{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,2,24]],"date-time":"2026-02-24T05:34:48Z","timestamp":1771911288307,"version":"3.50.1"},"reference-count":124,"publisher":"Springer Science and Business Media LLC","issue":"1","license":[{"start":{"date-parts":[[2024,11,14]],"date-time":"2024-11-14T00:00:00Z","timestamp":1731542400000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"},{"start":{"date-parts":[[2024,11,14]],"date-time":"2024-11-14T00:00:00Z","timestamp":1731542400000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"}],"funder":[{"name":"Stiftung Leibniz-Institut zur Analyse des Biodiversit\u00e4tswandels (LIB)"}],"content-domain":{"domain":["link.springer.com"],"crossmark-restriction":false},"short-container-title":["Sci Rep"],"abstract":"<jats:title>Abstract<\/jats:title><jats:p>The central nervous system (CNS) of Chelicerata has remained conserved since the Cambrian, yet few studies have examined its variability within chelicerate orders including Scorpiones. The scorpion CNS comprises the prosomal ganglion and opisthosomal ventral nerve cord. We visualize the scorpion CNS with microCT, explore morphological variation across taxa, compare the scorpion CNS to other arachnids, and create a terminology glossary and literature review to assist future studies. Six scorpion species were microCT scanned. Scan quality varied and most structures in the prosomal ganglion could only be observed in <jats:italic>Paruroctonus becki<\/jats:italic> (Vaejovidae). Major nerves and the first opisthosomal ganglion were visible in nearly all taxa. We present the most detailed 3D-rendering of the scorpion prosomal ganglion to date. Our results corroborate existing research and find the scorpion CNS to be conserved. Nearly all structures reported previously in the prosomal ganglion were located in similar positions in <jats:italic>P. becki<\/jats:italic>, and nerve morphology was conserved across examined families. Despite similarities, we report differences from the literature, observe taxonomic variation in prosomal ganglion shape, and confirm positional variation for the first opisthosomal ganglion. This study serves as a starting point for microCT analysis of the scorpion CNS, and future work should include more distantly related, size variable taxa to better elucidate these findings.<\/jats:p>","DOI":"10.1038\/s41598-024-76917-6","type":"journal-article","created":{"date-parts":[[2024,11,14]],"date-time":"2024-11-14T12:42:58Z","timestamp":1731588178000},"update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":2,"title":["Revisiting the scorpion central nervous system using microCT"],"prefix":"10.1038","volume":"14","author":[{"given":"Stephanie F.","family":"Loria","sequence":"first","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Valentin L.","family":"Ehrenthal","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Lauren A.","family":"Esposito","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]}],"member":"297","published-online":{"date-parts":[[2024,11,14]]},"reference":[{"issue":"20","key":"76917_CR1","doi-asserted-by":"publisher","first-page":"R989","DOI":"10.1016\/j.cub.2016.09.012","volume":"26","author":"NJ Strausfeld","year":"2016","unstructured":"Strausfeld, N. J., Ma, X. & Edgecombe, G. D. Fossils and the evolution of the arthropod brain. Curr. Biol. 26(20), R989-1000. https:\/\/doi.org\/10.1016\/j.cub.2016.09.012 (2016).","journal-title":"Curr. Biol."},{"issue":"4","key":"76917_CR2","doi-asserted-by":"publisher","first-page":"659","DOI":"10.1144\/jgs2017-103","volume":"175","author":"C Yang","year":"2018","unstructured":"Yang, C., Li, X. H., Zhu, M., Condon, D. J. & Chen, J. Geochronological constraint on the Cambrian Chengjiang biota, South China. J. Geol. Soc. 175(4), 659\u2013666. https:\/\/doi.org\/10.1144\/jgs2017-103 (2018).","journal-title":"J. Geol. Soc."},{"issue":"1","key":"76917_CR3","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1186\/s12983-022-00459-8","volume":"19","author":"K Frankowski","year":"2022","unstructured":"Frankowski, K., Miyazaki, K. & Brenneis, G. A microCT-based atlas of the central nervous system and midgut in sea spiders (Pycnogonida) sheds first light on evolutionary trends at the family level. Front. Zool. 19(1), 1\u201334. https:\/\/doi.org\/10.1186\/s12983-022-00459-8 (2022).","journal-title":"Front. Zool."},{"key":"76917_CR4","doi-asserted-by":"crossref","unstructured":"Contreras-F\u00e9lix, G. A., del-Pozo, O. G. & Navarrete-Heredia, J. L. A new species of Vaejovis from the mountains of west Mexico (Scorpiones: Vaejovidae). Dugesiana 30(2), 229\u2013245 (2023).","DOI":"10.32870\/dugesiana.v30i2.7306"},{"key":"76917_CR5","first-page":"1","volume":"82","author":"KS Babu","year":"1965","unstructured":"Babu, K. S. Anatomy of the central nervous system of arachnids. Zoologische Jahrbucher, Abteilung f\u00fcr Anatomie und Ontogenie. 82, 1\u2013154 (1965).","journal-title":"Zoologische Jahrbucher, Abteilung f\u00fcr Anatomie und Ontogenie."},{"issue":"2","key":"76917_CR6","doi-asserted-by":"publisher","first-page":"156","DOI":"10.1016\/j.asd.2016.11.003","volume":"46","author":"PO Steinhoff","year":"2017","unstructured":"Steinhoff, P. O. et al. The synganglion of the jumping spider Marpissa muscosa (Arachnida: Salticidae): Insights from histology, immunohistochemistry and microCT analysis. Arthropod Struct. Dev. 46(2), 156\u2013170. https:\/\/doi.org\/10.1016\/j.asd.2016.11.003 (2017).","journal-title":"Arthropod Struct. Dev."},{"issue":"1","key":"76917_CR7","doi-asserted-by":"publisher","first-page":"26","DOI":"10.1186\/s40851-019-0137-z","volume":"5","author":"A Sombke","year":"2019","unstructured":"Sombke, A., Klann, A. E., Lipke, E. & Wolf, H. Primary processing neuropils associated with the malleoli of camel spiders (Arachnida, Solifugae): A re-evaluation of axonal pathways. Zool. Lett. 5(1), 26. https:\/\/doi.org\/10.1186\/s40851-019-0137-z (2019).","journal-title":"Zool. Lett."},{"key":"76917_CR8","first-page":"9","volume-title":"The Biology of Scorpions","author":"JT Hjelle","year":"1990","unstructured":"Hjelle, J. T. Anatomy and morphology. In The Biology of Scorpions (ed. Polis, G. A.) 9\u201363 (Stanford University Press, 1990)."},{"key":"76917_CR9","first-page":"341","volume-title":"The Biology of Scorpions","author":"TM Root","year":"1990","unstructured":"Root, T. M. Neurobiology. In The Biology of Scorpions (ed. Polis, G. A.) 341\u2013413 (Stanford University Press, 1990)."},{"key":"76917_CR10","doi-asserted-by":"publisher","first-page":"29","DOI":"10.1186\/1742-9994-7-29","volume":"7","author":"S Richter","year":"2010","unstructured":"Richter, S. et al. Invertebrate neurophylogeny: Suggested terms and definitions for a neuroanatomical glossary. Front. Zool. 7, 29. https:\/\/doi.org\/10.1186\/1742-9994-7-29 (2010).","journal-title":"Front. Zool."},{"issue":"12","key":"76917_CR11","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pone.0243753","volume":"15","author":"D Drozd","year":"2020","unstructured":"Drozd, D., Wolf, H. & Stemme, T. Structure of the pecten neuropil pathway and its innervation by bimodal peg afferents in two scorpion species. PLoS One 15(12), e0243753. https:\/\/doi.org\/10.1371\/journal.pone.0243753 (2020).","journal-title":"PLoS One"},{"key":"76917_CR12","doi-asserted-by":"publisher","first-page":"395","DOI":"10.1007\/s00427-006-0085-4","volume":"216","author":"G Scholtz","year":"2006","unstructured":"Scholtz, G. & Edgecombe, G. D. The evolution of arthropod heads: reconciling morphological, developmental and palaeontological evidence. Dev. Genes Evol. 216, 395\u2013415. https:\/\/doi.org\/10.1007\/s00427-006-0085-4 (2006).","journal-title":"Dev. Genes Evol."},{"issue":"4","key":"76917_CR13","doi-asserted-by":"publisher","first-page":"317","DOI":"10.1111\/j.1463-6395.2007.00284.x","volume":"88","author":"J Bitsch","year":"2007","unstructured":"Bitsch, J. & Bitsch, C. The segmental organization of the head region in Chelicerata: a critical review of recent studies and hypotheses. Acta Zool. 88(4), 317\u2013335. https:\/\/doi.org\/10.1111\/j.1463-6395.2007.00284.x (2007).","journal-title":"Acta Zool."},{"key":"76917_CR14","doi-asserted-by":"publisher","unstructured":"Richter, S., Stein, M., Frase, T. & Szucsich, N. U. The Arthropod Head. In Arthropod Biology and Evolution: Molecules, Development, Morphology (eds Minelli, A. et al.) 223\u2013240 (Springer, 2013). https:\/\/doi.org\/10.1007\/978-3-642-36160-9_10","DOI":"10.1007\/978-3-642-36160-9_10"},{"key":"76917_CR15","doi-asserted-by":"publisher","unstructured":"Wolf, H. In Structure and Evolution of Invertebrate Nervous Systems (eds Schmidt-Rhaesa, A. et al.) 443\u2013452 (Oxford University Press, 2015). https:\/\/doi.org\/10.1093\/acprof:oso\/9780199682201.003.0037","DOI":"10.1093\/acprof:oso\/9780199682201.003.0037"},{"key":"76917_CR16","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1186\/1742-9994-11-28","volume":"11","author":"BJ Klu\u00dfmann-Fricke","year":"2014","unstructured":"Klu\u00dfmann-Fricke, B. J., Pomrehn, S. W. & Wirkner, C. S. A wonderful network unravelled\u2014Detailed description of capillaries in the prosomal ganglion of scorpions. Front. Zool. 11, 1\u20135. https:\/\/doi.org\/10.1186\/1742-9994-11-28 (2014).","journal-title":"Front. Zool."},{"issue":"7","key":"76917_CR17","doi-asserted-by":"publisher","first-page":"1642","DOI":"10.1002\/cne.25045","volume":"529","author":"I Sinakevitch","year":"2021","unstructured":"Sinakevitch, I., Long, S. M. & Gronenberg, W. The central nervous system of whip spiders (Amblypygi): Large mushroom bodies receive olfactory and visual input. J. Comp. Neurol. 529(7), 1642\u20131658. https:\/\/doi.org\/10.1002\/cne.25045 (2021).","journal-title":"J. Comp. Neurol."},{"key":"76917_CR18","doi-asserted-by":"publisher","first-page":"40","DOI":"10.1186\/1742-9994-10-40","volume":"10","author":"T Lehmann","year":"2013","unstructured":"Lehmann, T. & Melzer, R. R. Looking like Limulus?\u2014Retinula axons and visual neuropils of the median and lateral eyes of scorpions. Front. Zool. 10, 40. https:\/\/doi.org\/10.1186\/1742-9994-10-40 (2013).","journal-title":"Front. Zool."},{"issue":"1","key":"76917_CR19","doi-asserted-by":"publisher","first-page":"70","DOI":"10.1002\/jmor.1060","volume":"250","author":"RD Farley","year":"2001","unstructured":"Farley, R. D. Development of segments and appendages in embryos of the desert scorpion Paruroctonus mesaensis (Scorpiones: Vaejovidae). J. Morphol. 250(1), 70\u201388. https:\/\/doi.org\/10.1002\/jmor.1060 (2001).","journal-title":"J. Morphol."},{"issue":"18","key":"76917_CR20","doi-asserted-by":"publisher","first-page":"10665","DOI":"10.1073\/pnas.95.18.10665","volume":"95","author":"WG Damen","year":"1998","unstructured":"Damen, W. G., Hausdorf, M., Seyfarth, E. A. & Tautz, D. A conserved mode of head segmentation in arthropods revealed by the expression pattern of Hox genes in a spider. Proc. Natl. Acad. Sci. 95(18), 10665\u201310670. https:\/\/doi.org\/10.1073\/pnas.95.18.10665 (1998).","journal-title":"Proc. Natl. Acad. Sci."},{"key":"76917_CR21","doi-asserted-by":"publisher","first-page":"9","DOI":"10.1007\/s00427-002-0285-5","volume":"213","author":"B Mittmann","year":"2003","unstructured":"Mittmann, B. & Scholtz, G. Development of the nervous system in the\" head\" of Limulus polyphemus (Chelicerata: Xiphosura): Morphological evidence for a correspondence between the segments of the chelicerae and of the (first) antennae of Mandibulata. Dev. Genes Evol. 213, 9\u201317. https:\/\/doi.org\/10.1007\/s00427-002-0285-5 (2003).","journal-title":"Dev. Genes Evol."},{"issue":"282","key":"76917_CR22","doi-asserted-by":"publisher","first-page":"20150698","DOI":"10.1098\/rspb.2015.0698","volume":"2015","author":"PP Sharma","year":"1808","unstructured":"Sharma, P. P. et al. A conserved genetic mechanism specifies deutocerebral appendage identity in insects and arachnids. Proc. R. Soc. B Biol. Sci. 2015(282), 20150698. https:\/\/doi.org\/10.1098\/rspb.2015.0698 (1808).","journal-title":"Proc. R. Soc. B Biol. Sci."},{"key":"76917_CR23","doi-asserted-by":"crossref","unstructured":"Buxton, B. H. Notes on the anatomy of arachnids. J. Morphol. 29(1), 1\u201325, pl. I\u2013III (1917).","DOI":"10.1002\/jmor.1050290102"},{"issue":"4","key":"76917_CR24","doi-asserted-by":"publisher","first-page":"249","DOI":"10.1002\/cne.900350402","volume":"35","author":"B Hanstr\u00f6m","year":"1923","unstructured":"Hanstr\u00f6m, B. Further notes on the central nervous system of arachnids: Scorpions, phalangids, and trap-door spiders. J. Comp. Neurol. 35(4), 249\u2013274. https:\/\/doi.org\/10.1002\/cne.900350402 (1923).","journal-title":"J. Comp. Neurol."},{"issue":"3","key":"76917_CR25","doi-asserted-by":"publisher","first-page":"151","DOI":"10.1016\/j.zool.2011.10.002","volume":"115","author":"H Wolf","year":"2012","unstructured":"Wolf, H. & Harzsch, S. Serotonin-immunoreactive neurons in scorpion pectine neuropils: Similarities to insect and crustacean primary olfactory centres?. Zoology 115(3), 151\u2013159. https:\/\/doi.org\/10.1016\/j.zool.2011.10.002 (2012).","journal-title":"Zoology"},{"key":"76917_CR26","doi-asserted-by":"crossref","unstructured":"Babu, K. S. Patterns of arrangement and connectivity in the central nervous system of arachnids. In Neurobiology of Arachnids 3\u201319 (Springer, 1985).","DOI":"10.1007\/978-3-642-70348-5_1"},{"issue":"6","key":"76917_CR27","doi-asserted-by":"publisher","first-page":"545","DOI":"10.1016\/j.asd.2012.06.002","volume":"41","author":"BJ Klu\u00dfmann-Fricke","year":"2012","unstructured":"Klu\u00dfmann-Fricke, B. J., Prendini, L. & Wirkner, C. S. Evolutionary morphology of the hemolymph vascular system in scorpions: A character analysis. Arthropod Struct. Dev. 41(6), 545\u2013560. https:\/\/doi.org\/10.1016\/j.asd.2012.06.002 (2012).","journal-title":"Arthropod Struct. Dev."},{"issue":"220","key":"76917_CR28","doi-asserted-by":"publisher","first-page":"101","DOI":"10.1098\/rspl.1882.0019","volume":"34","author":"ERVI Lankester","year":"1882","unstructured":"Lankester, E. R. V. I. Note on the difference in the position of the ganglia of the ventral nerve-cord in three species of scorpion. Proc. R. Soc. Lond. 34(220), 101\u2013104. https:\/\/doi.org\/10.1098\/rspl.1882.0019 (1882).","journal-title":"Proc. R. Soc. Lond."},{"key":"76917_CR29","doi-asserted-by":"crossref","unstructured":"Lankester, E. R. Part V. Notes on certain points in the anatomy and generic characters of scorpions. Trans. Zool. Soc. Lond. 11, 372\u2013384, pl. LXXXII\u2013LXXXIII (1885).","DOI":"10.1111\/j.1096-3642.1885.tb00346.x"},{"key":"76917_CR30","doi-asserted-by":"crossref","unstructured":"Laurie, M. Notes on the anatomy of some scorpions, and its bearing on the classification of the order. Ann. Mag. Nat. Hist. (Ser. 6). 17, 185\u2013193 (1896).","DOI":"10.1080\/00222939608680350"},{"key":"76917_CR31","unstructured":"Millot, J. & Vachon, M. Ordre des scorpions. In Trait\u00e9 de zoologie (ed Grasse, P.-P.) 386\u2013436 (Masson et Cie, 1949)."},{"key":"76917_CR32","doi-asserted-by":"publisher","first-page":"2918","DOI":"10.1002\/cne.25384","volume":"530","author":"D Drozd","year":"2022","unstructured":"Drozd, D., Wolf, H. & Stemme, T. Mechanosensory pathways of scorpion pecten hair sensillae \u2014Adjustment of body height and pecten position. J. Comp. Neurol. 530, 2918\u20132937. https:\/\/doi.org\/10.1002\/cne.25384 (2022).","journal-title":"J. Comp. Neurol."},{"key":"76917_CR33","doi-asserted-by":"publisher","unstructured":"Newport, G. VIII. On the structure, relations, and development of the nervous and circulatory systems, and on the existence of a complete circulation of the blood in vessels, in Myriapoda and macrourous Arachnida.\u2014First series. Philos. Trans. R. Soc. Lond. 133, 243\u2013302. https:\/\/doi.org\/10.1098\/rstl.1843.0013 (1843)","DOI":"10.1098\/rstl.1843.0013"},{"key":"76917_CR34","first-page":"561","volume":"14","author":"L Dufour","year":"1856","unstructured":"Dufour, L. Histoire anatomique et physiologique des scorpions. M\u00e9moires Pr\u00e9sent\u00e9s par Divers Savants \u00e0 l\u2019Acad\u00e9mie des Sciences de l\u2019Institut Imp\u00e9rial de France 14, 561\u2013653 (1856).","journal-title":"M\u00e9moires Pr\u00e9sent\u00e9s par Divers Savants \u00e0 l'Acad\u00e9mie des Sciences de l\u2019Institut Imp\u00e9rial de France"},{"key":"76917_CR35","first-page":"1492","volume":"102","author":"G Saint-Remy","year":"1886","unstructured":"Saint-Remy, G. Recherches sur la structure du cerveau du scorpion. Comptes Rendus de l\u2019Acad\u00e9mie des Sciences. 102, 1492\u20131494 (1886).","journal-title":"Comptes Rendus de l\u2019Acad\u00e9mie des Sciences."},{"key":"76917_CR36","unstructured":"Saint-Remy, G. Structure des centres nerveux chez le scorpion. Bulletin de la Soci\u00e9t\u00e9 des Sciences de Nancy (2) 8(20), xxix (1886)."},{"key":"76917_CR37","unstructured":"Saint-Remy, G. Contribution \u00e0 l'\u00e9tude du cerveau chez les Arthropodes trach\u00e9ates. Archives de Zoologie Exp\u00e9rimentale et G\u00e9nerale. Ser. 2 Supp. 5(2), 1\u2013274 (1887)."},{"key":"76917_CR38","unstructured":"Saint-Remy, G. Structure du cerveau chez le scorpion et la scolopendre. Bulletin de la Soci\u00e9t\u00e9 des Sciences de Nancy (2) 9(21), xxxi\u2013xxxii (1887)."},{"key":"76917_CR39","doi-asserted-by":"publisher","unstructured":"Laurie, M. The embryology of the scorpion (Euscorpius italicus). Q. J. Microsc. Sci. New Ser. 31, 105\u2013141, pl. XIII\u2013XVIII. https:\/\/doi.org\/10.1242\/jcs.s2-31.122.105 (1890).","DOI":"10.1242\/jcs.s2-31.122.105"},{"key":"76917_CR40","doi-asserted-by":"publisher","unstructured":"Laurie, M. Some points in the development of Scorpio fulvipes. Q. J. Microsc. Sci. New Ser. 32, 587\u2013597, pl. XL. https:\/\/doi.org\/10.1242\/jcs.s2-32.128.587 (1891).","DOI":"10.1242\/jcs.s2-32.128.587"},{"issue":"123","key":"76917_CR41","doi-asserted-by":"publisher","first-page":"317","DOI":"10.1242\/jcs.s2-31.123.317","volume":"2","author":"W Patten","year":"1890","unstructured":"Patten, W. Memoirs: On the origin of vertebrates from arachnids. J. Cell Sci. 2(123), 317\u2013378. https:\/\/doi.org\/10.1242\/jcs.s2-31.123.317 (1890).","journal-title":"J. Cell Sci."},{"key":"76917_CR42","unstructured":"Brauer, A. Beitr\u00e4ge zur Kenntnis der Entwicklungsgeschichte des Skorpions.\u00a0I. Zeitschrift f\u00fcr Wissenschaftliche Zoologie 57, 402\u2013432, pl. XIX\u2013XX (1894)."},{"key":"76917_CR43","unstructured":"Brauer, A. 1895. Beitr\u00e4ge zur Kenntnis der Entwicklungsgeschichte des Skorpions.\u00a0II. Zeitschrift f\u00fcr Wissenschaftliche Zoologie 59, 351\u2013435, pl. XXI\u2013XXV (1895)."},{"issue":"1","key":"76917_CR44","doi-asserted-by":"publisher","first-page":"38","DOI":"10.2307\/1535778","volume":"8","author":"JF McClendon","year":"1904","unstructured":"McClendon, J. F. On the anatomy and embryology of the nervous system of the scorpion. Biol. Bull. 8(1), 38\u201355 (1904).","journal-title":"Biol. Bull."},{"key":"76917_CR45","unstructured":"Abd-el-Wahab A. Some notes on the segmentation of the scorpion, Buthus quinquestriatius (H.&E.). Proc. Egypt. Acad. Sci. 7, 75\u201391 (1952)."},{"key":"76917_CR46","unstructured":"Police, G. Ricerche sul Sistema nervosa dell\u2019 Euscorpius italicus. Atti della Reale Accademia delle Scienze Fisiche e Matematiche\u2014Societ\u00e0 Reale di Napoli. Ser. 2 10(7), 1\u201310, figs. 1\u20134 (1901)."},{"key":"76917_CR47","unstructured":"Police, G. Sui centri nervosi sottointestinali dell\u2019 Euscorpius italicus. Bolletino della Societ\u00e0 di Naturalisti Napoli. Ser. 1 15, 1\u201324, figs. 1\u201312 (1902)."},{"key":"76917_CR48","unstructured":"Police, G. Sul sistema nervoso stomatogastrico dello scorpione. Archivio Zoologico: Pubblicato sotto gli auspicii della unione zoologica italiana 1, 179\u2013200, figs. 1\u201310 (1903)."},{"key":"76917_CR49","unstructured":"Police, G. Sui centri nervosi dei cheliceri e del rostro nello skorpione. Bolletino della Societ\u00e0 di Naturalisti Napoli. Ser. 1 18, 130\u2013135 (1904)."},{"issue":"3","key":"76917_CR50","first-page":"49","volume":"23","author":"WA Hilton","year":"1931","unstructured":"Hilton, W. A. Nervous system and sense organs of Scorpionida. J. Entomol. Zool. 23(3), 49\u201355 (1931).","journal-title":"J. Entomol. Zool."},{"key":"76917_CR51","unstructured":"Werner, F. Scorpiones, Pedipalpi. In Klassen und Ordnungen des Tierreichs: 5, IV, 8, Lief. 1, 2 (Scorpiones) (ed Bronn, H. G.) 1\u2013316 (Akademische Verlagsgesellschaft, 1934)."},{"key":"76917_CR52","unstructured":"K\u00e4stner, A. 1. Ordnung der Arachnida: Scorpiones. In Handbuch der Zoologie (ed Krumbach, T.) 117\u2013240 (Walter de Gruyter Verlag, 1941)."},{"key":"76917_CR53","doi-asserted-by":"crossref","unstructured":"Haller, B. \u00dcber das Zentralnervensystem des Skorpions und der Spinnen. Archiv f\u00fcr mikroskopische Anatomie 79(1), 504\u2013524, pl. XXVI (1912).","DOI":"10.1007\/BF02982863"},{"key":"76917_CR54","unstructured":"Holmgren, N. Zur vergleichenden Anatomie des Gehirns: Von Polychaeten. Onychophoren, Xiphosuren, Arachniden, Crustaceen, Myriapoden und Insekten. Vorstudien zu einer Phylogenie der Arthropoden. Kungliga Svenska Vetenskapsakademiens Handlingar. Stockholm 56(1), 1\u2013303, pl. I\u2013XII (1916)."},{"key":"76917_CR55","unstructured":"Hanstr\u00f6m, B. XX. Kapitel. Chelicerata. In Vergleichende Anatomie des Nervensystems der wirbellosen Tiere: Unter Ber\u00fccksichtigung seiner Funktion 361\u2013420 (Springer, 1928)."},{"key":"76917_CR56","unstructured":"Gottlieb, K. \u00dcber das Gehirn des Skorpions. Zeitschrift f\u00fcr wissenschaftliche Zoologie 127, 185\u2013243, pl. III\u2013IV (1926)."},{"issue":"4","key":"76917_CR57","first-page":"121","volume":"14","author":"L Henry","year":"1949","unstructured":"Henry, L. The nervous system and the segmentation of the head in the scorpion (Arachnida). Microentomology 14(4), 121\u2013126 (1949).","journal-title":"Microentomology"},{"key":"76917_CR58","unstructured":"Ch\u2019eng-Pin, P. Morphology and anatomy of the Chinese scorpion Buthus martensi Karsch. Peking Nat. Hist. Bull. 14(2), 103\u2013117, pl. I\u2013II (1939)."},{"key":"76917_CR59","unstructured":"Pampapathi Rao, K. Some aspects of the electrical activity in the central nervous system of the scorpion, Heterometrus swammerdami. In Proceedings of the 16th International Congress of Zoology, Washington DC 69 (1963)."},{"key":"76917_CR60","doi-asserted-by":"publisher","first-page":"523","DOI":"10.1016\/0010-406X(65)90316-6","volume":"16","author":"H Pasantes","year":"1965","unstructured":"Pasantes, H., Tapia, R., Ortega, B. & Massieu, G. Free amino acids and activity of some pyridoxal phosphate-dependent enzymes in the nervous system of three arthropod species. Comp. Biochem. Physiol. 16, 523\u2013529. https:\/\/doi.org\/10.1016\/0010-406X(65)90316-6 (1965).","journal-title":"Comp. Biochem. Physiol."},{"key":"76917_CR61","doi-asserted-by":"publisher","first-page":"1047","DOI":"10.1038\/2131047a0","volume":"11","author":"K Pampapathi Rao","year":"1967","unstructured":"Pampapathi Rao, K. & Gropalakrishna, T. Blood borne factors in circadian rhythm of activity. Nature 11, 1047\u20131048. https:\/\/doi.org\/10.1038\/2131047a0 (1967).","journal-title":"Nature"},{"key":"76917_CR62","doi-asserted-by":"publisher","first-page":"821","DOI":"10.1007\/BF01897897","volume":"25","author":"SAT Venkatachari","year":"1969","unstructured":"Venkatachari, S. A. T. & Devarajulu, N. V. Choline esterase activity in the nervous system and the innervated organs of the scorpion, Heterometrus fulvipes. Experientia 25, 821\u2013822. https:\/\/doi.org\/10.1007\/BF01897897 (1969).","journal-title":"Experientia"},{"key":"76917_CR63","doi-asserted-by":"publisher","first-page":"165","DOI":"10.1242\/jeb.53.1.165","volume":"53","author":"P Sanjeeva-Reddy","year":"1970","unstructured":"Sanjeeva-Reddy, P. & Pampathi, R. K. The central course of the hair afferents and the pattern of contralateral activation in the central nervous system of the scorpion, Heterometrus fulvipes. J. Exp. Biol. 53, 165\u2013170. https:\/\/doi.org\/10.1242\/jeb.53.1.165 (1970).","journal-title":"J. Exp. Biol."},{"issue":"2","key":"76917_CR64","first-page":"341","volume":"164","author":"J Mercier","year":"1970","unstructured":"Mercier, J. & Dessaigne, S. Influence exerc\u00e9e par quelques drogues psycholeptiques sur le comportement du Scorpion (Androctonus australis Hector). Comptes Rendus des S\u00e9ances de la Soci\u00e9t\u00e9 de Biologie et de ses Filiales 164(2), 341\u2013344 (1970).","journal-title":"Comptes Rendus des S\u00e9ances de la Soci\u00e9t\u00e9 de Biologie et de ses Filiales"},{"issue":"6","key":"76917_CR65","first-page":"1368","volume":"165","author":"J Mercier","year":"1971","unstructured":"Mercier, J. & Dessaigne, S. Influence exerc\u00e9e par la gallamine sur la motricit\u00e9 et les r\u00e9flexes de la Mante religieuse et du Scorpion. Comptes Rendus des S\u00e9ances de la Soci\u00e9t\u00e9 de Biologie et de ses Filiales 165(6), 1368\u20131371 (1971).","journal-title":"Comptes Rendus des S\u00e9ances de la Soci\u00e9t\u00e9 de Biologie et de ses Filiales"},{"key":"76917_CR66","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1002\/jmor.1051310102","volume":"131","author":"M Habibulla","year":"1970","unstructured":"Habibulla, M. Neurosecretion in the scorpion Heterometrus swammerdami. J. Morphol. 131, 1\u201316. https:\/\/doi.org\/10.1002\/jmor.1051310102 (1970).","journal-title":"J. Morphol."},{"key":"76917_CR67","doi-asserted-by":"publisher","first-page":"253","DOI":"10.1016\/0016-6480(71)90133-X","volume":"17","author":"M Habibulla","year":"1971","unstructured":"Habibulla, M. Neurosecretion in the brain of a scorpion Heterometrus swammerdami\u2014A histochemical study. Gen. Comp. Endocrinol. 17, 253\u2013255. https:\/\/doi.org\/10.1016\/0016-6480(71)90133-X (1971).","journal-title":"Gen. Comp. Endocrinol."},{"key":"76917_CR68","doi-asserted-by":"publisher","first-page":"499","DOI":"10.1016\/0305-0491(71)90195-7","volume":"39","author":"M Habibulla","year":"1971","unstructured":"Habibulla, M. Effect of temperature on proteins and amino acids of the cephalothoracic nerve mass of the scorpion Heterometrus swammerdami. Comp. Biochem. Physiol. Part B 39, 499\u2013502. https:\/\/doi.org\/10.1016\/0305-0491(71)90195-7 (1971).","journal-title":"Comp. Biochem. Physiol. Part B"},{"key":"76917_CR69","doi-asserted-by":"publisher","unstructured":"Pampapathi Rao, K. & Habibulla, M. Correlation between neurosecretion and some physiological functions of the scorpion Heterometrus swammerdami. Proc. Indian Acad. Sci. Sec. B 77(4), 148\u2013155. https:\/\/doi.org\/10.1007\/BF03045590 (1973).","DOI":"10.1007\/BF03045590"},{"key":"76917_CR70","doi-asserted-by":"publisher","unstructured":"Geethabali, P. R. K. A metasomatic neural photoreceptor in the scorpion. J. Exp. Biol. 58(1), 189\u2013196. https:\/\/doi.org\/10.1242\/jeb.58.1.189 (1973).","DOI":"10.1242\/jeb.58.1.189"},{"issue":"10","key":"76917_CR71","first-page":"1239","volume":"168","author":"M Goyffon","year":"1974","unstructured":"Goyffon, M., Richard, M. & Vennet, R. Activit\u00e9 \u00e9lectrique c\u00e9r\u00e9brale spontan\u00e9e et comportement moteur du scorpion: Int\u00e9r\u00eat pharmacologique. Comptes Rendus des S\u00e9ances de la Soci\u00e9t\u00e9 de Biologie et de ses Filiales 168(10), 1239 (1974).","journal-title":"Comptes Rendus des S\u00e9ances de la Soci\u00e9t\u00e9 de Biologie et de ses Filiales"},{"key":"76917_CR72","first-page":"101","volume":"36","author":"M Goyffon","year":"1975","unstructured":"Goyffon, M. & Niaussat, P. Interrelations entre m\u00e9canismes cholinergiques et monoaminergiques dans le d\u00e9terminisme d l\u2019activit\u00e9 \u00e9lecrique c\u00e9r\u00e9brale spontan\u00e9e du scorpion. Annales d\u2019Endocrinologie (Paris) 36, 101\u2013102 (1975).","journal-title":"Annales d\u2019Endocrinologie (Paris)"},{"key":"76917_CR73","unstructured":"Goyffon, M., Luyckx, J. & Vachon, M. Sur l\u2019existence d\u2019une activit\u00e9 \u00e9lectrique rythmique spontan\u00e9e du syst\u00e8me nerveux c\u00e9phalique de scorpion. Comptes Rendus Hebdomadaires des S\u00e9ances de l\u2019Acad\u00e9mie des Sciences (Paris), Ser. D 280, 873\u2013876 (1975)."},{"key":"76917_CR74","unstructured":"Geethabali. After discharge in the ventral nerve cord of the scorpion Heterometrus. Curr. Sci. 44(19), 705\u2013706 (1975)."},{"key":"76917_CR75","doi-asserted-by":"crossref","unstructured":"Geethabali. Motor excitation with reference to neural photoreception in scorpions. Life Sci. 18(9), 1009\u20131112 (1976).","DOI":"10.1016\/0024-3205(76)90423-9"},{"key":"76917_CR76","doi-asserted-by":"publisher","unstructured":"Geethabali. Central course of photic input in the ventral nerve cord of scorpion (Heterometrus fulvipes). Experientia 32(3), 345\u2013347. https:\/\/doi.org\/10.1007\/BF01940829 (1976).","DOI":"10.1007\/BF01940829"},{"issue":"5","key":"76917_CR77","doi-asserted-by":"publisher","first-page":"690","DOI":"10.1007\/BF01946580","volume":"33","author":"K Raghavaiah","year":"1977","unstructured":"Raghavaiah, K., Sreeramachandramurthy, R., Ramamurthi, P., Satyam, P. & Chandrasekharam, V. The effect of epinephrine and the hyperglycemic factor of the scorpion\u2019s cephalothoracic ganglionic mass (CTGM) on the phosphorylase activity of the hepatopancreas of the scorpion Heterometrus fulvipes. Experientia 33(5), 690\u2013691. https:\/\/doi.org\/10.1007\/BF01946580 (1977).","journal-title":"Experientia"},{"key":"76917_CR78","doi-asserted-by":"publisher","first-page":"238","DOI":"10.1007\/BF02124087","volume":"33","author":"N Vasantha","year":"1977","unstructured":"Vasantha, N., Venkatachari, S. A. T., Murali Mohan, P. & Babu, K. S. On the possible mode of action of neurohormones on cholines-terase activity in the ventral nerve cord of scorpion, Heterometrus fulvipes. Experientia 33, 238\u2013239. https:\/\/doi.org\/10.1007\/BF02124087 (1977).","journal-title":"Experientia"},{"key":"76917_CR79","doi-asserted-by":"publisher","first-page":"65","DOI":"10.1016\/0306-4492(78)90013-8","volume":"59","author":"M Goyffon","year":"1978","unstructured":"Goyffon, M. Amines biogenes et activit\u00e9 \u00e9lectrique spontan\u00e9e du syst\u00e8me nerveux prosomien du scorpion. Comp. Biochem. Physiol. Part C 59, 65\u201374. https:\/\/doi.org\/10.1016\/0306-4492(78)90013-8 (1978).","journal-title":"Comp. Biochem. Physiol. Part C"},{"issue":"5","key":"76917_CR80","first-page":"209","volume":"2","author":"M Goyffon","year":"1979","unstructured":"Goyffon, M. & Francaz, J.-M. Activit\u00e9 \u00e9lectrique c\u00e9r\u00e9brale spontan\u00e9e du scorpion et amino-acides neurotransmetteurs. Revue Arachnologique 2(5), 209\u2013212 (1979).","journal-title":"Revue Arachnologique"},{"key":"76917_CR81","doi-asserted-by":"publisher","first-page":"59","DOI":"10.1016\/0306-4492(80)90072-6","volume":"66","author":"M Goyffon","year":"1980","unstructured":"Goyffon, M., Drouet, J. & Francaz, J.-M. Neurotransmitter aminoacids and spontaneous electrical activity of the prosimian nervous system of the scorpion. Comp. Biochem. Physiol. Part C 66, 59\u201364. https:\/\/doi.org\/10.1016\/0306-4492(80)90072-6 (1980).","journal-title":"Comp. Biochem. Physiol. Part C"},{"key":"76917_CR82","doi-asserted-by":"publisher","first-page":"31","DOI":"10.1007\/BF00613745","volume":"140","author":"RF Bowerman","year":"1980","unstructured":"Bowerman, R. F. & Burrows, M. The morphology and physiology of some walking leg motor neurons in a scorpion. J. Comp. Physiol. A 140, 31\u201342. https:\/\/doi.org\/10.1007\/BF00613745 (1980).","journal-title":"J. Comp. Physiol. A"},{"key":"76917_CR83","doi-asserted-by":"publisher","first-page":"495","DOI":"10.1007\/BF00191715","volume":"174","author":"KS Babu","year":"1994","unstructured":"Babu, K. S. & Jacobdoss, P. Central afferent pathways of long hair sensilla in the ventral nerve cord of the Indian black scorpion, Heterometrus fulvipes Koch. J. Comp. Physiol. A 174, 495\u2013505. https:\/\/doi.org\/10.1007\/BF00191715 (1994).","journal-title":"J. Comp. Physiol. A"},{"key":"76917_CR84","doi-asserted-by":"publisher","first-page":"1033","DOI":"10.1007\/s00359-007-0254-6","volume":"193","author":"N Kladt","year":"2007","unstructured":"Kladt, N., Wolf, H. & Heinzel, H. G. Mechanoreception by cuticular sensilla on the pectines of the scorpion Pandinus cavimanus. J. Comp. Physiol. A 193, 1033\u20131043 (2007).","journal-title":"J. Comp. Physiol. A"},{"key":"76917_CR85","doi-asserted-by":"publisher","first-page":"9","DOI":"10.1007\/s10158-019-0228-8","volume":"19","author":"KL Hughes","year":"2019","unstructured":"Hughes, K. L. & Gaffin, D. D. Investigating sensory processing in the pectines of the striped bark scorpion, Centruroides vittatus. Invertebr. Neurosci. 19, 9. https:\/\/doi.org\/10.1007\/s10158-019-0228-8 (2019).","journal-title":"Invertebr. Neurosci."},{"issue":"3","key":"76917_CR86","doi-asserted-by":"publisher","first-page":"225","DOI":"10.1007\/BF03185013","volume":"91","author":"K Yellamma","year":"1982","unstructured":"Yellamma, K., Subhashini, K., Murali Mohan, P. & Babu, K. S. Microanatomy of the 7th abdominal ganglion and its peripheral nerves in the scorpion, Heterometrus fulvipes. Proc. Anim. Sci. 91(3), 225\u2013234. https:\/\/doi.org\/10.1007\/BF03185013 (1982).","journal-title":"Proc. Anim. Sci."},{"key":"76917_CR87","unstructured":"Lucas, S., Eichstedt, V. D. & Bucherl, W. Sobre o Sistema nervosa de Tityus serrulatus\u2013Titynae\u2013Buthidae\u2013Scorpiones. Mem\u00f3rias do Instituto de Butantan. 32, 15\u201326 (1965)."},{"key":"76917_CR88","first-page":"244","volume":"22","author":"NJ Lane","year":"1980","unstructured":"Lane, N. J. & Harrison, J. B. An unusual form of tight junction in the nervous-system of the scorpion. Eur. J. Cell Biol. 22, 244\u2013244 (1980).","journal-title":"Eur. J. Cell Biol."},{"key":"76917_CR89","doi-asserted-by":"publisher","unstructured":"Khalil, A., Ismail, S. & El-Bakary, Z. Studies on the morphology and histology of the central nervous system of the adult of the Egyptian scorpion Buthus quinquestriatus (H.E.) II\u2014General Histology. Assiut Vet. Med. J. 14(28), 75\u201383. https:\/\/doi.org\/10.21608\/avmj.1985.190235 (1985).","DOI":"10.21608\/avmj.1985.190235"},{"issue":"1","key":"76917_CR90","doi-asserted-by":"publisher","first-page":"502","DOI":"10.1111\/j.1749-6632.1998.tb10614.x","volume":"855","author":"PH Brownell","year":"1998","unstructured":"Brownell, P. H. Glomerular cytoarchitectures in chemosensory systems of arachnids. Ann. N. Y. Acad. Sci. 855(1), 502\u2013507. https:\/\/doi.org\/10.1111\/j.1749-6632.1998.tb10614.x (1998).","journal-title":"Ann. N. Y. Acad. Sci."},{"key":"76917_CR91","doi-asserted-by":"publisher","unstructured":"Horn, A. C. M. & Achaval, M. The gross anatomy of the nervous system of Bothriurus bonariensis (L.C. Koch, 1842) (Scorpiones, Bothriuridae). Braz. J. Biol. 62, 253\u2013262. https:\/\/doi.org\/10.1590\/S1519-69842002000200009 (2002).","DOI":"10.1590\/S1519-69842002000200009"},{"issue":"5","key":"76917_CR92","doi-asserted-by":"publisher","first-page":"401","DOI":"10.1002\/jmor.10512","volume":"268","author":"CS Wirkner","year":"2007","unstructured":"Wirkner, C. S. & Prendini, L. Comparative morphology of the hemolymph vascular system in scorpions\u2014A survey using corrosion casting, MicroCT, and 3D-reconstruction. J. Morphol. 268(5), 401\u2013413. https:\/\/doi.org\/10.1002\/jmor.10512 (2007).","journal-title":"J. Morphol."},{"issue":"1","key":"76917_CR93","doi-asserted-by":"publisher","first-page":"67","DOI":"10.1016\/j.asd.2007.05.003","volume":"37","author":"H Wolf","year":"2008","unstructured":"Wolf, H. The pectine organs of the scorpion, Vaejovis spinigerus: Structure and (glomerular) central projections. Arthropod Struct. Dev. 37(1), 67\u201380. https:\/\/doi.org\/10.1016\/j.asd.2007.05.003 (2008).","journal-title":"Arthropod Struct. Dev."},{"issue":"6","key":"76917_CR94","doi-asserted-by":"publisher","first-page":"753","DOI":"10.1016\/j.asd.2017.10.002","volume":"46","author":"H Wolf","year":"2017","unstructured":"Wolf, H. Scorpions pectines\u2014Idiosyncratic chemo-and mechanosensory organs. Arthropod Struct. Dev. 46(6), 753\u2013764. https:\/\/doi.org\/10.1016\/j.asd.2017.10.002 (2017).","journal-title":"Arthropod Struct. Dev."},{"key":"76917_CR95","doi-asserted-by":"publisher","unstructured":"Wolf, H. & Harzsch, S. Evolution of the arthropod neuromuscular system. 1. Arrangement of muscles and innervation in the walking legs of a scorpion: Vaejovis spinigerus (Wood, 1863) Vaejovidae, Scorpiones, Arachnida. Arthropod Struct. Dev. 31(3), 185\u2013202. https:\/\/doi.org\/10.1016\/S1467-8039(02)00043-9 (2002).","DOI":"10.1016\/S1467-8039(02)00043-9"},{"key":"76917_CR96","doi-asserted-by":"publisher","unstructured":"Wolf, H. & Harzsch, S. Evolution of the arthropod neuromuscular system. 2. Inhibitory innervation of the walking legs of a scorpion: Vaejovis spinigerus (Wood, 1863), Vaejovidae, Scorpiones, Arachnida. Arthropod Struct. Dev. 31(3), 203\u2013215. https:\/\/doi.org\/10.1016\/S1467-8039(02)00044-0 (2002).","DOI":"10.1016\/S1467-8039(02)00044-0"},{"issue":"1","key":"76917_CR97","doi-asserted-by":"publisher","first-page":"38","DOI":"10.1016\/j.cub.2014.10.049","volume":"25","author":"GH Wolff","year":"2015","unstructured":"Wolff, G. H. & Strausfeld, N. J. Genealogical correspondence of mushroom bodies across invertebrate phyla. Curr. Biol. 25(1), 38\u201344. https:\/\/doi.org\/10.1016\/j.cub.2014.10.049 (2015).","journal-title":"Curr. Biol."},{"issue":"7","key":"76917_CR98","doi-asserted-by":"publisher","first-page":"731","DOI":"10.2108\/zsj.21.731","volume":"21","author":"K Mori","year":"2004","unstructured":"Mori, K. & Kuramoto, T. Photosensitivity of the central nervous system of Limulus polyphemus. Zool. Sci. 21(7), 731\u2013737. https:\/\/doi.org\/10.2108\/zsj.21.731 (2004).","journal-title":"Zool. Sci."},{"key":"76917_CR99","doi-asserted-by":"publisher","unstructured":"Franz-Guess, S. & Starck, J. M. Microscopic Anatomy of Eukoenenia spelaea (Peyerimhoff, 1902) (Arachnida: Palpigradi: Eukoeneniidae). Bonn Zool. Bull. Suppl. 65, 1\u2013125. https:\/\/doi.org\/10.20363\/BZB-S-2020.65 (2020).","DOI":"10.20363\/BZB-S-2020.65"},{"key":"76917_CR100","doi-asserted-by":"publisher","first-page":"205","DOI":"10.1007\/s10493-006-9039-9","volume":"40","author":"M van Wijk","year":"2006","unstructured":"van Wijk, M., Wadman, W. J. & Sabelis, M. W. Gross morphology of the central nervous system of a phytoseiid mite. Exp. Appl. Acarol. 40, 205\u2013216. https:\/\/doi.org\/10.1007\/s10493-006-9039-9 (2006).","journal-title":"Exp. Appl. Acarol."},{"issue":"1","key":"76917_CR101","doi-asserted-by":"publisher","first-page":"17","DOI":"10.9729\/AM.2018.48.1.17","volume":"48","author":"YK Park","year":"2018","unstructured":"Park, Y. K., Gu, H. Y., Kwon, H. J., Kim, H. & Moon, M. J. Fine structure of the neuroganglia in the central nervous system of the harvestman Leiobunum japonicum (Arachnida: Opiliones). Appl. Microsc. 48(1), 17\u201326. https:\/\/doi.org\/10.9729\/AM.2018.48.1.17 (2018).","journal-title":"Appl. Microsc."},{"issue":"1","key":"76917_CR102","doi-asserted-by":"publisher","first-page":"25","DOI":"10.3390\/insects13010025","volume":"13","author":"T Stemme","year":"2021","unstructured":"Stemme, T. & Pfeffer, S. E. Anatomy of the nervous system in Chelifer cancroides (Arachnida: Pseudoscorpiones) with a distinct sensory pathway associated with the pedipalps. Insects 13(1), 25. https:\/\/doi.org\/10.3390\/insects13010025 (2021).","journal-title":"Insects"},{"issue":"7","key":"76917_CR103","first-page":"1","volume":"11","author":"J Millot","year":"1945","unstructured":"Millot, J. L\u2019anatomie interne des Ricinulei (Arachnides). Annales des Sciences Naturelles, Zoologie. 11(7), 1\u201329 (1945).","journal-title":"Annales des Sciences Naturelles, Zoologie."},{"key":"76917_CR104","unstructured":"van der Hammen, L. Comparative studies in Chelicerata: I. The Cryptognomae (Ricinulei, Architarbi and Anactinotrichida). Zoologische Verhandelingen 174(1), 3\u201362 (1979)."},{"key":"76917_CR105","doi-asserted-by":"publisher","first-page":"23","DOI":"10.1016\/j.asd.2019.06.002","volume":"51","author":"T Lehmann","year":"2019","unstructured":"Lehmann, T. & Melzer, R. R. The visual system of Thelyphonida (whip scorpions): support for Arachnopulmonata. Arthropod Struct. Dev. 51, 23\u201331. https:\/\/doi.org\/10.1016\/j.asd.2019.06.002 (2019).","journal-title":"Arthropod Struct. Dev."},{"key":"76917_CR106","doi-asserted-by":"publisher","first-page":"52","DOI":"10.1186\/s12983-018-0293-6","volume":"15","author":"T Lehmann","year":"2018","unstructured":"Lehmann, T. & Melzer, R. R. Also looking like Limulus?\u2013Retinula axons and visual neuropils of Amblypygi (whip spiders). Front. Zool. 15, 52. https:\/\/doi.org\/10.1186\/s12983-018-0293-6 (2018).","journal-title":"Front. Zool."},{"key":"76917_CR107","doi-asserted-by":"publisher","first-page":"50","DOI":"10.1186\/s12983-016-0182-9","volume":"13","author":"T Lehmann","year":"2016","unstructured":"Lehmann, T., Lodde-Bensch, E., Melzer, R. R. & Metz, M. The visual system of harvestmen (Opiliones, Arachnida, Chelicerata)\u2014A re-examination. Front. Zool. 13, 50. https:\/\/doi.org\/10.1186\/s12983-016-0182-9 (2016).","journal-title":"Front. Zool."},{"key":"76917_CR108","doi-asserted-by":"publisher","DOI":"10.1016\/j.asd.2020.101024","volume":"60","author":"T Lehmann","year":"2021","unstructured":"Lehmann, T. & Melzer, R. R. Outsourcing a visual neuropil\u2014The central visual system of the median eyes of Galeodes granti Pocock, 1903 (Arachnida: Solifugae). Arthropod Struct. Dev. 60, 101024. https:\/\/doi.org\/10.1016\/j.asd.2020.101024 (2021).","journal-title":"Arthropod Struct. Dev."},{"key":"76917_CR109","doi-asserted-by":"publisher","first-page":"164","DOI":"10.1016\/j.jcz.2017.11.014","volume":"273","author":"T Lehmann","year":"2018","unstructured":"Lehmann, T. & Melzer, R. R. A tiny visual system\u2014Retinula axons and visual neuropils of Neobisium carcinoides (Hermann, 1804) (Chelicerata, Arachnida, Pseudoscorpiones). Zoologischer Anzeiger. 273, 164\u2013172. https:\/\/doi.org\/10.1016\/j.jcz.2017.11.014 (2018).","journal-title":"Zoologischer Anzeiger."},{"issue":"5","key":"76917_CR110","doi-asserted-by":"publisher","first-page":"347","DOI":"10.1016\/j.asd.2008.01.008","volume":"37","author":"U Homberg","year":"2008","unstructured":"Homberg, U. Evolution of the central complex in the arthropod brain with respect to the visual system. Arthropod Struct. Dev. 37(5), 347\u2013362. https:\/\/doi.org\/10.1016\/j.asd.2008.01.008 (2008).","journal-title":"Arthropod Struct. Dev."},{"issue":"1","key":"76917_CR111","doi-asserted-by":"publisher","first-page":"29","DOI":"10.1111\/j.1463-6395.2009.00422.x","volume":"91","author":"R Loesel","year":"2010","unstructured":"Loesel, R. & Heuer, C. M. The mushroom bodies\u2013prominent brain centres of arthropods and annelids with enigmatic evolutionary origin. Acta Zool. 91(1), 29\u201334. https:\/\/doi.org\/10.1111\/j.1463-6395.2009.00422.x (2010).","journal-title":"Acta Zool."},{"issue":"2","key":"76917_CR112","doi-asserted-by":"publisher","first-page":"259","DOI":"10.1002\/cne.24945","volume":"529","author":"SM Long","year":"2021","unstructured":"Long, S. M. Variations on a theme: Morphological variation in the secondary eye visual pathway across the order of Araneae. J. Comp. Neurol. 529(2), 259\u2013280. https:\/\/doi.org\/10.1002\/cne.24945 (2021).","journal-title":"J. Comp. Neurol."},{"issue":"6","key":"76917_CR113","doi-asserted-by":"publisher","first-page":"521","DOI":"10.1016\/j.asd.2011.07.002","volume":"40","author":"R Quesada","year":"2011","unstructured":"Quesada, R. et al. The allometry of CNS size and consequences of miniaturization in orb-weaving and cleptoparasitic spiders. Arthropod Struct. Dev. 40(6), 521\u2013529. https:\/\/doi.org\/10.1016\/j.asd.2011.07.002 (2011).","journal-title":"Arthropod Struct. Dev."},{"key":"76917_CR114","doi-asserted-by":"publisher","unstructured":"Santib\u00e1\u00f1ez\u2010L\u00f3pez, C. E., Ojanguren\u2010Affilastro, A. A., Graham, M. R. & Sharma, P. P. Congruence between ultraconserved element\u2010based matrices and phylotranscriptomic datasets in the scorpion Tree of Life. Cladistics 39(6), 533\u2013547. https:\/\/doi.org\/10.1111\/cla.12551 (2023).","DOI":"10.1111\/cla.12551"},{"issue":"1","key":"76917_CR115","doi-asserted-by":"publisher","first-page":"71","DOI":"10.1111\/cla.12067","volume":"31","author":"L Monod","year":"2015","unstructured":"Monod, L. & Prendini, L. Evidence for Eurogondwana: The roles of dispersal, extinction and vicariance in the evolution and biogeography of Indo-Pacific Hormuridae (Scorpiones: Scorpionoidea). Cladistics 31(1), 71\u2013111. https:\/\/doi.org\/10.1111\/cla.12067 (2015).","journal-title":"Cladistics"},{"issue":"2","key":"76917_CR116","doi-asserted-by":"publisher","first-page":"231","DOI":"10.1002\/jmor.1051840212","volume":"184","author":"RD Farley","year":"1985","unstructured":"Farley, R. D. & Chan, D. J. The ultrastructure of the cardiac ganglion of the desert scorpion, Paruroctonus mesaensis (Scorpionida: Vaejovidae). J. Morphol. 184(2), 231\u2013252. https:\/\/doi.org\/10.1002\/jmor.1051840212 (1985).","journal-title":"J. Morphol."},{"issue":"14","key":"76917_CR117","doi-asserted-by":"publisher","first-page":"2474","DOI":"10.1002\/cne.25343","volume":"530","author":"FA Rivera-Quiroz","year":"2022","unstructured":"Rivera-Quiroz, F. A. & Miller, J. A. Micro-CT visualization of the CNS: Performance of different contrast-enhancing techniques for documenting the spider brain. J. Comp. Neurol. 530(14), 2474\u20132485. https:\/\/doi.org\/10.1002\/cne.25343 (2022).","journal-title":"J. Comp. Neurol."},{"issue":"8","key":"76917_CR118","doi-asserted-by":"publisher","first-page":"1281","DOI":"10.1002\/cne.23741","volume":"523","author":"A Sombke","year":"2015","unstructured":"Sombke, A., Lipke, E., Michalik, P., Uhl, G. & Harzsch, S. Potential and limitations of X-Ray micro-computed tomography in arthropod neuroanatomy: A methodological and comparative survey. J. Comp. Neurol. 523(8), 1281\u20131295. https:\/\/doi.org\/10.1002\/cne.23741 (2015).","journal-title":"J. Comp. Neurol."},{"issue":"6","key":"76917_CR119","doi-asserted-by":"publisher","first-page":"889","DOI":"10.1111\/joa.12449","volume":"228","author":"PM Gignac","year":"2016","unstructured":"Gignac, P. M. et al. Diffusible iodine-based contrast-enhanced computed tomography (diceCT): An emerging tool for rapid, high-resolution, 3-D imaging of metazoan soft tissues. J. Anat. 228(6), 889\u2013909. https:\/\/doi.org\/10.1111\/joa.12449 (2016).","journal-title":"J. Anat."},{"key":"76917_CR120","doi-asserted-by":"publisher","first-page":"11","DOI":"10.1186\/1472-6793-9-11","volume":"9","author":"BD Metscher","year":"2009","unstructured":"Metscher, B. D. MicroCT for comparative morphology: Simple staining methods allow high-contrast 3D imaging of diverse non-mineralized animal tissues. BMC Physiol. 9, 11. https:\/\/doi.org\/10.1186\/1472-6793-9-11 (2009).","journal-title":"BMC Physiol."},{"issue":"1","key":"76917_CR121","doi-asserted-by":"publisher","first-page":"77","DOI":"10.1016\/S1467-8039(02)00017-8","volume":"31","author":"R Loesel","year":"2002","unstructured":"Loesel, R., N\u00e4ssel, D. R. & Strausfeld, N. J. Common design in a unique midline neuropil in the brains of arthropods. Arthropod Struct. Dev. 31(1), 77\u201391. https:\/\/doi.org\/10.1016\/S1467-8039(02)00017-8 (2002).","journal-title":"Arthropod Struct. Dev."},{"key":"76917_CR122","doi-asserted-by":"publisher","first-page":"1857","DOI":"10.1098\/rspb.2006.3536","volume":"273","author":"NJ Strausfeld","year":"2006","unstructured":"Strausfeld, N. J., Strausfeld, C., Stowe, S., Rowell, D. & Loesel, R. Arthropod phylogeny: Onychophoran brain organization suggests an archaic relationship with a chelicerate stem lineage. Proc. R. Soc. B Biol. Sci. 273, 1857\u20131866. https:\/\/doi.org\/10.1098\/rspb.2006.3536 (2006).","journal-title":"Proc. R. Soc. B Biol. Sci."},{"issue":"4","key":"76917_CR123","doi-asserted-by":"publisher","first-page":"325","DOI":"10.1002\/jemt.10140","volume":"58","author":"DD Gaffin","year":"2002","unstructured":"Gaffin, D. D. Electrophysiological analysis of synaptic interactions within peg sensilla of scorpion pectines. Microsc. Res. Tech. 58(4), 325\u2013334. https:\/\/doi.org\/10.1002\/jemt.10140 (2002).","journal-title":"Microsc. Res. Tech."},{"issue":"1792","key":"76917_CR124","doi-asserted-by":"publisher","first-page":"20140661","DOI":"10.1098\/rspb.2014.0661","volume":"281","author":"PP Sharma","year":"2014","unstructured":"Sharma, P. P., Schwager, E. E., Extavour, C. G. & Wheeler, W. C. Hox gene duplications correlate with posterior heteronomy in scorpions. Proc. R. Soc. B Biol. Sci. 281(1792), 20140661. https:\/\/doi.org\/10.1098\/rspb.2014.0661 (2014).","journal-title":"Proc. R. Soc. B Biol. Sci."}],"container-title":["Scientific Reports"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.nature.com\/articles\/s41598-024-76917-6.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/www.nature.com\/articles\/s41598-024-76917-6","content-type":"text\/html","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/www.nature.com\/articles\/s41598-024-76917-6.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2024,11,14]],"date-time":"2024-11-14T13:07:37Z","timestamp":1731589657000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.nature.com\/articles\/s41598-024-76917-6"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2024,11,14]]},"references-count":124,"journal-issue":{"issue":"1","published-online":{"date-parts":[[2024,12]]}},"alternative-id":["76917"],"URL":"https:\/\/doi.org\/10.1038\/s41598-024-76917-6","relation":{},"ISSN":["2045-2322"],"issn-type":[{"value":"2045-2322","type":"electronic"}],"subject":[],"published":{"date-parts":[[2024,11,14]]},"assertion":[{"value":"18 January 2024","order":1,"name":"received","label":"Received","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"17 October 2024","order":2,"name":"accepted","label":"Accepted","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"14 November 2024","order":3,"name":"first_online","label":"First Online","group":{"name":"ArticleHistory","label":"Article History"}},{"order":1,"name":"Ethics","group":{"name":"EthicsHeading","label":"Declarations"}},{"value":"The authors declare no competing interests.","order":2,"name":"Ethics","group":{"name":"EthicsHeading","label":"Competing interests"}},{"value":"No animals used in this study are regulated by the Institutional Animal Care & Use Committee, the International Union for Conservation of Nature or the Convention on International Trade in Endangered Species of Wild Fauna and Flora. Approval for experimental guidelines is not applicable.","order":3,"name":"Ethics","group":{"name":"EthicsHeading","label":"Ethics approval and consent to participate"}}],"article-number":"27961"}}