{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,17]],"date-time":"2026-10-17T10:06:28Z","timestamp":1792231588373,"version":"4.4.0"},"reference-count":52,"publisher":"Springer Science and Business Media LLC","issue":"1","license":[{"start":{"date-parts":[[2018,1,26]],"date-time":"2018-01-26T00:00:00Z","timestamp":1516924800000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"},{"start":{"date-parts":[[2018,1,26]],"date-time":"2018-01-26T00:00:00Z","timestamp":1516924800000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"}],"content-domain":{"domain":["link.springer.com"],"crossmark-restriction":false},"short-container-title":["Sci Rep"],"abstract":"<jats:title>Abstract<\/jats:title><jats:p>Here we show that the most venomous spiders in the world are phylogenetically misplaced. Australian atracine spiders (family Hexathelidae), including the notorious Sydney funnel-web spider <jats:italic>Atrax robustus<\/jats:italic>, produce venom peptides that can kill people. Intriguingly, eastern Australian mouse spiders (family Actinopodidae) are also medically dangerous, possessing venom peptides strikingly similar to <jats:italic>Atrax<\/jats:italic> hexatoxins. Based on the standing morphology-based classification, mouse spiders are hypothesized distant relatives of atracines, having diverged over 200 million years ago. Using sequence-capture phylogenomics, we instead show convincingly that hexathelids are non-monophyletic, and that atracines are sister to actinopodids. Three new mygalomorph lineages are elevated to the family level, and a revised circumscription of Hexathelidae is presented. Re-writing this phylogenetic story has major implications for how we study venom evolution in these spiders, and potentially genuine consequences for antivenom development and bite treatment research. More generally, our research provides a textbook example of the applied importance of modern phylogenomic research.<\/jats:p>","DOI":"10.1038\/s41598-018-19946-2","type":"journal-article","created":{"date-parts":[[2018,1,22]],"date-time":"2018-01-22T10:40:31Z","timestamp":1516617631000},"update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":64,"title":["Phylogenomic reclassification of the world\u2019s most venomous spiders (Mygalomorphae, Atracinae), with implications for venom evolution"],"prefix":"10.1038","volume":"8","author":[{"given":"Marshal","family":"Hedin","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-9163-9277","authenticated-orcid":false,"given":"Shahan","family":"Derkarabetian","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Mart\u00edn J.","family":"Ram\u00edrez","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Cor","family":"Vink","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jason E.","family":"Bond","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"297","published-online":{"date-parts":[[2018,1,26]]},"reference":[{"key":"19946_CR1","doi-asserted-by":"publisher","first-page":"785","DOI":"10.1046\/j.1440-1681.2002.03741.x","volume":"29","author":"GM Nicholson","year":"2002","unstructured":"Nicholson, G. M. & Graudins, A. Spiders of medical importance in the Asia-Pacific: Atracotoxin, latrotoxin and related spider neurotoxins. Clin. Exp. Pharm. Physio. 29, 785\u2013794 (2002).","journal-title":"Clin. Exp. Pharm. Physio."},{"key":"19946_CR2","doi-asserted-by":"publisher","first-page":"302","DOI":"10.5694\/mja15.00279","volume":"203","author":"GK Isbister","year":"2015","unstructured":"Isbister, G. K. et al. Catecholamine-induced cardiomyopathy resulting from life-threatening funnel-web spider envenoming. Medical J. Australia. 203, 302\u2013304 (2015).","journal-title":"Medical J. Australia."},{"key":"19946_CR3","doi-asserted-by":"crossref","first-page":"407","DOI":"10.5694\/j.1326-5377.2005.tb06760.x","volume":"182","author":"GK Isbister","year":"2005","unstructured":"Isbister, G. K. et al. Funnel-web spider bite: a systematic review of recorded clinical cases. Medical J. Australia. 182, 407\u2013411 (2005).","journal-title":"Medical J. Australia."},{"key":"19946_CR4","doi-asserted-by":"publisher","first-page":"587","DOI":"10.1016\/j.toxicon.2004.02.006","volume":"43","author":"GM Nicholson","year":"2004","unstructured":"Nicholson, G. M., Little, M. J. & Birinyi-Strachan, L. C. Structure and function of \u03b4-atracotoxins: lethal neurotoxins targeting the voltage-gated sodium channel. Toxicon. 43, 587\u2013599 (2004).","journal-title":"Toxicon."},{"key":"19946_CR5","doi-asserted-by":"publisher","first-page":"601","DOI":"10.1016\/j.toxicon.2004.02.010","volume":"43","author":"H Tedford","year":"2004","unstructured":"Tedford, H., Sollod, B., Maggio, F. & King, G. F. Australian funnel-web spiders: Master insecticide chemists. Toxicon. 43, 601\u2013618 (2004).","journal-title":"Toxicon."},{"issue":"177","key":"19946_CR6","first-page":"1","volume":"15","author":"SS Pineda","year":"2014","unstructured":"Pineda, S. S. et al. Diversification of a single ancestral gene into a successful toxin superfamily in highly venomous Australian funnel-web spiders. BMC Genomics. 15(177), 1\u2013177.16 (2014).","journal-title":"BMC Genomics."},{"key":"19946_CR7","doi-asserted-by":"crossref","unstructured":"King, G. F. & Hardy, M. C. Spider-venom peptides: structure, pharmacology, and potential for control of insect pests. Ann. Rev. Ent. 58, 475\u2013496 (2013).","DOI":"10.1146\/annurev-ento-120811-153650"},{"key":"19946_CR8","doi-asserted-by":"publisher","first-page":"4366","DOI":"10.3390\/toxins7104366","volume":"7","author":"V Herzig","year":"2015","unstructured":"Herzig, V. & King, G. F. The cystine knot is responsible for the exceptional stability of the insecticidal spider toxin \u03c9-Hexatoxin-Hv1a. Toxins. 7, 4366\u20134380 (2015).","journal-title":"Toxins."},{"key":"19946_CR9","doi-asserted-by":"publisher","first-page":"201614728","DOI":"10.1073\/pnas.1614728114","volume":"114","author":"IR Chassagnon","year":"2017","unstructured":"Chassagnon, I. R. et al. Potent neuroprotection after stroke afforded by a double-knot spider-venom peptide that inhibits acid-sensing ion channel 1a. Proc. Nat. Acad. Sci. 114, 201614728 (2017).","journal-title":"Proc. Nat. Acad. Sci."},{"key":"19946_CR10","first-page":"113","volume":"5","author":"MR Gray","year":"1988","unstructured":"Gray, M. R. Aspects of the systematics of the Australian funnel web spiders (Araneae: Hexathelidae: Atracinae) based upon morphological and electrophoretic data. Australian Ent. Soc. Misc. Pub. 5, 113\u2013125 (1988).","journal-title":"Australian Ent. Soc. Misc. Pub."},{"key":"19946_CR11","doi-asserted-by":"publisher","first-page":"285","DOI":"10.3853\/j.0067-1975.62.2010.1556","volume":"62","author":"MR Gray","year":"2010","unstructured":"Gray, M. R. A revision of the Australian funnel-web spiders (Hexathelidae: Atracinae). Records Australian Mus. 62, 285\u2013392 (2010).","journal-title":"Records Australian Mus."},{"key":"19946_CR12","first-page":"1","volume":"182","author":"RJ Raven","year":"1985","unstructured":"Raven, R. J. The spider infraorder Mygalomorphae (Araneae): Cladistics and systematics. Bull. Amer. Mus. Nat. Hist. 182, 1\u2013180 (1985).","journal-title":"Bull. Amer. Mus. Nat. Hist."},{"issue":"6","key":"19946_CR13","doi-asserted-by":"publisher","first-page":"e38753","DOI":"10.1371\/journal.pone.0038753","volume":"7","author":"JE Bond","year":"2012","unstructured":"Bond, J. E., Hendrixson, B. E., Hamilton, C. A. & Hedin, M. A reconsideration of the classification of the spider infraorder Mygalomorphae (Arachnida: Araneae) based on three nuclear genes and morphology. PLoS One. 7(6), e38753 (2012).","journal-title":"PLoS One."},{"key":"19946_CR14","doi-asserted-by":"crossref","unstructured":"Rash, L. D., Birinyi-Strachan, L. C., Nicholson, G. M. & Hodgson, W. C. Neurotoxic activity of venom from the Australian Eastern mouse spider (Missulena bradleyi) involves modulation of sodium channel gating. British J. Pharm. 130, 1817\u20131824 (2000).","DOI":"10.1038\/sj.bjp.0703494"},{"issue":"1\u20132","key":"19946_CR15","doi-asserted-by":"publisher","first-page":"211","DOI":"10.1016\/S0014-5793(03)01175-X","volume":"554","author":"SJ Gunning","year":"2003","unstructured":"Gunning, S. J. et al. Isolation of \u03b4-missulenatoxin-Mb1a, the major vertebrate-active spider \u03b4-toxin from the venom of Missulena bradleyi (Actinopodidae). FEBS Letters. 554(1\u20132), 211\u2013218 (2003).","journal-title":"FEBS Letters."},{"key":"19946_CR16","doi-asserted-by":"crossref","first-page":"225","DOI":"10.5694\/j.1326-5377.2004.tb05890.x","volume":"180","author":"GK Isbister","year":"2004","unstructured":"Isbister, G. K. Mouse spider bites (Missulena spp.) and their medical importance. Medical J. Australia. 180, 225\u2013227 (2004).","journal-title":"Medical J. Australia."},{"key":"19946_CR17","doi-asserted-by":"publisher","first-page":"394","DOI":"10.1016\/j.ympev.2006.07.018","volume":"42","author":"NA Ayoub","year":"2007","unstructured":"Ayoub, N. A., Garb, J. E., Hedin, M. & Hayashi, C. Y. Utility of the nuclear protein-coding gene, elongation factor-1gamma (EF-1\u03b3), for spider systematics, emphasizing family level relationships of tarantulas and their kin (Araneae: Mygalomorphae). Mol. Phylo. Evol. 42, 394\u2013409 (2007).","journal-title":"Mol. Phylo. Evol."},{"key":"19946_CR18","doi-asserted-by":"crossref","unstructured":"Opatova, V. & Arnedo, M. From Gondwana to Europe: Inferring the origins of Mediterranean Macrothele spiders (Araneae: Hexathelidae) and the limits of the family Hexathelidae. Invert. Syst. 28, 361\u2013374 (2014).","DOI":"10.1071\/IS14004"},{"key":"19946_CR19","doi-asserted-by":"publisher","first-page":"212","DOI":"10.1186\/s12862-016-0769-y","volume":"16","author":"CA Hamilton","year":"2016","unstructured":"Hamilton, C. A., Lemmon, A. R., Lemmon, E. M. & Bond, J. E. Expanding anchored hybrid enrichment to resolve both deep and shallow relationships within the spider tree of life. BMC Evol. Biol. 16, 212 (2016).","journal-title":"BMC Evol. Biol."},{"key":"19946_CR20","doi-asserted-by":"publisher","first-page":"574","DOI":"10.1111\/cla.12182","volume":"33","author":"WC Wheeler","year":"2017","unstructured":"Wheeler, W. C. et al. The spider tree of life: phylogeny of Araneae based on target-gene analyses from an extensive taxon sampling. Cladistics. 33, 574\u2013616 (2017).","journal-title":"Cladistics."},{"key":"19946_CR21","unstructured":"World Spider Catalog. Natural History Museum Bern, version 18. 5. http:\/\/wsc.nmbe.ch. 10.24436\/2 (2017)."},{"key":"19946_CR22","first-page":"1","volume":"3056","author":"P Goloboff","year":"1993","unstructured":"Goloboff, P. A reanalysis of mygalomorph spider families (Araneae). Am. Mus. Novitates. 3056, 1\u201332 (1993).","journal-title":"Am. Mus. Novitates."},{"key":"19946_CR23","doi-asserted-by":"publisher","first-page":"454","DOI":"10.1016\/j.ympev.2006.05.017","volume":"41","author":"M Hedin","year":"2006","unstructured":"Hedin, M. & Bond, J. E. Molecular phylogenetics of the spider infraorder Mygalomorphae using nuclear rRNA genes (18S and 28S): Conflict and agreement with the current system of classification. Mol. Phylo. Evol. 41, 454\u2013471 (2006).","journal-title":"Mol. Phylo. Evol."},{"issue":"8","key":"19946_CR24","doi-asserted-by":"publisher","first-page":"e0161531","DOI":"10.1371\/journal.pone.0161531","volume":"11","author":"BB Blaimer","year":"2016","unstructured":"Blaimer, B. B., Lloyd, M. W., Guillory, W. X. & Brady, S. G. Sequence capture and phylogenetic utility of genomic ultraconserved elements obtained from pinned insect specimens. PLoS ONE. 11(8), e0161531 (2016).","journal-title":"PLoS ONE."},{"key":"19946_CR25","doi-asserted-by":"publisher","first-page":"1189","DOI":"10.1111\/1755-0998.12466","volume":"16","author":"JE McCormack","year":"2016","unstructured":"McCormack, J. E., Tsai, W. L. E. & Faircloth, B. C. Sequence capture of ultraconserved elements from bird museum specimens. Mol. Ecol. Res. 16, 1189\u20131203 (2016).","journal-title":"Mol. Ecol. Res."},{"key":"19946_CR26","doi-asserted-by":"publisher","unstructured":"Ruane, S. & Austin, C. C. Phylogenomics using formalin-fixed and 100+ year-old intractable natural history specimens. Mol. Ecol. Res. https:\/\/doi.org\/10.1111\/1755-0998.12655 (2017).","DOI":"10.1111\/1755-0998.12655"},{"key":"19946_CR27","first-page":"251","volume":"8","author":"RJ Raven","year":"1980","unstructured":"Raven, R. J. The evolution and biogeography of the mygalomorph spider family Hexathelidae (Araneae, Chelicerata). J. Arachnology. 8, 251\u2013266 (1980).","journal-title":"J. Arachnology."},{"key":"19946_CR28","doi-asserted-by":"publisher","first-page":"292","DOI":"10.1016\/j.jprot.2013.01.002","volume":"80","author":"A Palagi","year":"2013","unstructured":"Palagi, A. et al. Unravelling the complex venom landscapes of lethal Australian funnel-web spiders (Hexathelidae: Atracinae) using LC-MALDI-TOF mass spectrometry. J. Proteomics. 80, 292\u2013310 (2013).","journal-title":"J. Proteomics."},{"key":"19946_CR29","doi-asserted-by":"publisher","first-page":"259","DOI":"10.1016\/S0041-0101(01)00210-0","volume":"40","author":"A Graudins","year":"2002","unstructured":"Graudins, A., Wilson, D., Alewood, P. F., Broady, K. W. & Nicholson, G. M. Cross-reactivity of Sydney funnel-web spider antivenom: neutralization of the in vitro toxicity of other Australian funnel-web (Atrax and Hadronyche) spider venoms. Toxicon. 40, 259\u2013266 (2002).","journal-title":"Toxicon."},{"key":"19946_CR30","doi-asserted-by":"publisher","first-page":"133","DOI":"10.1016\/j.toxicon.2003.11.009","volume":"43","author":"G Isbister","year":"2004","unstructured":"Isbister, G. & Gray, M. Bites by Australian mygalomorph spiders (Araneae, Mygalomorphae), including funnel-web spiders (Atracinae) and mouse spiders (Actinopodidae: Missulena spp). Toxicon. 43, 133\u201340 (2004).","journal-title":"Toxicon."},{"key":"19946_CR31","unstructured":"Mullen, G. & Vetter, R. S. Spiders (Araneae), in Medical and Veterinary Entomology, Mullen, G., Durden, L., eds. Academic Press, New York, 413\u2013432 (2009)."},{"issue":"1","key":"19946_CR32","doi-asserted-by":"publisher","first-page":"218","DOI":"10.1111\/j.1469-7998.1901.tb08176.x","volume":"71","author":"HR Hogg","year":"1901","unstructured":"Hogg, H. R. On Australian and New Zealand spiders of the suborder Mygalomorphae. Proc. Zool. Soc. London. 71(1), 218\u2013279 (1901).","journal-title":"Proc. Zool. Soc. London."},{"issue":"1\u201372","key":"19946_CR33","first-page":"126","volume":"2","author":"RR Forster","year":"1968","unstructured":"Forster, R. R. The spiders of New Zealand. Part II. Ctenizidae, Dipluridae. Otago Mus. Bull. 2(1\u201372), 126\u2013180 (1968).","journal-title":"Otago Mus. Bull."},{"key":"19946_CR34","doi-asserted-by":"publisher","first-page":"812","DOI":"10.1111\/1755-0998.12621","volume":"17","author":"J Starrett","year":"2017","unstructured":"Starrett, J. et al. High phylogenetic utility of an Ultraconserved element probe set designed for Arachnida. Mol. Ecol. Res. 17, 812\u2013823 (2017).","journal-title":"Mol. Ecol. Res."},{"key":"19946_CR35","doi-asserted-by":"publisher","unstructured":"Glenn, T. C. et al. Adapterama I: universal stubs and primers for thousands of dual-indexed Illumina libraries (iTru & iNext). bioRxiv. https:\/\/doi.org\/10.1101\/049114 (2016).","DOI":"10.1101\/049114"},{"key":"19946_CR36","doi-asserted-by":"publisher","first-page":"1103","DOI":"10.1111\/2041-210X.12754","volume":"8","author":"BC Faircloth","year":"2017","unstructured":"Faircloth, B. C. Identifying conserved genomic elements and designing universal probe sets to enrich them. Methods Ecol. Evol. 8, 1103\u20131112 (2017).","journal-title":"Methods Ecol. Evol."},{"key":"19946_CR37","doi-asserted-by":"publisher","first-page":"786","DOI":"10.1093\/bioinformatics\/btv646","volume":"32","author":"BC Faircloth","year":"2016","unstructured":"Faircloth, B. C. PHYLUCE is a software package for the analysis of conserved genomic loci. Bioinformatics. 32, 786\u2013788 (2016).","journal-title":"Bioinformatics."},{"key":"19946_CR38","unstructured":"Faircloth, B. C. Illumiprocessor: a trimmomatic wrapper for parallel adapter and quality trimming. http:\/\/dx.doi.org\/10.6079\/J9ILL (2013)."},{"key":"19946_CR39","doi-asserted-by":"publisher","first-page":"821","DOI":"10.1101\/gr.074492.107","volume":"18","author":"DR Zerbino","year":"2008","unstructured":"Zerbino, D. R. & Birney, E. Velvet: Algorithms for de novo short read assembly using de Bruijn graphs. Genome Res. 18, 821\u2013829 (2008).","journal-title":"Genome Res."},{"key":"19946_CR40","doi-asserted-by":"publisher","first-page":"644","DOI":"10.1038\/nbt.1883","volume":"29","author":"MG Grabherr","year":"2011","unstructured":"Grabherr, M. G. et al. Full-length transcriptome assembly from RNA-Seq data without a reference genome. Nat. Biotech. 29, 644\u2013652 (2011).","journal-title":"Nat. Biotech."},{"key":"19946_CR41","doi-asserted-by":"publisher","first-page":"772","DOI":"10.1093\/molbev\/mst010","volume":"30","author":"D Katoh","year":"2013","unstructured":"Katoh, D. & Standley, D. M. MAFFT multiple sequence alignment software version 7: improvements in performance and usability. Mol. Biol. Evol. 30, 772\u2013780 (2013).","journal-title":"Mol. Biol. Evol."},{"key":"19946_CR42","doi-asserted-by":"publisher","first-page":"540","DOI":"10.1093\/oxfordjournals.molbev.a026334","volume":"17","author":"J Castresana","year":"2000","unstructured":"Castresana, J. Selection of conserved blocks from multiple alignments for their use in phylogenetic analysis. Mol. Biol. Evol. 17, 540\u2013552 (2000).","journal-title":"Mol. Biol. Evol."},{"key":"19946_CR43","doi-asserted-by":"publisher","first-page":"564","DOI":"10.1080\/10635150701472164","volume":"56","author":"G Talavera","year":"2007","unstructured":"Talavera, G. & Castresana, J. Improvement of phylogenies after removing divergent and ambiguously aligned blocks from protein sequence alignments. Syst. Biol. 56, 564\u2013577 (2007).","journal-title":"Syst. Biol."},{"key":"19946_CR44","doi-asserted-by":"publisher","first-page":"1312","DOI":"10.1093\/bioinformatics\/btu033","volume":"30","author":"A Stamatakis","year":"2014","unstructured":"Stamatakis, A. RAxML version 8: a tool for phylogenetic analysis and post-analysis of large phylogenies. Bioinformatics. 30, 1312\u20131313 (2014).","journal-title":"Bioinformatics."},{"key":"19946_CR45","doi-asserted-by":"publisher","first-page":"539","DOI":"10.1093\/sysbio\/sys029","volume":"61","author":"F Ronquist","year":"2012","unstructured":"Ronquist, F. et al. MrBayes 3.2: efficient Bayesian phylogenetic inference and model choice across a large model space. Syst. Biol. 61, 539\u2013542 (2012).","journal-title":"Syst. Biol."},{"key":"19946_CR46","doi-asserted-by":"crossref","unstructured":"Miller, M. A., Pfeiffer, W. & Schwartz, T. Creating the CIPRES Science Gateway for inference of large phylogenetic trees in Proceedings of the Gateway Computing Environments Workshop (GCE), 14 Nov. 2010, New Orleans, LA pp 1\u20138 (2010).","DOI":"10.1109\/GCE.2010.5676129"},{"key":"19946_CR47","doi-asserted-by":"publisher","first-page":"1695","DOI":"10.1093\/molbev\/mss020","volume":"29","author":"R Lanfear","year":"2012","unstructured":"Lanfear, R., Calcott, B., Ho, S. Y. & Guindon, S. PartitionFinder: combined selection of partitioning schemes and substitution models for phylogenetic analyses. Mol. Biol. Evol. 29, 1695\u20131701 (2012).","journal-title":"Mol. Biol. Evol."},{"key":"19946_CR48","doi-asserted-by":"publisher","first-page":"i541","DOI":"10.1093\/bioinformatics\/btu462","volume":"30","author":"S Mirarab","year":"2014","unstructured":"Mirarab, S. et al. ASTRAL: genome-scale coalescent-based species tree estimation. Bioinformatics. 30, i541\u2013i548 (2014).","journal-title":"Bioinformatics."},{"key":"19946_CR49","doi-asserted-by":"publisher","first-page":"i44","DOI":"10.1093\/bioinformatics\/btv234","volume":"31","author":"S Mirarab","year":"2015","unstructured":"Mirarab, S. & Warnow, T. ASTRAL-II: coalescent-based species tree estimation with many hundreds of taxa and thousands of genes. Bioinformatics. 31, i44\u2013i52 (2015).","journal-title":"Bioinformatics."},{"key":"19946_CR50","doi-asserted-by":"publisher","first-page":"3317","DOI":"10.1093\/bioinformatics\/btu530","volume":"30","author":"J Chifman","year":"2014","unstructured":"Chifman, J. & Kubatko, L. Quartet inference from SNP data under the coalescent. Bioinformatics. 30, 3317\u20133324 (2014).","journal-title":"Bioinformatics."},{"key":"19946_CR51","doi-asserted-by":"publisher","first-page":"35","DOI":"10.1016\/j.jtbi.2015.03.006","volume":"374","author":"J Chifman","year":"2015","unstructured":"Chifman, J. & Kubatko, L. Identifiability of the unrooted species tree topology under the coalescent model with time-reversible substitution processes, site-specific rate variation, and invariable sites. J. Theoret. Biol. 374, 35\u201347 (2015).","journal-title":"J. Theoret. Biol."},{"key":"19946_CR52","unstructured":"Swofford, D. L. PAUP*: phylogenetic analysis using parsimony, version 4.0 b10 (2003)."}],"container-title":["Scientific Reports"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.nature.com\/articles\/s41598-018-19946-2","content-type":"text\/html","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/www.nature.com\/articles\/s41598-018-19946-2.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/www.nature.com\/articles\/s41598-018-19946-2.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2022,12,21]],"date-time":"2022-12-21T01:28:49Z","timestamp":1671586129000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.nature.com\/articles\/s41598-018-19946-2"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2018,1,26]]},"references-count":52,"journal-issue":{"issue":"1","published-online":{"date-parts":[[2018,12]]}},"alternative-id":["19946"],"URL":"https:\/\/doi.org\/10.1038\/s41598-018-19946-2","relation":{},"ISSN":["2045-2322"],"issn-type":[{"value":"2045-2322","type":"electronic"}],"subject":[],"published":{"date-parts":[[2018,1,26]]},"assertion":[{"value":"10 November 2017","order":1,"name":"received","label":"Received","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"10 January 2018","order":2,"name":"accepted","label":"Accepted","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"26 January 2018","order":3,"name":"first_online","label":"First Online","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"The authors declare that they have no competing interests.","order":1,"name":"Ethics","group":{"name":"EthicsHeading","label":"Competing Interests"}}],"article-number":"1636"}}