{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2025,12,8]],"date-time":"2025-12-08T17:30:07Z","timestamp":1765215007333,"version":"3.46.0"},"reference-count":55,"publisher":"Oxford University Press (OUP)","issue":"12","license":[{"start":{"date-parts":[[2025,11,26]],"date-time":"2025-11-26T00:00:00Z","timestamp":1764115200000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/"}],"funder":[{"name":"Youth Fund of the National Natural Science Foundation of China","award":["32200345"],"award-info":[{"award-number":["32200345"]}]},{"DOI":"10.13039\/501100002858","name":"China Postdoctoral Science Foundation","doi-asserted-by":"publisher","award":["2022M710878"],"award-info":[{"award-number":["2022M710878"]}],"id":[{"id":"10.13039\/501100002858","id-type":"DOI","asserted-by":"publisher"}]},{"name":"Guangzhou Basic and Applied Basic Research Foundation","award":["2023A04J0770"],"award-info":[{"award-number":["2023A04J0770"]}]},{"DOI":"10.13039\/501100012166","name":"National Key Research and Development Program of China","doi-asserted-by":"publisher","award":["2022YFF1301601"],"award-info":[{"award-number":["2022YFF1301601"]}],"id":[{"id":"10.13039\/501100012166","id-type":"DOI","asserted-by":"publisher"}]},{"name":"PI Project of Southern Marine Science and Engineering Guangdong Laboratory","award":["GML2021GD0805"],"award-info":[{"award-number":["GML2021GD0805"]}]},{"DOI":"10.13039\/501100001809","name":"National Natural Science Foundation of China","doi-asserted-by":"publisher","award":["32270453","31772448","32270442"],"award-info":[{"award-number":["32270453","31772448","32270442"]}],"id":[{"id":"10.13039\/501100001809","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":[],"published-print":{"date-parts":[[2025,11,28]]},"abstract":"<jats:title>Abstract<\/jats:title>\n                  <jats:p>The testicular descent leading to the exteriorization of male gonads is a complex sexual dimorphic process observed in numerous mammals, whereas some species, such as elephants and cetaceans, retain the lifelong ascrotal testes. Despite the implication of Wnt signaling in testicular development and descent, the evolutionary mechanisms underlying ascrotal testes have not been adequately addressed. Here, we examined selection signatures and unique amino acid substitutions in genes of the Wnt signaling pathway. We identified an ascrotal mammal-specific substitution (S406G) in zinc and ring finger 3 (ZNRF3) located within the disheveled (DVL)-interaction region. Functional assays showed that this substitution enhances the affinity of ascrotal ZNRF3 for DVLs and suppresses Wnt3a-induced Wnt signaling. Comparative transcriptomics of the gubernaculum between male fetal rats from wild-type and spontaneous cryptorchid (orl) strains revealed upregulation of Ctnnb1 and Gsk3b in orl rats, along with ascrotal-specific evolutionary changes in regulatory elements. Collectively, these findings suggest that Wnt signaling may be dampened in ascrotal mammals. This study provides insights into the pathogenesis of cryptorchidism in humans and domestic animals.<\/jats:p>","DOI":"10.1093\/molbev\/msaf310","type":"journal-article","created":{"date-parts":[[2025,11,25]],"date-time":"2025-11-25T12:50:47Z","timestamp":1764075047000},"source":"Crossref","is-referenced-by-count":0,"title":["Evolutionary Dampening of Wnt Signaling May Contribute to Naturally Ascrotal Testes in Mammals"],"prefix":"10.1093","volume":"42","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-2301-5171","authenticated-orcid":false,"given":"Simin","family":"Chai","sequence":"first","affiliation":[{"name":"Southern Marine Science and Engineering Guangdong Laboratory (Guangzhou) , Guangzhou, Guangdong ,","place":["China"]}],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0009-0003-2601-8620","authenticated-orcid":false,"given":"Cuijuan","family":"Zhong","sequence":"additional","affiliation":[{"name":"Jiangsu Key Laboratory for the Biodiversity Conservation and Sustainable Utilization in the Middle and Lower Reaches of the Yangtze River Basin, College of Life Sciences, Nanjing Normal University , Nanjing, Jiangsu ,","place":["China"]}],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0009-0001-1321-2307","authenticated-orcid":false,"given":"Zhenhua","family":"Zhang","sequence":"additional","affiliation":[{"name":"Jiangsu Key Laboratory for the Biodiversity Conservation and Sustainable Utilization in the Middle and Lower Reaches of the Yangtze River Basin, College of Life Sciences, Nanjing Normal University , Nanjing, Jiangsu ,","place":["China"]}],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0009-0007-6682-5864","authenticated-orcid":false,"given":"Daiqing","family":"Yin","sequence":"additional","affiliation":[{"name":"Southern Marine Science and Engineering Guangdong Laboratory (Guangzhou) , Guangzhou, Guangdong ,","place":["China"]}],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-0283-0986","authenticated-orcid":false,"given":"Shixia","family":"Xu","sequence":"additional","affiliation":[{"name":"Jiangsu Key Laboratory for the Biodiversity Conservation and Sustainable Utilization in the Middle and Lower Reaches of the Yangtze River Basin, College of Life Sciences, Nanjing Normal University , Nanjing, Jiangsu ,","place":["China"]}],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-8429-5963","authenticated-orcid":false,"given":"Wenhua","family":"Ren","sequence":"additional","affiliation":[{"name":"Jiangsu Key Laboratory for the Biodiversity Conservation and Sustainable Utilization in the Middle and Lower Reaches of the Yangtze River Basin, College of Life Sciences, Nanjing Normal University , Nanjing, Jiangsu ,","place":["China"]}],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-6285-6937","authenticated-orcid":false,"given":"Guang","family":"Yang","sequence":"additional","affiliation":[{"name":"Southern Marine Science and Engineering Guangdong Laboratory (Guangzhou) , Guangzhou, Guangdong ,","place":["China"]},{"name":"Jiangsu Key Laboratory for the Biodiversity Conservation and Sustainable Utilization in the Middle and Lower Reaches of the Yangtze River Basin, College of Life Sciences, Nanjing Normal University , Nanjing, Jiangsu ,","place":["China"]}],"role":[{"role":"author","vocabulary":"crossref"}]}],"member":"286","published-online":{"date-parts":[[2025,11,26]]},"reference":[{"key":"2025120812271856200_msaf310-B1","doi-asserted-by":"publisher","first-page":"btad623","DOI":"10.1093\/bioinformatics\/btad623","article-title":"CAAStools: a toolbox to identify and test convergent amino acid substitutions","volume":"39","author":"Barteri","year":"2023","journal-title":"Bioinformatics"},{"key":"2025120812271856200_msaf310-B2","doi-asserted-by":"publisher","first-page":"352","DOI":"10.2164\/jandrol.107.003970","article-title":"Altered expression of muscle-and cytoskeleton-related genes in a rat strain with inherited cryptorchidism","volume":"29","author":"Barthold","year":"2008","journal-title":"J Androl"},{"key":"2025120812271856200_msaf310-B3","doi-asserted-by":"publisher","first-page":"141","DOI":"10.1095\/biolreprod.113.112953","article-title":"Transcriptome analysis of the dihydrotestosterone-exposed fetal rat gubernaculum identifies common androgen and insulin-like 3 targets","volume":"89","author":"Barthold","year":"2013","journal-title":"Biol Reprod"},{"key":"2025120812271856200_msaf310-B4","doi-asserted-by":"publisher","first-page":"351","DOI":"10.1007\/s00441-017-2698-5","article-title":"Defective Wnt3 expression by testicular Sertoli cells compromise male fertility","volume":"371","author":"Basu","year":"2018","journal-title":"Cell Tissue Res"},{"key":"2025120812271856200_msaf310-B5","doi-asserted-by":"publisher","first-page":"W193","DOI":"10.1093\/nar\/gkaa382","article-title":"EpiRegio: analysis and retrieval of regulatory elements linked to genes","volume":"48","author":"Baumgarten","year":"2020","journal-title":"Nucleic Acids Res"},{"key":"2025120812271856200_msaf310-B6","doi-asserted-by":"publisher","first-page":"97","DOI":"10.1111\/j.1365-2605.2010.01074.x","article-title":"Human testicular insulin-like factor 3: in relation to development, reproductive hormones and andrological disorders","volume":"34","author":"Bay","year":"2011","journal-title":"Int J Androl"},{"key":"2025120812271856200_msaf310-B7","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1186\/s12862-021-01753-5","article-title":"Rapid evolution and molecular convergence in cryptorchidism-related genes associated with inherently undescended testes in mammals","volume":"21","author":"Chai","year":"2021","journal-title":"BMC Ecol Evol"},{"author":"Chen","key":"2025120812271856200_msaf310-B8","doi-asserted-by":"publisher","DOI":"10.1016\/j.molp.2020.06.009"},{"key":"2025120812271856200_msaf310-B9","doi-asserted-by":"publisher","first-page":"D988","DOI":"10.1093\/nar\/gkab1049","article-title":"Ensembl 2022","volume":"50","author":"Cunningham","year":"2022","journal-title":"Nucleic Acids Res."},{"key":"2025120812271856200_msaf310-B10","doi-asserted-by":"publisher","first-page":"155","DOI":"10.1016\/j.gene.2015.04.066","article-title":"Wnt signaling in testis development: unnecessary or essential?","volume":"565","author":"Dong","year":"2015","journal-title":"Gene"},{"key":"2025120812271856200_msaf310-B11","doi-asserted-by":"publisher","first-page":"643","DOI":"10.1007\/s00383-010-2604-7","article-title":"The testicular descent in the rat: a scanning electron microscopic study","volume":"26","author":"Fiegel","year":"2010","journal-title":"Pediatric Surg Int"},{"key":"2025120812271856200_msaf310-B12","doi-asserted-by":"publisher","first-page":"725","DOI":"10.1093\/oxfordjournals.molbev.a040153","article-title":"A codon-based model of nucleotide substitution for protein-coding DNA sequences","volume":"11","author":"Goldman","year":"1994","journal-title":"Mol Biol Evol"},{"key":"2025120812271856200_msaf310-B13","doi-asserted-by":"publisher","first-page":"221","DOI":"10.1093\/molbev\/msp259","article-title":"SeaView version 4: a multiplatform graphical user interface for sequence alignment and phylogenetic tree building","volume":"27","author":"Gouy","year":"2010","journal-title":"Mol Biol Evol"},{"volume-title":"Cryptorchidism: management and implications","year":"2012","author":"Hadziselimovic","key":"2025120812271856200_msaf310-B14"},{"key":"2025120812271856200_msaf310-B15","doi-asserted-by":"publisher","first-page":"195","DOI":"10.1038\/nature11019","article-title":"ZNRF3 promotes Wnt receptor turnover in an R-spondin-sensitive manner","volume":"485","author":"Hao","year":"2012","journal-title":"Nature"},{"key":"2025120812271856200_msaf310-B16","doi-asserted-by":"publisher","first-page":"1573","DOI":"10.1016\/j.jpedsurg.2012.08.038","article-title":"Wnt signalling in testicular descent: a candidate mechanism for cryptorchidism in Robinow syndrome","volume":"48","author":"Harisis","year":"2013","journal-title":"J Pediatric Surg"},{"key":"2025120812271856200_msaf310-B17","doi-asserted-by":"publisher","first-page":"1086","DOI":"10.1093\/molbev\/msz049","article-title":"Bayesian detection of convergent rate changes of conserved noncoding elements on phylogenetic trees","volume":"36","author":"Hu","year":"2019","journal-title":"Mol Biol Evol"},{"author":"Hutson","key":"2025120812271856200_msaf310-B18"},{"key":"2025120812271856200_msaf310-B19","doi-asserted-by":"publisher","first-page":"259","DOI":"10.1210\/edrv.18.2.0298","article-title":"Anatomical and functional aspects of testicular descent and cryptorchidism","volume":"18","author":"Hutson","year":"1997","journal-title":"Endocr Rev"},{"key":"2025120812271856200_msaf310-B20","doi-asserted-by":"publisher","first-page":"115","DOI":"10.1177\/1756287209105266","article-title":"The role of the gubernaculum in the descent and undescent of the testis","volume":"1","author":"Hutson","year":"2009","journal-title":"Therapeutic Adv Urol"},{"key":"2025120812271856200_msaf310-B21","doi-asserted-by":"crossref","DOI":"10.1007\/978-3-319-25910-9","volume-title":"Descent of the testis","author":"Hutson","year":"2016"},{"key":"2025120812271856200_msaf310-B22","doi-asserted-by":"publisher","first-page":"522","DOI":"10.1016\/j.molcel.2015.03.015","article-title":"Dishevelled promotes Wnt receptor degradation through recruitment of ZNRF3\/RNF43 E3 ubiquitin ligases","volume":"58","author":"Jiang","year":"2015","journal-title":"Mol Cell"},{"key":"2025120812271856200_msaf310-B23","doi-asserted-by":"publisher","first-page":"774","DOI":"10.1095\/biolreprod.110.085175","article-title":"Insulin-like 3 exposure of the fetal rat gubernaculum modulates expression of genes involved in neural pathways1","volume":"83","author":"Johnson","year":"2010","journal-title":"Biol Reprod"},{"key":"2025120812271856200_msaf310-B24","doi-asserted-by":"publisher","first-page":"10866","DOI":"10.1073\/pnas.1834480100","article-title":"Wnt4 overexpression disrupts normal testicular vasculature and inhibits testosterone synthesis by repressing steroidogenic factor 1\/\u03b2-catenin synergy","volume":"100","author":"Jordan","year":"2003","journal-title":"Proc Natl Acad Sci U S A."},{"key":"2025120812271856200_msaf310-B25","doi-asserted-by":"publisher","first-page":"170","DOI":"10.1210\/me.2010-0330","article-title":"Suppression of insulin-like3 receptor reveals the role of \u03b2-catenin and Notch signaling in gubernaculum development","volume":"25","author":"Kaftanovskaya","year":"2011","journal-title":"Mol Endocrinol"},{"key":"2025120812271856200_msaf310-B26","doi-asserted-by":"publisher","first-page":"39","DOI":"10.1002\/path.4161","article-title":"Left-sided cryptorchidism in mice with Wilms\u2019 tumour 1 gene deletion in gubernaculum testis","volume":"230","author":"Kaftanovskaya","year":"2013","journal-title":"J Pathol"},{"key":"2025120812271856200_msaf310-B27","doi-asserted-by":"publisher","first-page":"D493","DOI":"10.1093\/nar\/gkh103","article-title":"The UCSC Table Browser data retrieval tool","volume":"32","author":"Karolchik","year":"2004","journal-title":"Nucleic Acids Res"},{"key":"2025120812271856200_msaf310-B28","doi-asserted-by":"publisher","first-page":"772","DOI":"10.1093\/molbev\/mst010","article-title":"MAFFT multiple sequence alignment software version 7: improvements in performance and usability","volume":"30","author":"Katoh","year":"2013","journal-title":"Mol Biol Evol"},{"key":"2025120812271856200_msaf310-B29","doi-asserted-by":"publisher","first-page":"4042","DOI":"10.1158\/1078-0432.CCR-06-2316","article-title":"WNT signaling pathway and stem cell signaling network","volume":"13","author":"Katoh","year":"2007","journal-title":"Clin Cancer Res"},{"key":"2025120812271856200_msaf310-B30","doi-asserted-by":"publisher","first-page":"27","DOI":"10.1007\/s12038-010-0005-7","article-title":"The evolutionary history of testicular externalization and the origin of the scrotum","volume":"35","author":"Kleisner","year":"2010","journal-title":"J Biosci"},{"key":"2025120812271856200_msaf310-B31","doi-asserted-by":"publisher","first-page":"msac174","DOI":"10.1093\/molbev\/msac174","article-title":"TimeTree 5: an expanded resource for species divergence times","volume":"39","author":"Kumar","year":"2022","journal-title":"Mol Biol Evol"},{"key":"2025120812271856200_msaf310-B32","doi-asserted-by":"publisher","first-page":"W774","DOI":"10.1093\/nar\/gkac238","article-title":"Washu epigenome browser update 2022","volume":"50","author":"Li","year":"2022","journal-title":"Nucleic Acids Res"},{"key":"2025120812271856200_msaf310-B33","doi-asserted-by":"publisher","first-page":"781","DOI":"10.1146\/annurev.cellbio.20.010403.113126","article-title":"The Wnt signaling pathway in development and disease","volume":"20","author":"Logan","year":"2004","journal-title":"Annu Rev Cell Dev Biol"},{"key":"2025120812271856200_msaf310-B34","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1186\/s13059-014-0550-8","article-title":"Moderated estimation of fold change and dispersion for RNA-seq data with DESeq2","volume":"15","author":"Love","year":"2014","journal-title":"Genome Biol"},{"key":"2025120812271856200_msaf310-B35","doi-asserted-by":"publisher","first-page":"801","DOI":"10.1111\/jeb.12373","article-title":"Cool sperm: why some placental mammals have a scrotum","volume":"27","author":"Lovegrove","year":"2014","journal-title":"J Evol Biol"},{"key":"2025120812271856200_msaf310-B36","doi-asserted-by":"publisher","first-page":"1365","DOI":"10.1093\/molbev\/msv035","article-title":"Gene-wide identification of episodic selection","volume":"32","author":"Murrell","year":"2015","journal-title":"Mol Biol Evol"},{"volume-title":"Biology of Chinese tree shrews (Tupaia belangeri chinensis)","year":"1991","author":"Peng","key":"2025120812271856200_msaf310-B37"},{"key":"2025120812271856200_msaf310-B38","first-page":"365","article-title":"Observations on the gubernaculum during descent of the testis","volume":"16","author":"Radhakrishnan","year":"1979","journal-title":"Invest Urol"},{"key":"2025120812271856200_msaf310-B39","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1186\/s12301-023-00342-w","article-title":"The curious case of testicular descent: factors controlling testicular descent with a note on cryptorchidism","volume":"29","author":"Reny","year":"2023","journal-title":"African J Urol"},{"key":"2025120812271856200_msaf310-B40","doi-asserted-by":"publisher","first-page":"861","DOI":"10.1093\/molbev\/msz015","article-title":"Orthomam v10: scaling-up orthologous coding sequence and exon alignments with more than one hundred mammalian genomes","volume":"36","author":"Scornavacca","year":"2019","journal-title":"Mol Biol Evol"},{"volume-title":"Observationes nonnullae de testiculorum ex abdomine in scrotum descensu et partium genitalium anomaliis","year":"1817","author":"Seiler","key":"2025120812271856200_msaf310-B41"},{"key":"2025120812271856200_msaf310-B42","doi-asserted-by":"publisher","first-page":"dev201093","DOI":"10.1242\/dev.201093","article-title":"Loss of WNT4 in the gubernaculum causes unilateral cryptorchidism and fertility defects","volume":"149","author":"Seth","year":"2022","journal-title":"Development"},{"key":"2025120812271856200_msaf310-B43","doi-asserted-by":"publisher","first-page":"1034","DOI":"10.1101\/gr.3715005","article-title":"Evolutionarily conserved elements in vertebrate, insect, worm, and yeast genomes","volume":"15","author":"Siepel","year":"2005","journal-title":"Genome Res"},{"key":"2025120812271856200_msaf310-B44","doi-asserted-by":"publisher","first-page":"110","DOI":"10.24272\/j.issn.2095-8137.2017.017","article-title":"FasParser: a package for manipulating sequence data","volume":"38","author":"Sun","year":"2017","journal-title":"Zool Res"},{"key":"2025120812271856200_msaf310-B45","doi-asserted-by":"publisher","first-page":"564","DOI":"10.1080\/10635150701472164","article-title":"Improvement of phylogenies after removing divergent and ambiguously aligned blocks from protein sequence alignments","volume":"56","author":"Talavera","year":"2007","journal-title":"Syst Biol"},{"key":"2025120812271856200_msaf310-B46","doi-asserted-by":"publisher","first-page":"12","DOI":"10.1186\/1750-2187-6-12","article-title":"Canonical Wnt signaling is involved in switching from cell proliferation to myogenic differentiation of mouse myoblast cells","volume":"6","author":"Tanaka","year":"2014","journal-title":"J Mol Signal"},{"key":"2025120812271856200_msaf310-B47","doi-asserted-by":"publisher","first-page":"9697","DOI":"10.1073\/pnas.0804709105","article-title":"A genome-wide RNAi screen for Wnt\/\u03b2-catenin pathway components identifies unexpected roles for TCF transcription factors in cancer","volume":"105","author":"Tang","year":"2008","journal-title":"Proc Natl Acad Sci U S A."},{"key":"2025120812271856200_msaf310-B48","doi-asserted-by":"publisher","first-page":"602","DOI":"10.1016\/j.tcb.2012.07.008","article-title":"Wnt signaling in myogenesis","volume":"22","author":"von Maltzahn","year":"2012","journal-title":"Trends Cell Biol"},{"key":"2025120812271856200_msaf310-B49","doi-asserted-by":"publisher","first-page":"D113","DOI":"10.1093\/nar\/gkx988","article-title":"HEDD: human enhancer disease database","volume":"46","author":"Wang","year":"2018","journal-title":"Nucleic Acids Res"},{"key":"2025120812271856200_msaf310-B50","doi-asserted-by":"publisher","first-page":"162","DOI":"10.1007\/BF00176060","article-title":"The phylogeny of testicular descent","volume":"6","author":"Williams","year":"1991","journal-title":"Pediatric Surg Int"},{"key":"2025120812271856200_msaf310-B51","doi-asserted-by":"publisher","first-page":"1586","DOI":"10.1093\/molbev\/msm088","article-title":"PAML 4: phylogenetic analysis by maximum likelihood","volume":"24","author":"Yang","year":"2007","journal-title":"Mol Biol Evol"},{"key":"2025120812271856200_msaf310-B52","doi-asserted-by":"publisher","first-page":"7652","DOI":"10.12659\/MSM.917110","article-title":"Long non-coding RNA plasmacytoma variant translocation 1 (PVT1) enhances proliferation, migration, and epithelial-mesenchymal transition (EMT) of pituitary adenoma cells by activating \u03b2-catenin, c-Myc, and cyclin D1 expression","volume":"25","author":"Zhang","year":"2019","journal-title":"Med Scie Monitor"},{"key":"2025120812271856200_msaf310-B53","doi-asserted-by":"publisher","first-page":"132.e1","DOI":"10.1016\/j.jpurol.2015.02.007","article-title":"The variations in the AXIN1 gene and susceptibility to cryptorchidism","volume":"11","author":"Zhou","year":"2015","journal-title":"J Pediatric Urol"},{"key":"2025120812271856200_msaf310-B54","doi-asserted-by":"publisher","first-page":"71","DOI":"10.1097\/MCD.0000000000000401","article-title":"A novel variant in WNT5A responsible for Robinow syndrome in a male fetus with limb shortening and agenesis of the penis","volume":"31","author":"Zhu","year":"2022","journal-title":"Clin Dysmorphol"},{"key":"2025120812271856200_msaf310-B55","doi-asserted-by":"publisher","first-page":"681","DOI":"10.1210\/mend.13.5.0272","article-title":"Targeted disruption of the Insl3 gene causes bilateral cryptorchidism","volume":"13","author":"Zimmermann","year":"1999","journal-title":"Mol Endocrinol"}],"container-title":["Molecular Biology and Evolution"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/academic.oup.com\/mbe\/advance-article-pdf\/doi\/10.1093\/molbev\/msaf310\/65534399\/msaf310.pdf","content-type":"application\/pdf","content-version":"am","intended-application":"syndication"},{"URL":"https:\/\/academic.oup.com\/mbe\/article-pdf\/42\/12\/msaf310\/65534399\/msaf310.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"syndication"},{"URL":"https:\/\/academic.oup.com\/mbe\/article-pdf\/42\/12\/msaf310\/65534399\/msaf310.pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,12,8]],"date-time":"2025-12-08T17:27:32Z","timestamp":1765214852000},"score":1,"resource":{"primary":{"URL":"https:\/\/academic.oup.com\/mbe\/article\/doi\/10.1093\/molbev\/msaf310\/8342993"}},"subtitle":[],"editor":[{"given":"Xuming","family":"Zhou","sequence":"additional","affiliation":[],"role":[{"role":"editor","vocabulary":"crossref"}]}],"short-title":[],"issued":{"date-parts":[[2025,11,26]]},"references-count":55,"journal-issue":{"issue":"12","published-print":{"date-parts":[[2025,11,28]]}},"URL":"https:\/\/doi.org\/10.1093\/molbev\/msaf310","relation":{},"ISSN":["0737-4038","1537-1719"],"issn-type":[{"type":"print","value":"0737-4038"},{"type":"electronic","value":"1537-1719"}],"subject":[],"published-other":{"date-parts":[[2025,12]]},"published":{"date-parts":[[2025,11,26]]},"article-number":"msaf310"}}