chain Monte Carlo (MCMC) chains were run for five million generations and ten independent runs. The reconstructed state was sampled every 1000 generations. The fixed model JC + G (Jukes-Cantor + Gamma) was used. Data for species range distribution were obtained from the Global Biodiversity Information Facility (www.gbif.org) and are shown in Table 3 . Six major biogeographic areas were defined following Olson et al ( 2001 ): A, Desert and Xeric shrubland; B, Flooded Grassland and Savannas; C, Temperate Coniferous Forests; D, Montane Grasslands and Shrublands; E, Temperate Broadleaf and Mixed Forests; F, Temperate Grasslands, Savannas and Shrublands (Fig. 2 ). Results Phylogeny and divergence times Phylogenetic analyses have been conducted on 737 bp of the cytb belonging to 17 murid rodent species (Supplementary material). The AIC identified GTR + I + G as the best-fit model for cytb . BI and ML analyses have produced similar topologies with well-supported clades (Fig. 3 ). The main difference rests in the clade support, which is generally higher in the ML analysis. Considering the number of nodes that are supported by posterior probability ≥ 0.95 or bootstrap value ≥ 70, ML analysis yields a slightly more robust topology (25 well-supported nodes) when compared to the BI tree (20 well-supported nodes). The results of the phylogenetic analyses recovered the monophyly of the family Muridae. Its most recent common ancestor (MRCA) originated during the Early Miocene circa 18.86 Ma. The origin of the subfamily Murinae is here set up approximately at 13.49 Ma (Fig. 4 ). Rattus rattus is the most basal taxon of our tree (Fig. 3 ). The next branch splits into two highly supported monophyletic clades ( Mus / Apodemus ) and diverges from Rattus some 13.49 Ma. The divergence between the tribes Murini and Apodemini ( Mus / Apodemus ) is here set up approximately at 11.91 Ma. The tribe Apodemini, represented here by the genus Apodemus , is well supported and its origin is set up at about 8.84 Ma. Our analysis evidence that Apodemus mystacinus is the most basal species inside this tribe. It is followed successively by Apodemus witherbyi and Apodemus uralensis . The latter one, being sister species to the clade constituted by Apodemus ponticus and Apodemus hyrcanicus , which diverged about 3.55 Ma. The tribe Murini, represented here by the genus Mus , is also well supported. Its most basal taxon is Mus macedonicus , which originated approximately 2.48 Ma. This species is sister species to the clade constituted by Mus musculus and Mus domesticus , which diverged about 0.47 Ma. The clade Gerbillinae is well supported and its origin is here set up circa 10.16 Ma (Fig. 4 ). The topology of our tree shows Gerbillus nanus as the most basal taxon inside gerbillines (Fig. 3 ). This taxon is sister species of the clade that includes Meriones spp. and Rhombomys opimus. Our results, concur with those obtained by Steppan and Schenk ( 2017 ), according to which Rhombomys opimus could in fact belong to the genus Meriones . If so, the clade Meriones would be monophyletic according to our results and its origin could be set up circa 7.08 Ma. Our results showed that the clade Meriones is well supported and split into two main evolutionary lineages ( M. libycus + more derived Meriones ) and ( M. persicus + more derived Meriones ). Rhombomys opimus is sister species of Meriones vinogradovi (Fig. 3 ). Overall, the crown age for the Muridae family is here estimated as being Early Miocene in time whereas most of the speciation events in the family seems to have occurred during the Late Miocene and Pliocene (Figs. 3 and 4 ). Based on cytb the mean genetic divergence among murid rodent species in this study was 22% and the mean genetic divergence between Murinae and Gerbillinae was 25%. Maximum genetic divergence was revealed between Rattus rattus and Rhombomy opimus (29%) whereas the minimum between Mus musculus and Mus domesticus (2%), (Table 4). Historical biogeography The results of our historical biogeographical analyses (Fig. 5 ) showed that the divergence within the murid rodent species in Iran (node 33) may have originated in central and east of Iran (Desert and Xeric Shrubland ecoregion), from where they may have dispersed towards more temperate areas (Temperate Coniferous Forest; Temperate Broadleaf and Mixed Forest; and Temperate Grasslands, Savannas and Shrublands) (Fig. 5 c). Dispersal events mostly occurred within ecoregions (mainly in the Temperate Broadleaf and Mixed Forests ecoregion) than between ecoregions. Gerbillinae in Iran (node 24) may have originated in the central and east areas of the country (Desert and Xeric Shrubland ecoregion), from where dispersal events led to the colonization of their current ranges. Murinae (stemming from node 32) seems to have been originated in central, west and north of Iran (Desert and Xeric Shrubland, Temperate Coniferous forests, Temperate Broadleaf and Mixed Forests, Temperate Grasslands, Savannas and Shrublands ecoregions), and from where dispersal events led to the colonization of current ranges. Overall, Murinae and Gerbillinae ancestors through 72 dispersal events expanded their ranges and diverged into different evolutionary lineages. The majority of the dispersal events occurred during the Pliocene and subsequently this family experienced the highest diversification (Figs. 4 and 5 b). Most of the dispersal events took place from the central and eastern regions of Iran (Desert and Xeric Shrubland) towards the north, west and northwestern parts where the ecoregions were predominantly Temperate Coniferous Forests, Temperate Broadleaf and Mixed Forests, and Temperate Grasslands, Savannas and Shrublands. Dispersal events were less numerous from north and northwest parts, which were dominated by Temperate Coniferous Forests, Temperate Grasslands, Savannas and Shrublands, towards the southwest of Iran to Flooded Grassland and Savannas. Moreover, most of the speciation events have occurred in the west and north of Iran (14 speciation events occurred in Temperate Broadleaf and Mixed Forests ecoregion) whereas the least number of speciation events took place in southwest of Iran (5 speciation events in the ecoregion corresponding to Flooded Grassland and Savannas). Discussion Our dataset represented more than 62% of known murid rodent species in Iran. We used four fossil calibrations for estimating divergence dates, using the Gerbil and Mus as prior for estimating divergence times led to an overestimation of the age of the family Muridae, inferring median ages for the origin of the family that were 21.80 and 28.33 Ma, respectively. In the same way, phylogenetic runs with uniform distribution prior, also led to overestimation of the ages, for instance when we used Apodemus fossil calibration with uniform distribution the median age for the origin of the family was set up to circa 22 Ma. The most accurate results were obtained using Apodemus and Murinae as prior for calibration, applying log normal distribution under uncorrelated relax log normal clock, our results agreed with recent molecular studies that used many fossil calibrations and genes (Steppan et al. 2004 ; Schenk et al. 2013; Kimura et al. 2015; Alhajari et al. 2015; Steppan and Schenk 2017 ; Aghova et al. 2018). We set up here the origin of the family Muridae approximately 18.86 Ma, which is in agreement with the results obtained by Steppan and Schenk ( 2017 ) and Aghová et al ( 2018 ), who inferred a similar age for the origin of Muridae (circa 19 Ma). Our work does not support the monophyly of Meriones except if we consider that Rhombomys belongs in fact to the genus Meriones . These results are in line with those obtained by molecular analyses performed by Alhajeri et al ( 2015 ) and Steppan and Schenk ( 2017 ), in which Meriones may not be a monophyletic group unless some taxonomical rearrangements are done such as the inclusion of Rhombomys in it. So, if we consider Rhombomys as being in fact Meriones , we may point out that this clade diverged into two main lineages. The first one, strongly supported in both BI and ML analyses, includes M. libycus , M. tristrami and M. crasus . Monophyly of this branch has also been recovered from previous molecular studies (Ito et al. 2010 ; Alhajeri et al. 2015 ; Steppan and Schenk 2017 ). The other lineage splits in two clades and it is less well supported. It includes M. persicus as sister species of ( M. meridianus , M. vinogradovi , “ Rh”. opimus ) (Figs. 3 and 4 ). According to Steppan and Schenk ( 2017 ), Rh. opimus and M. persicus are sister species whereas M. meridianus is in the most derived position of the clade. Alhajeri et al ( 2015 ) didn’t recover the monophyly of most of the genera inside Gerbillinae and pointed out the need of major revision of this group, particularly in that concerning the taxonomy of this group based on morphology. According to them the relationships among Rhombomys , Brachiones Thomas, 1925, Psammomys Cretzschmar, 1828 and Meriones were ambiguous with Meriones recovered as monophyletic in their Bayesian Inference but paraphyletic in their Maximum Likelihood analyses. Ito et al ( 2010 ) also found ambiguity in the relationships concerning these four genera. They suggested the non-monophyly of the genus Meriones and their results placed Meriones tamariscinus Pallas, 1773 as sister taxon to the clade comprising Brachiones przewalskii Büchner, 1889 and Rhombomys opimus , and the remaining species belonging to Meriones . They also found that Meriones tamariscinus is more related to Brachiones , Rhombomys and Psammomys than to the other species of Meriones . Ding et al ( 2022 ) also found ambiguous relationships between Meriones and its most closely related genera Rhombomys , Brachiones and Psammomys . According to them Meriones tamariscinus has a closer relationship with Rhombomys than to all other species of Meriones . Both in Brachiones and Rhombomys are monospecific genera for which their phylogenetic relationships are still unclear. It is possible that Meriones tamariscinus does not belong to the genus Meriones and that Rhombomys may be synonymized to Meriones in a near future. Overall, our topology and divergence dates for the main lineages are consistent with previous results of recent molecular studies that includes a dense sampling of gerbillines (Ito et al. 2010 ; Alhajari et al. 2015; Ding et al. 2022 ) and with those from Steppan and Schenk ( 2017 ) that includes 900 muroid species. During the Early Miocene (c. 20 Ma), the counter-clockwise rotation of the Afro-Arabian plate resulted in the collision between this plate and Eurasia (Akıncı et al. 2016 ) and results in the closure of the Tethys and the emergence of a land-bridge (the Gomphotherium land bridge) between them that enabled major continental faunal exchanges and led to substantial changes in global climate. The formation of this land-bridge prompted several independent phases of dispersals instead of one (Mein 2003 ; Sen 2013 ). However, the beginning of the main faunal interchange between both continents took place in the Early Miocene (approximately 20–19 Ma) but was followed by additional dispersal events during the Middle and Late Miocene (e.g., Murinae and Gerbillinae) as is evidenced by palaeontological data (Geraads 2001; Jacobs and Flynn 2005 ; Wessels 2009 ; López-Antoñanzas et al. 2013 , 2015 , 2019 ). According to our results, murines originate around 13.49 Ma (Fig. 4 ). The earliest fossil has been found in Pakistan some 14 Ma whereas their first record in Africa and Europe dated from circa 12 and 11 Ma respectively. So murines probably originated in southeast Asia where they started to diversify and to disperse to Africa, Eurasia and East Asia (López-Antoñanzas et al. 2019 and references there in). Our results evidence that gerbils originated at the beginning of the Late Miocene. The first record from an extant genus is Gerbilliscus Thomas, 1897 formerly Tatera Lataste, 1882, from the Late Miocene of Kenya and Ethiopia (Winkler et al. 2010 ). The origin of the Gerbillinae is still unknown and despite the fact that are considered to have evolved from the Myocricetodontinae (Winkler et al. 2010 ), no phylogenetic analyses at specific level including their potential ancestors (e.g., Abudhabia de Bruijn & Whybrow, 1994, Paradakkamys Lindsay, 1988, Myocricetodon Lavocat, 1952) has been carried out. This group of rodents is supposed to have African origins from where they dispersed to Asia at several times (Chevret and Dobigny 2005 ) Generally, Iran acts as a bridge for exchanging between the Afro-Arabian region and Eurasia and between Central Asia and the Mediterranean regions (Darvish et al. 2012 ; Amir Afzali et al. 2017 ; Amir Afzali et al. 2018 ; Yousefi et al. 2019 ). Our study showed most of the murid rodents of Iran resulted from the westward expansion of an ancestry during the Miocene and most of the speciation events in this family in Iran occurred during the Late Miocene and Pliocene, when Iran experienced orogeny events. The active geology of Iran, resulting from collisions of the Arabian with the Iranian plate that has led to rapid isolation of areas and changing hydrological networks (Allen et al. 2004 ; Hatzfeld et al. 2010 ). The rising of mountains ranges in Iran accelerated during the Middle to Late Miocene (15–5 Ma). This may have promoted allopatric speciation (Ahmadzadeh et al. 2013 ; Mee and Moore 2014 ), which according to our analyses would have been the result of dispersal events. Also, High topographic complexity in the mountains probably causes high habitat diversity and thus a large number of local niche spaces. This is expected to foster adaptation to different niches (i.e., ecological speciation) as well as to create local refugia for species during climatic fluctuations, reducing extinction risks (Ashcroft et al. 2012 ; Manafzadeh et al. 2017 ; Hashemzadeh Segherloo et al. 2018 ). Most of the speciation events inside murids occurred in west and north of Iran where the Alborz and Zagros mountains extend along. These mountains are two hotspots for Iran and act as endemism centers for different taxonomic groups (Noroozi et al. 2018 ). Besides, there is evidence from paleoclimatic and phylogeographic modeling that have highlighted Alborz and Zagros mountains acting as glacial refugia for species of insects, birds, reptiles and mammals during past climatic oscillations (Eskandarzadeh et al. 2018 ). Conclusion Here, we investigate the phylogenetic relationships and the historical biogeography of Iranian Muridae, providing a time calibrated phylogeny for this family of rodents in this region. Our results based on mitochondrial cytochrome b and a reduced number of species are mostly consistent with those based on several nuclear and mitochondrial markers (even up to 15 markers) and a much dense sampling of taxa. Our analyses evidence a time of origin of Muridae about 18.86 Ma. Iranian murids most likely originated in central and eastern Iran and mountain regions, particularly in Zagros and Alborz. These mountain ranges may have been center of speciation for this family of rodents in Iran by providing different ecological niches and local refugia during climate oscillations. Multiple colonization events may have taken place in Iran, which acted, repeatedly, as a corridor for faunal dispersal between Afro-Arabia and Eurasia and between Central Asia and the Mediterranean regions. Declarations Supplementary Information The online version contains supplementary material available at https://doi.org/XXX/XXXXXX. Acknowledgments In memory of Professor Jamshid Darvish, who founded animal systematics in Iran. Author contributions YAA has designed the study. Material preparation, data collection and analysis have been performed by YAA. The manuscript has been written by YAA and RLA, who supervised this study. All authors have red and approved the manuscript. Funding This research received support from the French National Research Agency (project RoMa ANR-22-CE02) Data availability All genetic sequences used in this study are publicly available in the GenBank. Also, all data analyzed during this study are included in this published article and its supplementary information files. Conflict of interest The authors have no competing interests to declare that are relevant to the content of this article. Ethics approval Not applicable. 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Family Subfamily Genus Species Muridae Murinae Apodemus Kaup, 1829 Apodemus hyrcanicus Vorontsov, Boyeskorov & Mezhzherin, 1992 A podemus mystacinus Danford & Alston, 1877 Apodemus ponticus Sviridenko, 1936 Apodemus uralensis Pallas, 1811 Apodemus witherbyi Thomas, 1902 Mus Clerck, 1757 Mus musculus Linnaeus, 1758 Mus domesticus Schwarz & Schwarz, 1943 Mus macedonicus Petrov & Ruzic, 1983 Rattus Fischer de Waldheim, 1803 Rattus rattus Linnaeus, 1758 Gerbillinae Gerbillus Desmarest, 1804 Gerbillus nanus Blanford, 1875 Meriones Illiger, 1811 Meriones crassus Sundevall, 1842 Meriones libycus Lichtenstein, 1823 Meriones meridianus Pallas, 1773 Meriones persicus Blanford, 1875 Meriones tristrami Thomas, 1892 Meriones vinogradovi Heptner, 1931 Rhombomys Wagner, 1841 Rhombomys opimus Lichtenstein, 1823 Table 2 Fossil calibrations used in this work for dating analyses. All ages are in million years before present. Taxon Min Age Max Age Log-StDev Offset Reference Apodemus 5.3 7.2 0.515 4.871 Steppan and Schenk 2017 Murinae 12.1 14.05 0.885 9.767 Steppan and Schenk 2017 Mus 5.3 7.2 0.515 4.871 Steppan and Schenk 2017 Gerbil 16 23.7 1.25 15.868 Schenk et al. 2013 Table 3 Murid rodent species range in terrestrial ecoregions of Iran. Species Ecoregions A B C D E F Mus domesticus * * * * * * Mus musculus * * * * * * Mus macedonicus * * * * * * Rattus rattus * * * Apodemus mystacinus * * * * Apodemus ponticus * * * * Apodemus witherbyi * * * * Apodemus hyrcanicus * Apodemus uralensis * * * * Gerbillus nanus * Rhombomys opimus * * * Meriones persicus * * * * * * Meriones vinogradovi * * * Meriones tristrami * * * * * * Meriones libycus * * * * * * Meriones meridianus * * Meriones crasus * * * * * * Table 4 is available in the Supplementary Files section. Supplementary Files Table4.doc Table 4 Percentage of mean genetic divergence between murid rodents based on cytb . SupplementaryMaterial.rar Cite Share Download PDF Status: Published Journal Publication published 12 Jan, 2024 Read the published version in Mammalian Biology → Version 1 posted Reviewers agreed at journal 02 Aug, 2023 Reviewers invited by journal 01 Aug, 2023 Editor assigned by journal 21 Jul, 2023 First submitted to journal 19 Jul, 2023 You are reading this latest preprint version Research Square lets you share your work early, gain feedback from the community, and start making changes to your manuscript prior to peer review in a journal. As a division of Research Square Company, we’re committed to making research communication faster, fairer, and more useful. We do this by developing innovative software and high quality services for the global research community. Our growing team is made up of researchers and industry professionals working together to solve the most critical problems facing scientific publishing. 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-3186974","acceptedTermsAndConditions":true,"allowDirectSubmit":false,"archivedVersions":[],"articleType":"Research Article","associatedPublications":[],"authors":[{"id":223206609,"identity":"69dc23a3-bde9-494d-8aed-44260d2b83ed","order_by":0,"name":"Yaser Amir Afzali","email":"","orcid":"","institution":"University of Tehran College of Science","correspondingAuthor":false,"submittingAuthor":false,"prefix":"","firstName":"Yaser","middleName":"Amir","lastName":"Afzali","suffix":""},{"id":223206610,"identity":"f50ac717-48ae-49b0-a0c8-e85689ac9064","order_by":1,"name":"Raquel López-Antoñanzas","email":"data:image/png;base64,iVBORw0KGgoAAAANSUhEUgAAAZAAAAAyAQMAAABI0h/eAAAABlBMVEX///8AAABVwtN+AAAACXBIWXMAAA7EAAAOxAGVKw4bAAABX0lEQVRIie3RMUvDQBQH8BcCcQm99YWW+hVOCilC8bOcBJIlolAoCiVUhLpI5xb8EHUpHVMOMtVVChVBCp0cIkLJUKovaROtFlwF8x8uj0t+vHc5gDx5/mzOoPRjr5wWpYIaP2rbrznoaYlpUUlWH0DXEmL/To5bGYEtUr2WAwyJsOKVDKH56FExf46antN/kMHsdVjTtT01CEE8pYOO7YbRJWLcBoJDMEcqqhwDedKf2g4fjW0aTLO6IOrZHK5Z1InwicutpSaRT4SJXPOJuCaO2pKIXgElEilhL2ZxmZDT0IdVTJwFipXn8DV5J8LeAERGkLrAuovSUtoxSb5UxYb4cRd1i8zrhzcc47NQ9440el23YVx25EFvatt4P7bis1RIfA5m3U2i81qZftQshIVkDJ2BES28/cLUCvBieFRmTM4gzEhyKcltIOyK0toU4usu/3aHu0mePHny/ON8ANSbfVrssmbZAAAAAElFTkSuQmCC","orcid":"https://orcid.org/0000-0003-3025-0476","institution":"Institute of Evolutionary Sciences: Institut des sciences de l'evolution de Montpellier","correspondingAuthor":true,"submittingAuthor":false,"prefix":"","firstName":"Raquel","middleName":"","lastName":"López-Antoñanzas","suffix":""}],"badges":[],"createdAt":"2023-07-20 05:00:04","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-3186974/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-3186974/v1","draftVersion":[],"editorialEvents":[{"content":"https://doi.org/10.1007/s42991-023-00390-3","type":"published","date":"2024-01-12T15:01:21+00:00"}],"editorialNote":"","failedWorkflow":false,"files":[{"id":41180744,"identity":"45340956-1747-4e01-a70e-3378e6864328","added_by":"auto","created_at":"2023-08-07 13:46:37","extension":"png","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":1872454,"visible":true,"origin":"","legend":"\u003cp\u003eSampling localities of Iranian murid rodent species used in this study.\u003c/p\u003e","description":"","filename":"Fig1Map.png","url":"https://assets-eu.researchsquare.com/files/rs-3186974/v1/54c171943f1c5728c76c94e4.png"},{"id":41180745,"identity":"dfb0ab8d-0fc9-4472-aeed-b9479c32c945","added_by":"auto","created_at":"2023-08-07 13:46:37","extension":"png","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":1078452,"visible":true,"origin":"","legend":"\u003cp\u003eTerrestrial ecoregions of Iran.\u003c/p\u003e","description":"","filename":"Fig2iranecoregion.png","url":"https://assets-eu.researchsquare.com/files/rs-3186974/v1/6d75cd7a9ce8940c42136026.png"},{"id":41180747,"identity":"5205830b-e25e-4edf-b7f2-2b8e232186f5","added_by":"auto","created_at":"2023-08-07 13:46:38","extension":"png","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":225045,"visible":true,"origin":"","legend":"\u003cp\u003eMaximum clade credibility tree of Muridae from Bayesian inference molecular data. Values at the nodes correspond to the posterior probabilities and bootstrap support, respectively.\u003c/p\u003e","description":"","filename":"Fig3BItree.png","url":"https://assets-eu.researchsquare.com/files/rs-3186974/v1/ce7d826acd503d6ff556310c.png"},{"id":41180749,"identity":"4c49e65f-4b88-4f65-abdf-d642984dc8b8","added_by":"auto","created_at":"2023-08-07 13:46:38","extension":"png","order_by":4,"title":"Figure 4","display":"","copyAsset":false,"role":"figure","size":175164,"visible":true,"origin":"","legend":"\u003cp\u003eTime-calibrated tree of Muridae rodents. Mean divergence times estimated using a relaxed molecular clock model on the available mitochondrial cytochrome b sequences with 4 fossil priors. The ages correspond to million years before present.\u003c/p\u003e","description":"","filename":"Fig4timetree.png","url":"https://assets-eu.researchsquare.com/files/rs-3186974/v1/65f65f5a49fd2117976a2c7a.png"},{"id":41182008,"identity":"1c373c6f-90d7-498e-bc91-8d35a5e99c58","added_by":"auto","created_at":"2023-08-07 13:54:37","extension":"png","order_by":5,"title":"Figure 5","display":"","copyAsset":false,"role":"figure","size":1209354,"visible":true,"origin":"","legend":"\u003cp\u003eBBM (Bayesian Binary MCMC) analyses implemented in RASP. a) Biogeographic ecoregions of Iran; b) Diagram showing the frequency of various ancestral origins (y-axis) against time (x-axis); c) Historical biogeography of Muridae, pie charts on each node depict the relative probabilities of ancestral ranges. Black areas account for phylogenetic uncertainty. Outgroup taxa have been pruned from the tree.\u003c/p\u003e","description":"","filename":"Fig5Historicalbiogeography.png","url":"https://assets-eu.researchsquare.com/files/rs-3186974/v1/99dd8b3a9607ea985b38db20.png"},{"id":49629031,"identity":"33590a6c-5a85-401f-ab72-4cb6041a5e0d","added_by":"auto","created_at":"2024-01-15 15:10:06","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":2343975,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-3186974/v1/bcbcb2ce-0dfc-41a9-896a-8f325ee118a4.pdf"},{"id":41180743,"identity":"4bf6347e-6ca9-4339-9c51-92448ba7d16e","added_by":"auto","created_at":"2023-08-07 13:46:37","extension":"doc","order_by":1,"title":"","display":"","copyAsset":false,"role":"supplement","size":66048,"visible":true,"origin":"","legend":"\u003cp\u003e\u003cstrong\u003eTable 4 \u003c/strong\u003ePercentage of mean genetic divergence between murid rodents based on \u003cem\u003ecytb\u003c/em\u003e.\u003c/p\u003e","description":"","filename":"Table4.doc","url":"https://assets-eu.researchsquare.com/files/rs-3186974/v1/56c2232c40bb72e277c1627e.doc"},{"id":41182007,"identity":"659342b8-8800-4ee8-ad53-6cee649ab32d","added_by":"auto","created_at":"2023-08-07 13:54:37","extension":"rar","order_by":2,"title":"","display":"","copyAsset":false,"role":"supplement","size":31794,"visible":true,"origin":"","legend":"","description":"","filename":"SupplementaryMaterial.rar","url":"https://assets-eu.researchsquare.com/files/rs-3186974/v1/394dae84d38694371a101ddf.rar"}],"financialInterests":"","formattedTitle":"Molecular phylogeny and historical biogeography of Iranian Murid rodents inferred from mitochondrial cytochrome b gene","fulltext":[{"header":"Introduction","content":"\u003cp\u003eThe Muridae with more than 1383 species is the largest and most evolutionarily successful family of mammals (Martin et al. \u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e2000\u003c/span\u003e; Aghov\u0026aacute; et al. \u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). Its species are distributed worldwide (except for Antarctica and New Zealand), where they occupy a wide range of habitats from tropical forests to tundra and arid deserts, and have adapted a wide range of lifestyles (Steppan et al. \u003cspan citationid=\"CR50\" class=\"CitationRef\"\u003e2004\u003c/span\u003e). Molecular data have been used for phylogenetic studies of murid rodents and other groups of small mammals with encouraging results (Michaux et al. \u003cspan citationid=\"CR39\" class=\"CitationRef\"\u003e2001\u003c/span\u003e; Ito et al. \u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e2010\u003c/span\u003e; Fabre et al. \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2013\u003c/span\u003e; Alhajeri et al. \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2015\u003c/span\u003e; Steppan and Schenk \u003cspan citationid=\"CR51\" class=\"CitationRef\"\u003e2017\u003c/span\u003e; Aghov\u0026aacute; et al. \u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; Upham et al. \u003cspan citationid=\"CR52\" class=\"CitationRef\"\u003e2019\u003c/span\u003e; Nicolas et al. \u003cspan citationid=\"CR41\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). These studies have revealed the existence of new lineages, questioned the validity of others or even described higher taxonomic ranks among groups. However, Muridae systematics is not completely understood with numerous changes occurring in the past decades (Ducroz et al. \u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e2001\u003c/span\u003e; Jansa and Weksler \u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e2004\u003c/span\u003e; Musser and Carleton \u003cspan citationid=\"CR40\" class=\"CitationRef\"\u003e2005\u003c/span\u003e; Alhajeri et al. \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2015\u003c/span\u003e; Aghov\u0026aacute; et al. \u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; Steppan and Schenk \u003cspan citationid=\"CR51\" class=\"CitationRef\"\u003e2017\u003c/span\u003e), and the relationships within and between its lineages are still poorly known (Martin et al. \u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e2000\u003c/span\u003e; Michaux et al. \u003cspan citationid=\"CR39\" class=\"CitationRef\"\u003e2001\u003c/span\u003e; Jansa and Weksler \u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e2004\u003c/span\u003e; Pag\u0026egrave;s et al. \u003cspan citationid=\"CR44\" class=\"CitationRef\"\u003e2016\u003c/span\u003e; Ge et al. \u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Nicolas et al. \u003cspan citationid=\"CR41\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). Moreover, phylogenetic knowledge of the Muridae family is critical to many areas of science because Muridae species are the most common model organisms for ecological, physiological and experimental research (Hashemian et al. \u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e2017\u003c/span\u003e; Steppan and Schenk \u003cspan citationid=\"CR51\" class=\"CitationRef\"\u003e2017\u003c/span\u003e; Edrisi et al. \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e2018\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eThe Irano-Anatolian biodiversity hotspot mainly extends through the high elevations of central and eastern Turkey, Armenia, northeast Iraq and Iran. Iran covers about 54% of the surface area of this biodiversity hotspot and five biodiversity hotspots within hotspots have been identified in this country (i.e., Zagros, Azerbaijan, Alborz, Central Alborz and Kopet Dagh-Khorasan; Noroozi et al. \u003cspan citationid=\"CR42\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). Moreover, Iran is at the crossroad of the Palearctic, Oriental and Saharo-Arabian zoogeographic realms (Holt et al. \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2013\u003c/span\u003e). So, the Zagros and Alborz mountains belong to the Palearctic realm, the central regions of the country are placed in the Saharo-Arabian realm, and the southeastern lowlands are situated in the Oriental realm (Zehzad et al. \u003cspan citationid=\"CR59\" class=\"CitationRef\"\u003e2002\u003c/span\u003e; Sindaco et al. \u003cspan citationid=\"CR48\" class=\"CitationRef\"\u003e2008\u003c/span\u003e; Djamali et al. \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e2011\u003c/span\u003e). In addition, Iran acts as a corridor between Central Asia and the Indian regions on one side and the Mediterranean and Arabian areas on the other side (Amir Afzali et al. \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; Yousefi et al. \u003cspan citationid=\"CR57\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). To date, 27 murid rodent species are known in Iran (Yousefi et al. \u003cspan citationid=\"CR57\" class=\"CitationRef\"\u003e2019\u003c/span\u003e; Yousefi et al. \u003cspan citationid=\"CR56\" class=\"CitationRef\"\u003e2022\u003c/span\u003e), however, there is lack of information about their historical biogeography and phylogenetic relationships in the region. So, in this study we aim to I) obtain a robust time calibrated phylogeny for murid rodents in Iran; II) elucidate their historical biogeography; and finally, III) based on the obtained results we discuss about phylogeny and biogeography of the group.\u003c/p\u003e"},{"header":"Material and methods","content":"\u003cdiv id=\"Sec3\" class=\"Section2\"\u003e \u003ch2\u003eTaxonomic Sampling\u003c/h2\u003e \u003cp\u003eTo investigate the phylogenetic relationships, we have reviewed all literature concerning murid rodents in Iran, and used all available mitochondrial cytochrome \u003cem\u003eb\u003c/em\u003e sequences (\u003cem\u003ecytb\u003c/em\u003e) in the GeneBank. We have used cyt\u003cem\u003eb\u003c/em\u003e sequences because they are relevant to the study of deep-time divergence in murid rodents, making this gene an ideal marker (Martin et al. \u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e2000\u003c/span\u003e; Michaux et al. \u003cspan citationid=\"CR39\" class=\"CitationRef\"\u003e2001\u003c/span\u003e; Steppan et al. \u003cspan citationid=\"CR50\" class=\"CitationRef\"\u003e2004\u003c/span\u003e; Schenk et al. 2013; Alhajeri et al. \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2015\u003c/span\u003e; Aghov\u0026aacute; et al. \u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; Steppan and Schenk \u003cspan citationid=\"CR51\" class=\"CitationRef\"\u003e2017\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eOur dataset encompasses 17 out of 27 murid rodent species in Iran, which are distributed in six genera belonging to two subfamilies: The Murinae (eight species) and the Gerbillinae (nine species) (Table\u0026nbsp;1; Supplementary material). Even though our taxonomic sampling is not complete, it reflects the diversity of murid rodents in Iran (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). Two cricetids, \u003cem\u003eSigmodon arizonae\u003c/em\u003e Mearns, 1890 and \u003cem\u003eCricetulus kamensis\u003c/em\u003e Satunin, 1903 have been selected as outgroups, as Cricetidae are well supported sister group of Muridae (Steppan and Schenk \u003cspan citationid=\"CR51\" class=\"CitationRef\"\u003e2017\u003c/span\u003e). The tree has been rooted using \u003cem\u003eSicista armenica\u003c/em\u003e Sokolov \u0026amp; Baskevich, 1988 a representative of the more distant family Sminthidae.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec4\" class=\"Section2\"\u003e \u003ch2\u003ePhylogenetic analysis\u003c/h2\u003e \u003cp\u003eSequences were aligned with the MUSCLE algorithm using the default parameters in MEGA-X (Kumar et al. \u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). The phylogenetic analyses were performed through Bayesian inference (BI) using BEAST 2 version 2.7.3. (Bouckaert et al. \u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e2019\u003c/span\u003e) and Maximum Likelihood (ML) using RAxML 8.2.12 (Kozlov et al. \u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). Models of nucleotide substitution for analyses were selected in the jModelTest 2.1.10 (Darriba et al. \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e2012\u003c/span\u003e) using the Akaike Information Criterion (AIC). Four Metropolis-coupled Markov Chain Monte Carlo (MC3) were run independently, using different random starting trees, for 10\u0026nbsp;million generations, sampling every 10000 generations from the posterior distribution. Each run was checked for adequate convergence using Tracer v1.7.0, first independently and then together. All parameter\u0026rsquo;s effective sample size (ESS) values were confirmed to be higher than 200. Then, we combined the results of log and tree files of independent runs using LogCombiner v.2.7.3. The Maximum Clade Credibility tree (MCC) was obtained using TreeAnnotator 2.7.3 after removal of 20% of the trees as burn-in. To estimate the maximum likelihood tree, analyses were conducted in RAxML 8.2.12, clade support was assessed using a non-parametric bootstrap procedure with 1000 replicates. Nodes supported by bootstrap value\u0026thinsp;\u0026ge;\u0026thinsp;70 and posterior probability\u0026thinsp;\u0026ge;\u0026thinsp;0.95 were considered strongly supported (Aghov\u0026aacute; et al. \u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). The mean genetic divergence among evolutionary lineages derived from phylogenetic analyses calculated with MEGA-X (Kumar et al. \u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e2018\u003c/span\u003e).\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec5\" class=\"Section2\"\u003e \u003ch2\u003eDivergence time analysis\u003c/h2\u003e \u003cp\u003eTo generate a time-calibrated topology, we used uncorrelated relaxed log normal clock implemented in BEAST 2 version 2.7.3. (Bouckaert et al. \u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). The obtained molecular dataset was reduced to include a similar number of sequences from each species level cluster identified in the phylogenetic analyses. Four calibration points were used to infer absolute divergence times (Table\u0026nbsp;2). Priors for fossil constraints were defined under lognormal and uniform distributions in two separate analyses. For each fossil calibration two independent analyses of 40\u0026nbsp;million generations were performed in BEAST 2 (one run with lognormal distribution and another run with uniform distribution). Convergence of independent runs was assessed by ESS values greater than 200 for each parameter. The software LogCombiner and TreeAnnotator (included in the BEAST 2 package) were used to generate the maximum clade credibility tree, representing the mean and 95% HPD interval for all nodal ages.\u003c/p\u003e \u003c/div\u003e\n\u003ch3\u003eHistorical biogeography\u003c/h3\u003e\n\u003cp\u003eBayesian Binary Markov Chain Monte Carlo (BBM) method for ancestral state has been performed on the maximum clade credibility tree in RASP 4.3 (Ronquist and Huelsenbeck \u003cspan citationid=\"CR46\" class=\"CitationRef\"\u003e2003\u003c/span\u003e; Ali et al. \u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e2012\u003c/span\u003e; Yu et al. \u003cspan citationid=\"CR58\" class=\"CitationRef\"\u003e2015\u003c/span\u003e). BBM accommodates phylogenetic uncertainty by averaging the ancestral reconstructions over a sample of user-supplied trees (Sosa et al. \u003cspan citationid=\"CR49\" class=\"CitationRef\"\u003e2016\u003c/span\u003e). The 12800 post-burn-in trees from the Bayesian Inference analyses using BEAST 2 were input into RASP to estimate the probabilities of ancestral areas at each node on the consensus tree. The maximum number of areas allowed for the ancestral ranges were constrained to six. The Markov Chain Monte Carlo (MCMC) chains were run for five million generations and ten independent runs. The reconstructed state was sampled every 1000 generations. The fixed model JC\u0026thinsp;+\u0026thinsp;G (Jukes-Cantor\u0026thinsp;+\u0026thinsp;Gamma) was used. Data for species range distribution were obtained from the Global Biodiversity Information Facility (www.gbif.org) and are shown in Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e3\u003c/span\u003e. Six major biogeographic areas were defined following Olson et al (\u003cspan citationid=\"CR43\" class=\"CitationRef\"\u003e2001\u003c/span\u003e): A, Desert and Xeric shrubland; B, Flooded Grassland and Savannas; C, Temperate Coniferous Forests; D, Montane Grasslands and Shrublands; E, Temperate Broadleaf and Mixed Forests; F, Temperate Grasslands, Savannas and Shrublands (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e).\u003c/p\u003e"},{"header":"Results","content":"\u003cdiv id=\"Sec8\" class=\"Section2\"\u003e \u003ch2\u003ePhylogeny and divergence times\u003c/h2\u003e \u003cp\u003ePhylogenetic analyses have been conducted on 737 bp of the \u003cem\u003ecytb\u003c/em\u003e belonging to 17 murid rodent species (Supplementary material). The AIC identified GTR\u0026thinsp;+\u0026thinsp;I\u0026thinsp;+\u0026thinsp;G as the best-fit model for \u003cem\u003ecytb\u003c/em\u003e. BI and ML analyses have produced similar topologies with well-supported clades (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003e). The main difference rests in the clade support, which is generally higher in the ML analysis. Considering the number of nodes that are supported by posterior probability\u0026thinsp;\u0026ge;\u0026thinsp;0.95 or bootstrap value\u0026thinsp;\u0026ge;\u0026thinsp;70, ML analysis yields a slightly more robust topology (25 well-supported nodes) when compared to the BI tree (20 well-supported nodes).\u003c/p\u003e \u003cp\u003eThe results of the phylogenetic analyses recovered the monophyly of the family Muridae. Its most recent common ancestor (MRCA) originated during the Early Miocene circa 18.86 Ma. The origin of the subfamily Murinae is here set up approximately at 13.49 Ma (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003e). \u003cem\u003eRattus rattus\u003c/em\u003e is the most basal taxon of our tree (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003e). The next branch splits into two highly supported monophyletic clades (\u003cem\u003eMus\u003c/em\u003e/\u003cem\u003eApodemus\u003c/em\u003e) and diverges from \u003cem\u003eRattus\u003c/em\u003e some 13.49 Ma. The divergence between the tribes Murini and Apodemini (\u003cem\u003eMus\u003c/em\u003e/\u003cem\u003eApodemus\u003c/em\u003e) is here set up approximately at 11.91 Ma. The tribe Apodemini, represented here by the genus \u003cem\u003eApodemus\u003c/em\u003e, is well supported and its origin is set up at about 8.84 Ma. Our analysis evidence that \u003cem\u003eApodemus mystacinus\u003c/em\u003e is the most basal species inside this tribe. It is followed successively by \u003cem\u003eApodemus witherbyi\u003c/em\u003e and \u003cem\u003eApodemus uralensis\u003c/em\u003e. The latter one, being sister species to the clade constituted by \u003cem\u003eApodemus ponticus\u003c/em\u003e and \u003cem\u003eApodemus hyrcanicus\u003c/em\u003e, which diverged about 3.55 Ma. The tribe Murini, represented here by the genus \u003cem\u003eMus\u003c/em\u003e, is also well supported. Its most basal taxon is \u003cem\u003eMus macedonicus\u003c/em\u003e, which originated approximately 2.48 Ma. This species is sister species to the clade constituted by \u003cem\u003eMus musculus\u003c/em\u003e and \u003cem\u003eMus domesticus\u003c/em\u003e, which diverged about 0.47 Ma.\u003c/p\u003e \u003cp\u003eThe clade Gerbillinae is well supported and its origin is here set up circa 10.16 Ma (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003e). The topology of our tree shows \u003cem\u003eGerbillus nanus\u003c/em\u003e as the most basal taxon inside gerbillines (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003e). This taxon is sister species of the clade that includes \u003cem\u003eMeriones\u003c/em\u003e spp. and \u003cem\u003eRhombomys opimus.\u003c/em\u003e Our results, concur with those obtained by Steppan and Schenk (\u003cspan citationid=\"CR51\" class=\"CitationRef\"\u003e2017\u003c/span\u003e), according to which \u003cem\u003eRhombomys opimus\u003c/em\u003e could in fact belong to the genus \u003cem\u003eMeriones\u003c/em\u003e. If so, the clade \u003cem\u003eMeriones\u003c/em\u003e would be monophyletic according to our results and its origin could be set up circa 7.08 Ma. Our results showed that the clade \u003cem\u003eMeriones\u003c/em\u003e is well supported and split into two main evolutionary lineages (\u003cem\u003eM. libycus\u003c/em\u003e\u0026thinsp;+\u0026thinsp;more derived \u003cem\u003eMeriones\u003c/em\u003e) and (\u003cem\u003eM. persicus\u003c/em\u003e\u0026thinsp;+\u0026thinsp;more derived \u003cem\u003eMeriones\u003c/em\u003e). \u003cem\u003eRhombomys opimus\u003c/em\u003e is sister species of \u003cem\u003eMeriones vinogradovi\u003c/em\u003e (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eOverall, the crown age for the Muridae family is here estimated as being Early Miocene in time whereas most of the speciation events in the family seems to have occurred during the Late Miocene and Pliocene (Figs.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003e and \u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eBased on \u003cem\u003ecytb\u003c/em\u003e the mean genetic divergence among murid rodent species in this study was 22% and the mean genetic divergence between Murinae and Gerbillinae was 25%. Maximum genetic divergence was revealed between \u003cem\u003eRattus rattus\u003c/em\u003e and \u003cem\u003eRhombomy opimus\u003c/em\u003e (29%) whereas the minimum between \u003cem\u003eMus musculus\u003c/em\u003e and \u003cem\u003eMus domesticus\u003c/em\u003e (2%), (Table\u0026nbsp;4).\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec9\" class=\"Section2\"\u003e \u003ch2\u003eHistorical biogeography\u003c/h2\u003e \u003cp\u003eThe results of our historical biogeographical analyses (Fig.\u0026nbsp;\u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e5\u003c/span\u003e) showed that the divergence within the murid rodent species in Iran (node 33) may have originated in central and east of Iran (Desert and Xeric Shrubland ecoregion), from where they may have dispersed towards more temperate areas (Temperate Coniferous Forest; Temperate Broadleaf and Mixed Forest; and Temperate Grasslands, Savannas and Shrublands) (Fig.\u0026nbsp;\u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e5\u003c/span\u003ec). Dispersal events mostly occurred within ecoregions (mainly in the Temperate Broadleaf and Mixed Forests ecoregion) than between ecoregions. Gerbillinae in Iran (node 24) may have originated in the central and east areas of the country (Desert and Xeric Shrubland ecoregion), from where dispersal events led to the colonization of their current ranges. Murinae (stemming from node 32) seems to have been originated in central, west and north of Iran (Desert and Xeric Shrubland, Temperate Coniferous forests, Temperate Broadleaf and Mixed Forests, Temperate Grasslands, Savannas and Shrublands ecoregions), and from where dispersal events led to the colonization of current ranges.\u003c/p\u003e \u003cp\u003eOverall, Murinae and Gerbillinae ancestors through 72 dispersal events expanded their ranges and diverged into different evolutionary lineages. The majority of the dispersal events occurred during the Pliocene and subsequently this family experienced the highest diversification (Figs.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003e and \u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e5\u003c/span\u003eb). Most of the dispersal events took place from the central and eastern regions of Iran (Desert and Xeric Shrubland) towards the north, west and northwestern parts where the ecoregions were predominantly Temperate Coniferous Forests, Temperate Broadleaf and Mixed Forests, and Temperate Grasslands, Savannas and Shrublands. Dispersal events were less numerous from north and northwest parts, which were dominated by Temperate Coniferous Forests, Temperate Grasslands, Savannas and Shrublands, towards the southwest of Iran to Flooded Grassland and Savannas. Moreover, most of the speciation events have occurred in the west and north of Iran (14 speciation events occurred in Temperate Broadleaf and Mixed Forests ecoregion) whereas the least number of speciation events took place in southwest of Iran (5 speciation events in the ecoregion corresponding to Flooded Grassland and Savannas).\u003c/p\u003e \u003c/div\u003e"},{"header":"Discussion","content":"\u003cp\u003eOur dataset represented more than 62% of known murid rodent species in Iran. We used four fossil calibrations for estimating divergence dates, using the Gerbil and \u003cem\u003eMus\u003c/em\u003e as prior for estimating divergence times led to an overestimation of the age of the family Muridae, inferring median ages for the origin of the family that were 21.80 and 28.33 Ma, respectively. In the same way, phylogenetic runs with uniform distribution prior, also led to overestimation of the ages, for instance when we used \u003cem\u003eApodemus\u003c/em\u003e fossil calibration with uniform distribution the median age for the origin of the family was set up to circa 22 Ma. The most accurate results were obtained using \u003cem\u003eApodemus\u003c/em\u003e and Murinae as prior for calibration, applying log normal distribution under uncorrelated relax log normal clock, our results agreed with recent molecular studies that used many fossil calibrations and genes (Steppan et al. \u003cspan citationid=\"CR50\" class=\"CitationRef\"\u003e2004\u003c/span\u003e; Schenk et al. 2013; Kimura et al. 2015; Alhajari et al. 2015; Steppan and Schenk \u003cspan citationid=\"CR51\" class=\"CitationRef\"\u003e2017\u003c/span\u003e; Aghova et al. 2018). We set up here the origin of the family Muridae approximately 18.86 Ma, which is in agreement with the results obtained by Steppan and Schenk (\u003cspan citationid=\"CR51\" class=\"CitationRef\"\u003e2017\u003c/span\u003e) and Aghov\u0026aacute; et al (\u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2018\u003c/span\u003e), who inferred a similar age for the origin of Muridae (circa 19 Ma).\u003c/p\u003e \u003cp\u003eOur work does not support the monophyly of \u003cem\u003eMeriones\u003c/em\u003e except if we consider that \u003cem\u003eRhombomys\u003c/em\u003e belongs in fact to the genus \u003cem\u003eMeriones\u003c/em\u003e. These results are in line with those obtained by molecular analyses performed by Alhajeri et al (\u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2015\u003c/span\u003e) and Steppan and Schenk (\u003cspan citationid=\"CR51\" class=\"CitationRef\"\u003e2017\u003c/span\u003e), in which \u003cem\u003eMeriones\u003c/em\u003e may not be a monophyletic group unless some taxonomical rearrangements are done such as the inclusion of \u003cem\u003eRhombomys\u003c/em\u003e in it. So, if we consider \u003cem\u003eRhombomys\u003c/em\u003e as being in fact \u003cem\u003eMeriones\u003c/em\u003e, we may point out that this clade diverged into two main lineages. The first one, strongly supported in both BI and ML analyses, includes \u003cem\u003eM. libycus\u003c/em\u003e, \u003cem\u003eM. tristrami\u003c/em\u003e and \u003cem\u003eM. crasus\u003c/em\u003e. Monophyly of this branch has also been recovered from previous molecular studies (Ito et al. \u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e2010\u003c/span\u003e; Alhajeri et al. \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2015\u003c/span\u003e; Steppan and Schenk \u003cspan citationid=\"CR51\" class=\"CitationRef\"\u003e2017\u003c/span\u003e). The other lineage splits in two clades and it is less well supported. It includes \u003cem\u003eM. persicus\u003c/em\u003e as sister species of (\u003cem\u003eM. meridianus\u003c/em\u003e, \u003cem\u003eM. vinogradovi\u003c/em\u003e, \u0026ldquo;\u003cem\u003eRh\u0026rdquo;. opimus\u003c/em\u003e) (Figs.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003e and \u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003e). According to Steppan and Schenk (\u003cspan citationid=\"CR51\" class=\"CitationRef\"\u003e2017\u003c/span\u003e), \u003cem\u003eRh. opimus\u003c/em\u003e and \u003cem\u003eM. persicus\u003c/em\u003e are sister species whereas \u003cem\u003eM. meridianus\u003c/em\u003e is in the most derived position of the clade. Alhajeri et al (\u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2015\u003c/span\u003e) didn\u0026rsquo;t recover the monophyly of most of the genera inside Gerbillinae and pointed out the need of major revision of this group, particularly in that concerning the taxonomy of this group based on morphology. According to them the relationships among \u003cem\u003eRhombomys\u003c/em\u003e, \u003cem\u003eBrachiones\u003c/em\u003e Thomas, 1925, \u003cem\u003ePsammomys\u003c/em\u003e Cretzschmar, 1828 and \u003cem\u003eMeriones\u003c/em\u003e were ambiguous with \u003cem\u003eMeriones\u003c/em\u003e recovered as monophyletic in their Bayesian Inference but paraphyletic in their Maximum Likelihood analyses. Ito et al (\u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e2010\u003c/span\u003e) also found ambiguity in the relationships concerning these four genera. They suggested the non-monophyly of the genus \u003cem\u003eMeriones\u003c/em\u003e and their results placed \u003cem\u003eMeriones tamariscinus\u003c/em\u003e Pallas, 1773 as sister taxon to the clade comprising \u003cem\u003eBrachiones przewalskii\u003c/em\u003e B\u0026uuml;chner, 1889 and \u003cem\u003eRhombomys opimus\u003c/em\u003e, and the remaining species belonging to \u003cem\u003eMeriones\u003c/em\u003e. They also found that \u003cem\u003eMeriones tamariscinus\u003c/em\u003e is more related to \u003cem\u003eBrachiones\u003c/em\u003e, \u003cem\u003eRhombomys\u003c/em\u003e and \u003cem\u003ePsammomys\u003c/em\u003e than to the other species of \u003cem\u003eMeriones\u003c/em\u003e. Ding et al (\u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e2022\u003c/span\u003e) also found ambiguous relationships between \u003cem\u003eMeriones\u003c/em\u003e and its most closely related genera \u003cem\u003eRhombomys\u003c/em\u003e, \u003cem\u003eBrachiones\u003c/em\u003e and \u003cem\u003ePsammomys\u003c/em\u003e. According to them \u003cem\u003eMeriones tamariscinus\u003c/em\u003e has a closer relationship with \u003cem\u003eRhombomys\u003c/em\u003e than to all other species of \u003cem\u003eMeriones\u003c/em\u003e. Both in \u003cem\u003eBrachiones\u003c/em\u003e and \u003cem\u003eRhombomys\u003c/em\u003e are monospecific genera for which their phylogenetic relationships are still unclear. It is possible that \u003cem\u003eMeriones tamariscinus\u003c/em\u003e does not belong to the genus \u003cem\u003eMeriones\u003c/em\u003e and that \u003cem\u003eRhombomys\u003c/em\u003e may be synonymized to \u003cem\u003eMeriones\u003c/em\u003e in a near future.\u003c/p\u003e \u003cp\u003eOverall, our topology and divergence dates for the main lineages are consistent with previous results of recent molecular studies that includes a dense sampling of gerbillines (Ito et al. \u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e2010\u003c/span\u003e; Alhajari et al. 2015; Ding et al. \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e2022\u003c/span\u003e) and with those from Steppan and Schenk (\u003cspan citationid=\"CR51\" class=\"CitationRef\"\u003e2017\u003c/span\u003e) that includes 900 muroid species.\u003c/p\u003e \u003cp\u003eDuring the Early Miocene (c. 20 Ma), the counter-clockwise rotation of the Afro-Arabian plate resulted in the collision between this plate and Eurasia (Akıncı et al. \u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e2016\u003c/span\u003e) and results in the closure of the Tethys and the emergence of a land-bridge (the \u003cem\u003eGomphotherium\u003c/em\u003e land bridge) between them that enabled major continental faunal exchanges and led to substantial changes in global climate. The formation of this land-bridge prompted several independent phases of dispersals instead of one (Mein \u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e2003\u003c/span\u003e; Sen \u003cspan citationid=\"CR47\" class=\"CitationRef\"\u003e2013\u003c/span\u003e). However, the beginning of the main faunal interchange between both continents took place in the Early Miocene (approximately 20\u0026ndash;19 Ma) but was followed by additional dispersal events during the Middle and Late Miocene (e.g., Murinae and Gerbillinae) as is evidenced by palaeontological data (Geraads 2001; Jacobs and Flynn \u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e2005\u003c/span\u003e; Wessels \u003cspan citationid=\"CR54\" class=\"CitationRef\"\u003e2009\u003c/span\u003e; L\u0026oacute;pez-Anto\u0026ntilde;anzas et al. \u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e2013\u003c/span\u003e, \u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e2015\u003c/span\u003e, \u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). According to our results, murines originate around 13.49 Ma (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003e). The earliest fossil has been found in Pakistan some 14 Ma whereas their first record in Africa and Europe dated from circa 12 and 11 Ma respectively. So murines probably originated in southeast Asia where they started to diversify and to disperse to Africa, Eurasia and East Asia (L\u0026oacute;pez-Anto\u0026ntilde;anzas et al. \u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e2019\u003c/span\u003e and references there in).\u003c/p\u003e \u003cp\u003eOur results evidence that gerbils originated at the beginning of the Late Miocene. The first record from an extant genus is \u003cem\u003eGerbilliscus\u003c/em\u003e Thomas, 1897 formerly \u003cem\u003eTatera\u003c/em\u003e Lataste, 1882, from the Late Miocene of Kenya and Ethiopia (Winkler et al. \u003cspan citationid=\"CR55\" class=\"CitationRef\"\u003e2010\u003c/span\u003e). The origin of the Gerbillinae is still unknown and despite the fact that are considered to have evolved from the Myocricetodontinae (Winkler et al. \u003cspan citationid=\"CR55\" class=\"CitationRef\"\u003e2010\u003c/span\u003e), no phylogenetic analyses at specific level including their potential ancestors (e.g., \u003cem\u003eAbudhabia\u003c/em\u003e de Bruijn \u0026amp; Whybrow, 1994, \u003cem\u003eParadakkamys\u003c/em\u003e Lindsay, 1988, \u003cem\u003eMyocricetodon\u003c/em\u003e Lavocat, 1952) has been carried out. This group of rodents is supposed to have African origins from where they dispersed to Asia at several times (Chevret and Dobigny \u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e2005\u003c/span\u003e)\u003c/p\u003e \u003cp\u003eGenerally, Iran acts as a bridge for exchanging between the Afro-Arabian region and Eurasia and between Central Asia and the Mediterranean regions (Darvish et al. \u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e2012\u003c/span\u003e; Amir Afzali et al. \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e2017\u003c/span\u003e; Amir Afzali et al. \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; Yousefi et al. \u003cspan citationid=\"CR57\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). Our study showed most of the murid rodents of Iran resulted from the westward expansion of an ancestry during the Miocene and most of the speciation events in this family in Iran occurred during the Late Miocene and Pliocene, when Iran experienced orogeny events. The active geology of Iran, resulting from collisions of the Arabian with the Iranian plate that has led to rapid isolation of areas and changing hydrological networks (Allen et al. \u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e2004\u003c/span\u003e; Hatzfeld et al. \u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e2010\u003c/span\u003e). The rising of mountains ranges in Iran accelerated during the Middle to Late Miocene (15\u0026ndash;5 Ma). This may have promoted allopatric speciation (Ahmadzadeh et al. \u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2013\u003c/span\u003e; Mee and Moore \u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e2014\u003c/span\u003e), which according to our analyses would have been the result of dispersal events. Also, High topographic complexity in the mountains probably causes high habitat diversity and thus a large number of local niche spaces. This is expected to foster adaptation to different niches (i.e., ecological speciation) as well as to create local refugia for species during climatic fluctuations, reducing extinction risks (Ashcroft et al. \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e2012\u003c/span\u003e; Manafzadeh et al. \u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e2017\u003c/span\u003e; Hashemzadeh Segherloo et al. \u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). Most of the speciation events inside murids occurred in west and north of Iran where the Alborz and Zagros mountains extend along. These mountains are two hotspots for Iran and act as endemism centers for different taxonomic groups (Noroozi et al. \u003cspan citationid=\"CR42\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). Besides, there is evidence from paleoclimatic and phylogeographic modeling that have highlighted Alborz and Zagros mountains acting as glacial refugia for species of insects, birds, reptiles and mammals during past climatic oscillations (Eskandarzadeh et al. \u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e2018\u003c/span\u003e).\u003c/p\u003e"},{"header":"Conclusion","content":"\u003cp\u003eHere, we investigate the phylogenetic relationships and the historical biogeography of Iranian Muridae, providing a time calibrated phylogeny for this family of rodents in this region. Our results based on mitochondrial cytochrome \u003cem\u003eb\u003c/em\u003e and a reduced number of species are mostly consistent with those based on several nuclear and mitochondrial markers (even up to 15 markers) and a much dense sampling of taxa. Our analyses evidence a time of origin of Muridae about 18.86 Ma. Iranian murids most likely originated in central and eastern Iran and mountain regions, particularly in Zagros and Alborz. These mountain ranges may have been center of speciation for this family of rodents in Iran by providing different ecological niches and local refugia during climate oscillations. Multiple colonization events may have taken place in Iran, which acted, repeatedly, as a corridor for faunal dispersal between Afro-Arabia and Eurasia and between Central Asia and the Mediterranean regions.\u003c/p\u003e"},{"header":"Declarations","content":"\u003cp\u003e\u003cstrong\u003eSupplementary Information\u0026nbsp;\u003c/strong\u003eThe online version contains supplementary material available at https://doi.org/XXX/XXXXXX.\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eAcknowledgments\u0026nbsp;\u003c/strong\u003eIn memory of Professor Jamshid Darvish, who founded animal systematics in Iran.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eAuthor contributions\u0026nbsp;\u003c/strong\u003eYAA has designed the study. Material preparation, data collection and analysis have been performed by YAA. The manuscript has been written by YAA and RLA, who\u0026nbsp;supervised this study. All authors have red and approved the manuscript.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eFunding\u0026nbsp;\u003c/strong\u003eThis research received support from the French National Research Agency (project RoMa ANR-22-CE02)\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eData availability\u0026nbsp;\u003c/strong\u003eAll genetic sequences used in this study are publicly available in the GenBank. Also, all data analyzed during this study are included in this published article and its supplementary information files.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eConflict of interest\u0026nbsp;\u003c/strong\u003eThe authors have no competing interests to declare that are relevant to the content of this article.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eEthics approval\u0026nbsp;\u003c/strong\u003eNot applicable.\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\n\u003cli\u003eAghov\u0026aacute; T, Kimura Y, Bryja J, Dobigny G, Granjon L, Kergoat GJ (2018) Fossils know it best: using a new set of fossil calibrations to improve the temporal phylogenetic framework of murid rodents (Rodentia: Muridae). 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Zoology in the Middle East 26:7\u0026ndash;10. https://doi.org/10.1080/09397140.2002.10637915\u003c/li\u003e\n\u003c/ol\u003e"},{"header":"Tables","content":"\u003cp\u003e\u003cstrong\u003eTable\u0026nbsp;\u003c/strong\u003e\u003cstrong\u003e1\u003c/strong\u003e Murid rodent species used in this study.\u003c/p\u003e\n\u003ctable border=\"0\" cellspacing=\"0\" cellpadding=\"0\" width=\"601\"\u003e\n \u003ctbody\u003e\n \u003ctr\u003e\n \u003ctd width=\"11.480865224625624%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cstrong\u003eFamily\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"13.643926788685524%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cstrong\u003eSubfamily\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"36.7720465890183%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cstrong\u003eGenus\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"38.10316139767055%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cstrong\u003eSpecies\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"11.480865224625624%\" rowspan=\"17\" valign=\"top\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003cp\u003eMuridae\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"13.643926788685524%\" rowspan=\"9\" valign=\"top\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003cp\u003eMurinae\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"36.7720465890183%\" rowspan=\"5\" valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003e\u0026nbsp;\u003c/em\u003e\u003c/p\u003e\n \u003cp\u003e\u003cem\u003e\u0026nbsp;\u003c/em\u003e\u003c/p\u003e\n \u003cp\u003e\u003cem\u003e\u0026nbsp;\u003c/em\u003e\u003c/p\u003e\n \u003cp\u003e\u003cem\u003eApodemus\u0026nbsp;\u003c/em\u003eKaup, 1829\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"38.10316139767055%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003eApodemus hyrcanicus\u003c/em\u003e Vorontsov, Boyeskorov \u0026amp; Mezhzherin, 1992\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"100%\" valign=\"top\"\u003e\n \u003cp\u003eA\u003cem\u003epodemus mystacinus\u0026nbsp;\u003c/em\u003eDanford \u0026amp; Alston, 1877\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"100%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003eApodemus ponticus\u0026nbsp;\u003c/em\u003eSviridenko, 1936\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"100%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003eApodemus uralensis\u003c/em\u003e Pallas, 1811\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"100%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003eApodemus witherbyi\u003c/em\u003e Thomas, 1902\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"49.111111111111114%\" rowspan=\"3\" valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003e\u0026nbsp;\u003c/em\u003e\u003c/p\u003e\n \u003cp\u003e\u003cem\u003eMus\u0026nbsp;\u003c/em\u003eClerck, 1757\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"50.888888888888886%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003eMus musculus\u003c/em\u003e Linnaeus, 1758\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"100%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003eMus domesticus\u0026nbsp;\u003c/em\u003eSchwarz \u0026amp; Schwarz, 1943\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"100%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003eMus macedonicus\u0026nbsp;\u003c/em\u003ePetrov \u0026amp; Ruzic, 1983\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"49.111111111111114%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003eRattus\u0026nbsp;\u003c/em\u003eFischer de Waldheim, 1803\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"50.888888888888886%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003eRattus rattus\u0026nbsp;\u003c/em\u003eLinnaeus, 1758\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"15.413533834586467%\" rowspan=\"8\" valign=\"top\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003cp\u003eGerbillinae\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"41.54135338345865%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003eGerbillus\u0026nbsp;\u003c/em\u003eDesmarest, 1804\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"43.045112781954884%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003eGerbillus nanus\u003c/em\u003e Blanford, 1875\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"49.111111111111114%\" rowspan=\"6\" valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003e\u0026nbsp;\u003c/em\u003e\u003c/p\u003e\n \u003cp\u003e\u003cem\u003e\u0026nbsp;\u003c/em\u003e\u003c/p\u003e\n \u003cp\u003e\u003cem\u003eMeriones\u0026nbsp;\u003c/em\u003eIlliger, 1811\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"50.888888888888886%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003eMeriones crassus\u0026nbsp;\u003c/em\u003eSundevall, 1842\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"100%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003eMeriones libycus\u003c/em\u003e Lichtenstein, 1823\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"100%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003eMeriones meridianus\u003c/em\u003e Pallas, 1773\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"100%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003eMeriones persicus\u003c/em\u003e Blanford, 1875\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"100%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003eMeriones tristrami\u003c/em\u003e Thomas, 1892\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"100%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003eMeriones vinogradovi\u003c/em\u003e Heptner, 1931\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"49.111111111111114%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003eRhombomys\u0026nbsp;\u003c/em\u003eWagner, 1841\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"50.888888888888886%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003eRhombomys opimus\u0026nbsp;\u003c/em\u003eLichtenstein, 1823\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003c/tbody\u003e\n\u003c/table\u003e\n\u003cp\u003e\u003cstrong\u003eTable\u0026nbsp;\u003c/strong\u003e\u003cstrong\u003e2\u003c/strong\u003e Fossil calibrations used in this work for dating analyses. All ages are in million years before present.\u003c/p\u003e\n\u003ctable border=\"0\" cellspacing=\"0\" cellpadding=\"0\" width=\"576\"\u003e\n \u003ctbody\u003e\n \u003ctr\u003e\n \u003ctd width=\"15.625%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cstrong\u003eTaxon\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"13.541666666666666%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cstrong\u003eMin Age\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"13.541666666666666%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cstrong\u003eMax Age\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"15.625%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cstrong\u003eLog-StDev\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.590277777777779%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cstrong\u003eOffset\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"31.07638888888889%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cstrong\u003eReference\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"15.625%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003eApodemus\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"13.541666666666666%\" valign=\"top\"\u003e\n \u003cp\u003e5.3\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"13.541666666666666%\" valign=\"top\"\u003e\n \u003cp\u003e7.2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"15.625%\" valign=\"top\"\u003e\n \u003cp\u003e0.515\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.590277777777779%\" valign=\"top\"\u003e\n \u003cp\u003e4.871\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"31.07638888888889%\" valign=\"top\"\u003e\n \u003cp\u003eSteppan and Schenk 2017\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"15.625%\" valign=\"top\"\u003e\n \u003cp\u003eMurinae\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"13.541666666666666%\" valign=\"top\"\u003e\n \u003cp\u003e12.1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"13.541666666666666%\" valign=\"top\"\u003e\n \u003cp\u003e14.05\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"15.625%\" valign=\"top\"\u003e\n \u003cp\u003e0.885\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.590277777777779%\" valign=\"top\"\u003e\n \u003cp\u003e9.767\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"31.07638888888889%\" valign=\"top\"\u003e\n \u003cp\u003eSteppan and Schenk 2017\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"15.625%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003eMus\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"13.541666666666666%\" valign=\"top\"\u003e\n \u003cp\u003e5.3\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"13.541666666666666%\" valign=\"top\"\u003e\n \u003cp\u003e7.2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"15.625%\" valign=\"top\"\u003e\n \u003cp\u003e0.515\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.590277777777779%\" valign=\"top\"\u003e\n \u003cp\u003e4.871\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"31.07638888888889%\" valign=\"top\"\u003e\n \u003cp\u003eSteppan and Schenk 2017\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"15.625%\" valign=\"top\"\u003e\n \u003cp\u003eGerbil\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"13.541666666666666%\" valign=\"top\"\u003e\n \u003cp\u003e16\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"13.541666666666666%\" valign=\"top\"\u003e\n \u003cp\u003e23.7\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"15.625%\" valign=\"top\"\u003e\n \u003cp\u003e1.25\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.590277777777779%\" valign=\"top\"\u003e\n \u003cp\u003e15.868\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"31.07638888888889%\" valign=\"top\"\u003e\n \u003cp\u003eSchenk et al. 2013\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003c/tbody\u003e\n\u003c/table\u003e\n\u003cp\u003e\u003cstrong\u003eTable 3\u003c/strong\u003e Murid rodent species range in terrestrial ecoregions of Iran.\u003c/p\u003e\n\u003ctable border=\"1\" cellspacing=\"0\" cellpadding=\"0\"\u003e\n \u003ctbody\u003e\n \u003ctr\u003e\n \u003ctd width=\"27.333333333333332%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cstrong\u003eSpecies\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"12.666666666666666%\" valign=\"top\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"11%\" valign=\"top\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"16.166666666666668%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cstrong\u003eEcoregions\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"11%\" valign=\"top\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"11.166666666666666%\" valign=\"top\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.666666666666666%\" valign=\"top\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"27.333333333333332%\" valign=\"top\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"12.666666666666666%\" valign=\"top\"\u003e\n \u003cp\u003eA\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"11%\" valign=\"top\"\u003e\n \u003cp\u003eB\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"16.166666666666668%\" valign=\"top\"\u003e\n \u003cp\u003eC\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"11%\" valign=\"top\"\u003e\n \u003cp\u003eD\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"11.166666666666666%\" valign=\"top\"\u003e\n \u003cp\u003eE\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.666666666666666%\" valign=\"top\"\u003e\n \u003cp\u003eF\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"27.333333333333332%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003eMus domesticus\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"12.666666666666666%\" valign=\"top\"\u003e\n \u003cp\u003e*\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"11%\" valign=\"top\"\u003e\n \u003cp\u003e*\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"16.166666666666668%\" valign=\"top\"\u003e\n \u003cp\u003e*\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"11%\" valign=\"top\"\u003e\n \u003cp\u003e*\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"11.166666666666666%\" valign=\"top\"\u003e\n \u003cp\u003e*\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.666666666666666%\" valign=\"top\"\u003e\n \u003cp\u003e*\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"27.333333333333332%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003eMus musculus\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"12.666666666666666%\" valign=\"top\"\u003e\n \u003cp\u003e*\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"11%\" valign=\"top\"\u003e\n \u003cp\u003e*\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"16.166666666666668%\" valign=\"top\"\u003e\n \u003cp\u003e*\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"11%\" valign=\"top\"\u003e\n \u003cp\u003e*\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"11.166666666666666%\" valign=\"top\"\u003e\n \u003cp\u003e*\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.666666666666666%\" valign=\"top\"\u003e\n \u003cp\u003e*\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"27.333333333333332%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003eMus macedonicus\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"12.666666666666666%\" valign=\"top\"\u003e\n \u003cp\u003e*\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"11%\" valign=\"top\"\u003e\n \u003cp\u003e*\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"16.166666666666668%\" valign=\"top\"\u003e\n \u003cp\u003e*\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"11%\" valign=\"top\"\u003e\n \u003cp\u003e*\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"11.166666666666666%\" valign=\"top\"\u003e\n \u003cp\u003e*\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.666666666666666%\" valign=\"top\"\u003e\n \u003cp\u003e*\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"27.333333333333332%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003eRattus rattus\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"12.666666666666666%\" valign=\"top\"\u003e\n \u003cp\u003e*\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"11%\" valign=\"top\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"16.166666666666668%\" valign=\"top\"\u003e\n \u003cp\u003e*\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"11%\" valign=\"top\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"11.166666666666666%\" valign=\"top\"\u003e\n \u003cp\u003e*\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.666666666666666%\" valign=\"top\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"27.333333333333332%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003eApodemus mystacinus\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"12.666666666666666%\" valign=\"top\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"11%\" valign=\"top\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"16.166666666666668%\" valign=\"top\"\u003e\n \u003cp\u003e*\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"11%\" valign=\"top\"\u003e\n \u003cp\u003e*\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"11.166666666666666%\" valign=\"top\"\u003e\n \u003cp\u003e*\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.666666666666666%\" valign=\"top\"\u003e\n \u003cp\u003e*\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"27.333333333333332%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003eApodemus ponticus\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"12.666666666666666%\" valign=\"top\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"11%\" valign=\"top\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"16.166666666666668%\" valign=\"top\"\u003e\n \u003cp\u003e*\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"11%\" valign=\"top\"\u003e\n \u003cp\u003e*\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"11.166666666666666%\" valign=\"top\"\u003e\n \u003cp\u003e*\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.666666666666666%\" valign=\"top\"\u003e\n \u003cp\u003e*\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"27.333333333333332%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003eApodemus witherbyi\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"12.666666666666666%\" valign=\"top\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"11%\" valign=\"top\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"16.166666666666668%\" valign=\"top\"\u003e\n \u003cp\u003e*\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"11%\" valign=\"top\"\u003e\n \u003cp\u003e*\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"11.166666666666666%\" valign=\"top\"\u003e\n \u003cp\u003e*\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.666666666666666%\" valign=\"top\"\u003e\n \u003cp\u003e*\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"27.333333333333332%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003eApodemus hyrcanicus\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"12.666666666666666%\" valign=\"top\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"11%\" valign=\"top\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"16.166666666666668%\" valign=\"top\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"11%\" valign=\"top\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"11.166666666666666%\" valign=\"top\"\u003e\n \u003cp\u003e*\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.666666666666666%\" valign=\"top\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"27.333333333333332%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003eApodemus uralensis\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"12.666666666666666%\" valign=\"top\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"11%\" valign=\"top\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"16.166666666666668%\" valign=\"top\"\u003e\n \u003cp\u003e*\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"11%\" valign=\"top\"\u003e\n \u003cp\u003e*\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"11.166666666666666%\" valign=\"top\"\u003e\n \u003cp\u003e*\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.666666666666666%\" valign=\"top\"\u003e\n \u003cp\u003e*\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"27.333333333333332%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003eGerbillus nanus\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"12.666666666666666%\" valign=\"top\"\u003e\n \u003cp\u003e*\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"11%\" valign=\"top\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"16.166666666666668%\" valign=\"top\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"11%\" valign=\"top\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"11.166666666666666%\" valign=\"top\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.666666666666666%\" valign=\"top\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"27.333333333333332%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003eRhombomys opimus\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"12.666666666666666%\" valign=\"top\"\u003e\n \u003cp\u003e*\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"11%\" valign=\"top\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"16.166666666666668%\" valign=\"top\"\u003e\n \u003cp\u003e*\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"11%\" valign=\"top\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"11.166666666666666%\" valign=\"top\"\u003e\n \u003cp\u003e*\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.666666666666666%\" valign=\"top\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"27.333333333333332%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003eMeriones persicus\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"12.666666666666666%\" valign=\"top\"\u003e\n \u003cp\u003e*\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"11%\" valign=\"top\"\u003e\n \u003cp\u003e*\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"16.166666666666668%\" valign=\"top\"\u003e\n \u003cp\u003e*\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"11%\" valign=\"top\"\u003e\n \u003cp\u003e*\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"11.166666666666666%\" valign=\"top\"\u003e\n \u003cp\u003e*\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.666666666666666%\" valign=\"top\"\u003e\n \u003cp\u003e*\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"27.333333333333332%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003eMeriones vinogradovi\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"12.666666666666666%\" valign=\"top\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"11%\" valign=\"top\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"16.166666666666668%\" valign=\"top\"\u003e\n \u003cp\u003e*\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"11%\" valign=\"top\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"11.166666666666666%\" valign=\"top\"\u003e\n \u003cp\u003e*\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.666666666666666%\" valign=\"top\"\u003e\n \u003cp\u003e*\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"27.333333333333332%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003eMeriones tristrami\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"12.666666666666666%\" valign=\"top\"\u003e\n \u003cp\u003e*\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"11%\" valign=\"top\"\u003e\n \u003cp\u003e*\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"16.166666666666668%\" valign=\"top\"\u003e\n \u003cp\u003e*\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"11%\" valign=\"top\"\u003e\n \u003cp\u003e*\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"11.166666666666666%\" valign=\"top\"\u003e\n \u003cp\u003e*\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.666666666666666%\" valign=\"top\"\u003e\n \u003cp\u003e*\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"27.333333333333332%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003eMeriones libycus\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"12.666666666666666%\" valign=\"top\"\u003e\n \u003cp\u003e*\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"11%\" valign=\"top\"\u003e\n \u003cp\u003e*\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"16.166666666666668%\" valign=\"top\"\u003e\n \u003cp\u003e*\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"11%\" valign=\"top\"\u003e\n \u003cp\u003e*\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"11.166666666666666%\" valign=\"top\"\u003e\n \u003cp\u003e*\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.666666666666666%\" valign=\"top\"\u003e\n \u003cp\u003e*\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"27.333333333333332%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003eMeriones meridianus\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"12.666666666666666%\" valign=\"top\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"11%\" valign=\"top\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"16.166666666666668%\" valign=\"top\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"11%\" valign=\"top\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"11.166666666666666%\" valign=\"top\"\u003e\n \u003cp\u003e*\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.666666666666666%\" valign=\"top\"\u003e\n \u003cp\u003e*\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"27.333333333333332%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003eMeriones crasus\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"12.666666666666666%\" valign=\"top\"\u003e\n \u003cp\u003e*\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"11%\" valign=\"top\"\u003e\n \u003cp\u003e*\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"16.166666666666668%\" valign=\"top\"\u003e\n \u003cp\u003e*\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"11%\" valign=\"top\"\u003e\n \u003cp\u003e*\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"11.166666666666666%\" valign=\"top\"\u003e\n \u003cp\u003e*\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"10.666666666666666%\" valign=\"top\"\u003e\n \u003cp\u003e*\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003c/tbody\u003e\n\u003c/table\u003e\n\u003cp\u003e\u003cbr\u003e\u003c/p\u003e\n\u003cp\u003eTable 4 is available in the Supplementary Files section.\u003c/p\u003e"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":false,"hideJournal":false,"highlight":"","institution":"","isAcceptedByJournal":true,"isAuthorSuppliedPdf":false,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":true,"isPdf":false,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","identity":"mammalian-biology","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"mamb","sideBox":"Learn more about [Mammalian Biology](https://link.springer.com/journal/42991)","snPcode":"42991","submissionUrl":"https://www.editorialmanager.com/mamb/default2.aspx","title":"Mammalian Biology","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false},"keywords":"Murinae, Gerbillinae, Bayesian molecular clock, Divergence time, Biogeography, Middle East","lastPublishedDoi":"10.21203/rs.3.rs-3186974/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-3186974/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003eThe family Muridae represents the largest, most diverse and successful of all groups of mammals. Here we infer the phylogenetic relationships and historical biogeography for the Iranian murid rodents, which consist in 17 species distributed in two subfamilies and six genera. Molecular dating analyses using a relaxed Bayesian molecular clock support the monophyly of Murinae and Gerbillinae and allow to set up a divergence date between them around 18.86 Ma (Million years ago). According to our results, murines may have originated approximately 13.49 Ma and the diversification of most of their evolutionary lineages may have taken place between 10\u0026ndash;4 Ma, which is consistent with the fossil record. Our results provide strong support for the tribes Apodemyini and Rattini (Murinae) but the monophyly of the genus \u003cem\u003eMeriones\u003c/em\u003e belonging to the Gerbillinae is questioned. Historical biogeographic analysis supports a Paleotropical origin for the Iranian murids, likely found in central and eastern Iran (Desert and Xeric Shrubland ecoregion). From there they dispersed to colonize the Afrotropical, Indomalayan and Palearctic realms. All in all, Iran seems to have acted as a corridor for faunal exchanges between the Afrotropic and Saharo-Arabian realms and the Indomalayan realm as well as between Central Asia and the Mediterranean regions.\u003c/p\u003e","manuscriptTitle":"Molecular phylogeny and historical biogeography of Iranian Murid rodents inferred from mitochondrial cytochrome b gene","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2023-08-07 13:46:33","doi":"10.21203/rs.3.rs-3186974/v1","editorialEvents":[{"type":"communityComments","content":0},{"type":"reviewerAgreed","content":"","date":"2023-08-02T07:06:46+00:00","index":0,"fulltext":""},{"type":"reviewersInvited","content":"","date":"2023-08-01T12:39:49+00:00","index":"","fulltext":""},{"type":"editorAssigned","content":"","date":"2023-07-22T02:07:28+00:00","index":"","fulltext":""},{"type":"submitted","content":"Mammalian Biology","date":"2023-07-20T00:59:51+00:00","index":"","fulltext":""}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"mammalian-biology","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"mamb","sideBox":"Learn more about [Mammalian Biology](https://link.springer.com/journal/42991)","snPcode":"42991","submissionUrl":"https://www.editorialmanager.com/mamb/default2.aspx","title":"Mammalian Biology","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false}}],"origin":"","ownerIdentity":"d2daece5-717b-422a-865c-17cfb8a0feb1","owner":[],"postedDate":"August 7th, 2023","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"published-in-journal","subjectAreas":[],"tags":[],"updatedAt":"2024-01-15T15:07:53+00:00","versionOfRecord":{"articleIdentity":"rs-3186974","link":"https://doi.org/10.1007/s42991-023-00390-3","journal":{"identity":"mammalian-biology","isVorOnly":false,"title":"Mammalian Biology"},"publishedOn":"2024-01-12 15:01:21","publishedOnDateReadable":"January 12th, 2024"},"versionCreatedAt":"2023-08-07 13:46:33","video":"","vorDoi":"10.1007/s42991-023-00390-3","vorDoiUrl":"https://doi.org/10.1007/s42991-023-00390-3","workflowStages":[]},"version":"v1","identity":"rs-3186974","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-3186974","identity":"rs-3186974","version":["v1"]},"buildId":"cBFmMYwuxLRRLfASyISRj","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}