{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,1,15]],"date-time":"2026-01-15T13:48:54Z","timestamp":1768484934346,"version":"3.49.0"},"reference-count":86,"publisher":"Springer Science and Business Media LLC","issue":"2","license":[{"start":{"date-parts":[[2023,7,27]],"date-time":"2023-07-27T00:00:00Z","timestamp":1690416000000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"},{"start":{"date-parts":[[2023,7,27]],"date-time":"2023-07-27T00:00:00Z","timestamp":1690416000000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"}],"funder":[{"DOI":"10.13039\/501100007051","name":"Uppsala University","doi-asserted-by":"crossref","id":[{"id":"10.13039\/501100007051","id-type":"DOI","asserted-by":"crossref"}]}],"content-domain":{"domain":["link.springer.com"],"crossmark-restriction":false},"short-container-title":["Dev Genes Evol"],"published-print":{"date-parts":[[2023,12]]},"abstract":"<jats:title>Abstract<\/jats:title><jats:p>Spiders represent an evolutionary successful group of chelicerate arthropods. The body of spiders is subdivided into two regions (tagmata). The anterior tagma, the prosoma, bears the head appendages and four pairs of walking legs. The segments of the posterior tagma, the opisthosoma, either lost their appendages during the course of evolution or their appendages were substantially modified to fulfill new tasks such as reproduction, gas exchange, and silk production. Previous work has shown that the homeotic Hox genes are involved in shaping the posterior appendages of spiders. In this paper, we investigate the expression of the posterior Hox genes in a tarantula that possesses some key differences of posterior appendages compared to true spiders, such as the lack of the anterior pair of spinnerets and a second set of book lungs instead of trachea. Based on the observed differences in posterior Hox gene expression in true spiders and tarantulas, we argue that subtle changes in the Hox gene expression of the Hox genes <jats:italic>abdA<\/jats:italic> and <jats:italic>AbdB<\/jats:italic> are possibly responsible for at least some of the morphological differences seen in true spiders <jats:italic>versus<\/jats:italic> tarantulas.<\/jats:p>","DOI":"10.1007\/s00427-023-00707-9","type":"journal-article","created":{"date-parts":[[2023,7,27]],"date-time":"2023-07-27T02:01:16Z","timestamp":1690423276000},"page":"107-121","update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":5,"title":["Expression of posterior Hox genes and opisthosomal appendage development in a mygalomorph spider"],"prefix":"10.1007","volume":"233","author":[{"given":"Ralf","family":"Janssen","sequence":"first","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Matthias","family":"Pechmann","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]}],"member":"297","published-online":{"date-parts":[[2023,7,27]]},"reference":[{"key":"707_CR1","doi-asserted-by":"crossref","unstructured":"Akam M, Averof M, Castelli-Gair J, Dawes R, Falciani F, Ferrier D (1994) The evolving role of Hox genes in arthropods. Development (Suppl):209\u2013215","DOI":"10.1242\/dev.1994.Supplement.209"},{"key":"707_CR2","doi-asserted-by":"publisher","unstructured":"Alberstat EJ, Chung K, Sun DA, Ray S, Patel NH (2022) Combinatorial interactions of Hox genes establish appendage diversity of the amphipod crustacean Parhyale hawaiensis. bioRxiv. https:\/\/doi.org\/10.1101\/2022.03.25.485717","DOI":"10.1101\/2022.03.25.485717"},{"issue":"1","key":"707_CR3","doi-asserted-by":"publisher","first-page":"51","DOI":"10.1016\/S0925-4773(96)00649-1","volume":"62","author":"AC Aplin","year":"1997","unstructured":"Aplin AC, Kaufman TC (1997) Homeotic transformation of legs to mouthparts by proboscipedia expression in Drosophila imaginal discs. Mech Dev 62(1):51\u201360","journal-title":"Mech Dev"},{"issue":"6539","key":"707_CR4","doi-asserted-by":"publisher","first-page":"420","DOI":"10.1038\/376420a0","volume":"376","author":"M Averof","year":"1995","unstructured":"Averof M, Akam M (1995) Hox genes and the diversification of insect and crustacean body plans. Nature 376(6539):420\u2013423","journal-title":"Nature"},{"issue":"1","key":"707_CR5","doi-asserted-by":"publisher","first-page":"42","DOI":"10.1111\/j.1525-142X.2005.05005.x","volume":"7","author":"C Brena","year":"2005","unstructured":"Brena C, Liu PZ, Minelli A, Kaufman TC (2005) Abd-B expression in Porcellio scaber Latreille, 1804 (Isopoda: Crustacea): conserved pattern versus novel roles in development and evolution. Evol Dev 7(1):42\u201350","journal-title":"Evol Dev"},{"issue":"5","key":"707_CR6","doi-asserted-by":"publisher","first-page":"332","DOI":"10.1046\/j.1525-142X.2001.01041.x","volume":"3","author":"GE Budd","year":"2001","unstructured":"Budd GE (2001) Why are arthropods segmented? Evol Dev. 3(5):332\u2013342","journal-title":"Evol Dev."},{"issue":"1","key":"707_CR7","doi-asserted-by":"publisher","first-page":"113","DOI":"10.1016\/0092-8674(86)90372-7","volume":"47","author":"SB Carroll","year":"1986","unstructured":"Carroll SB, Laymon RA, McCutcheon MA, Riley PD, Scott MP (1986) The localization and regulation of Antennapedia protein expression in Drosophila embryos. Cell 47(1):113\u2013122","journal-title":"Cell"},{"issue":"9","key":"707_CR8","doi-asserted-by":"publisher","first-page":"2973","DOI":"10.1242\/dev.121.9.2973","volume":"121","author":"J Castelli-Gair","year":"1995","unstructured":"Castelli-Gair J, Akam M (1995) How the Hox gene Ultrabithorax specifies two different segments: the significance of spatial and temporal regulation within metameres. Development 121(9):2973\u20132982","journal-title":"Development"},{"issue":"18","key":"707_CR9","doi-asserted-by":"publisher","first-page":"dev170480","DOI":"10.1242\/dev.170480","volume":"146","author":"E Clark","year":"2019","unstructured":"Clark E, Peel AD, Akam M (2019) Arthropod segmentation. Development. 146(18):dev170480","journal-title":"Development."},{"issue":"6214","key":"707_CR10","doi-asserted-by":"publisher","first-page":"432","DOI":"10.1038\/338432a0","volume":"338","author":"SM Cohen","year":"1989","unstructured":"Cohen SM, Br\u00f6nner G, K\u00fcttner F, J\u00fcrgens G, J\u00e4ckle H (1989) Distal-less encodes a homoeodomain protein required for limb development in Drosophila. Nature 338(6214):432\u2013434","journal-title":"Nature"},{"issue":"10","key":"707_CR11","doi-asserted-by":"publisher","first-page":"759","DOI":"10.1016\/S0960-9822(01)00222-6","volume":"11","author":"CE Cook","year":"2001","unstructured":"Cook CE, Smith ML, Telford MJ, Bastianello A, Akam M (2001) Hox genes and the phylogeny of the arthropods. Curr Biol 11(10):759\u2013763","journal-title":"Curr Biol"},{"issue":"24","key":"707_CR12","doi-asserted-by":"publisher","first-page":"5915","DOI":"10.1242\/dev.00835","volume":"130","author":"T Copf","year":"2003","unstructured":"Copf T, Rabet N, Celniker SE, Averof M (2003) Posterior patterning genes and the identification of a unique body region in the brine shrimp Artemia franciscana. Development 130(24):5915\u20135927","journal-title":"Development"},{"issue":"11","key":"707_CR13","doi-asserted-by":"publisher","first-page":"992","DOI":"10.1002\/bies.10180","volume":"24","author":"WG Damen","year":"2002","unstructured":"Damen WG (2002) fushi tarazu: a Hox gene changes its role. Bioessays 24(11):992\u2013995","journal-title":"Bioessays"},{"issue":"18","key":"707_CR14","doi-asserted-by":"publisher","first-page":"10665","DOI":"10.1073\/pnas.95.18.10665","volume":"95","author":"WG Damen","year":"1998","unstructured":"Damen WG, Hausdorf M, Seyfarth EA, Tautz D (1998) A conserved mode of head segmentation in arthropods revealed by the expression pattern of Hox genes in a spider. Proc Natl Acad Sci U S A 95(18):10665\u201310670","journal-title":"Proc Natl Acad Sci U S A"},{"issue":"1","key":"707_CR15","doi-asserted-by":"publisher","first-page":"85","DOI":"10.1002\/(SICI)1097-010X(19990415)285:1<85::AID-JEZ10>3.0.CO;2-N","volume":"285","author":"WG Damen","year":"1999","unstructured":"Damen WG, Tautz D (1999) Abdominal-B expression in a spider suggests a general role for Abdominal-B in specifying the genital structure. J Exp Zool 285(1):85\u201391","journal-title":"J Exp Zool"},{"issue":"2","key":"707_CR16","doi-asserted-by":"publisher","first-page":"323","DOI":"10.1242\/dev.108.2.323","volume":"108","author":"M Delorenzi","year":"1990","unstructured":"Delorenzi M, Bienz M (1990) Expression of Abdominal-B homeoproteins in Drosophila embryos. Development 108(2):323\u2013329","journal-title":"Development"},{"issue":"9","key":"707_CR17","doi-asserted-by":"publisher","first-page":"878","DOI":"10.1002\/bies.10319","volume":"25","author":"JS Deutsch","year":"2003","unstructured":"Deutsch JS, Mouchel-Vielh E (2003) Hox genes and the crustacean body plan. Bioessays 25(9):878\u2013887","journal-title":"Bioessays"},{"issue":"3-4","key":"707_CR18","doi-asserted-by":"publisher","first-page":"117","DOI":"10.1007\/s00427-010-0329-1","volume":"220","author":"BJ Eriksson","year":"2010","unstructured":"Eriksson BJ, Tait NN, Budd GE, Janssen R, Akam M (2010) Head patterning and Hox gene expression in an onychophoran and its implications for the arthropod head problem. Dev Genes Evol 220(3-4):117\u2013122","journal-title":"Dev Genes Evol"},{"key":"707_CR19","first-page":"17","volume-title":"Biology of spiders","author":"R Foelix","year":"2010","unstructured":"Foelix R (2010) Biology of spiders, 3rd edn. Oxford University Press, Oxford, pp 17\u201348","edition":"3"},{"issue":"1956","key":"707_CR20","first-page":"20211168","volume":"288","author":"G Gainett","year":"2021","unstructured":"Gainett G, Gonz\u00e1lez VL, Ballesteros JA, Setton EVW, Baker CM, Barolo Gargiulo L, Santib\u00e1\u00f1ez-L\u00f3pez CE, Coddington JA, Sharma PP (2021) The genome of a daddy-long-legs (Opiliones) illuminates the evolution of arachnid appendages. Proc Biol Sci 288(1956):20211168","journal-title":"Proc Biol Sci"},{"key":"707_CR21","doi-asserted-by":"publisher","first-page":"e1719","DOI":"10.7717\/peerj.1719","volume":"4","author":"NL Garrison","year":"2016","unstructured":"Garrison NL, Rodriguez J, Agnarsson I, Coddington JA, Griswold CE, Hamilton CA, Hedin M, Kocot KM, Ledford JM, Bond JE (2016) Spider phylogenomics: untangling the spider tree of life. PeerJ 4:e1719","journal-title":"PeerJ"},{"issue":"11-12","key":"707_CR22","doi-asserted-by":"publisher","first-page":"673","DOI":"10.1387\/ijdb.180162sg","volume":"62","author":"SJ Gaunt","year":"2018","unstructured":"Gaunt SJ (2018) Hox cluster genes and collinearities throughout the tree of animal life. Int J Dev Biol 62(11-12):673\u2013683","journal-title":"Int J Dev Biol"},{"issue":"4","key":"707_CR23","doi-asserted-by":"publisher","first-page":"657","DOI":"10.1242\/dev.102.4.657","volume":"102","author":"G Gibson","year":"1988","unstructured":"Gibson G, Gehring WJ (1988) Head and thoracic transformations caused by ectopic expression of Antennapedia during Drosophila development. Development 102(4):657\u2013675","journal-title":"Development"},{"issue":"8","key":"707_CR24","doi-asserted-by":"publisher","first-page":"547","DOI":"10.1016\/S0960-9822(06)00253-3","volume":"7","author":"JK Grenier","year":"1997","unstructured":"Grenier JK, Garber TL, Warren R, Whitington PM, Carroll S (1997) Evolution of the entire arthropod Hox gene set predated the origin and radiation of the onychophoran\/arthropod clade. Curr Biol 7(8):547\u2013553","journal-title":"Curr Biol"},{"key":"707_CR25","doi-asserted-by":"publisher","unstructured":"Harper A, Baudouin Gonzalez L, Sch\u00f6nauer A, Janssen R, Seiter M, Holzem M, Arif S, McGregor AP, Sumner-Rooney L (2021) Widespread retention of ohnologs in key developmental gene families following whole-genome duplication in arachnopulmonates. G3:jkab299. https:\/\/doi.org\/10.1093\/g3journal\/jkab299","DOI":"10.1093\/g3journal\/jkab299"},{"issue":"15","key":"707_CR26","doi-asserted-by":"publisher","first-page":"2655","DOI":"10.1242\/dev.078204","volume":"139","author":"M Hilbrant","year":"2012","unstructured":"Hilbrant M, Damen WG, McGregor AP (2012) Evolutionary crossroads in developmental biology: the spider Parasteatoda tepidariorum. Development 139(15):2655\u20132662","journal-title":"Development"},{"issue":"6","key":"707_CR27","doi-asserted-by":"publisher","first-page":"494","DOI":"10.1111\/gtc.12589","volume":"23","author":"C Hiruta","year":"2018","unstructured":"Hiruta C, Kakui K, Tollefsen KE, Iguchi T (2018) Targeted gene disruption by use of CRISPR\/Cas9 ribonucleoprotein complexes in the water flea Daphnia pulex. Genes Cells 23(6):494\u2013502","journal-title":"Genes Cells"},{"issue":"2","key":"707_CR28","doi-asserted-by":"publisher","first-page":"131","DOI":"10.1111\/j.1525-142X.2010.00399.x","volume":"12","author":"CC Hsia","year":"2010","unstructured":"Hsia CC, Par\u00e9 AC, Hannon M, Ronshaugen M, McGinnis W (2010) Silencing of an abdominal Hox gene during early development is correlated with limb development in a crustacean trunk. Evol Dev 12(2):131\u2013143","journal-title":"Evol Dev"},{"key":"707_CR29","doi-asserted-by":"publisher","first-page":"754","DOI":"10.1093\/bioinformatics\/17.8.754","volume":"17","author":"JP Huelsenbeck","year":"2001","unstructured":"Huelsenbeck JP, Ronquist F (2001) MRBAYES: bayesian inference of phylogenetic trees. Bioinformatics 17:754\u2013755","journal-title":"Bioinformatics"},{"issue":"17","key":"707_CR30","doi-asserted-by":"publisher","first-page":"3683","DOI":"10.1242\/dev.127.17.3683","volume":"127","author":"CL Hughes","year":"2000","unstructured":"Hughes CL, Kaufman TC (2000) RNAi analysis of deformed, proboscipedia and sex combs reduced in the milkweed bug Oncopeltus fasciatus: novel roles for Hox genes in the hemipteran head. Development 127(17):3683\u20133694","journal-title":"Development"},{"issue":"6","key":"707_CR31","doi-asserted-by":"publisher","first-page":"459","DOI":"10.1046\/j.1525-142X.2002.02034.x","volume":"4","author":"CL Hughes","year":"2002","unstructured":"Hughes CL, Kaufman TC (2002a) Hox genes and the evolution of the arthropod body plan. Evol Dev. 4(6):459\u2013499","journal-title":"Evol Dev."},{"issue":"5","key":"707_CR32","doi-asserted-by":"publisher","first-page":"1225","DOI":"10.1242\/dev.129.5.1225","volume":"129","author":"CL Hughes","year":"2002","unstructured":"Hughes CL, Kaufman TC (2002b) Exploring the myriapod body plan: expression patterns of the ten Hox genes in a centipede. Development 129(5):1225\u20131238","journal-title":"Development"},{"issue":"1","key":"707_CR33","doi-asserted-by":"publisher","first-page":"88","DOI":"10.1186\/s12862-018-1196-z","volume":"18","author":"R Janssen","year":"2018","unstructured":"Janssen R, Andersson E, Betn\u00e9r E, Bijl S, Fowler W, H\u00f6\u00f6k L, Leyhr J, Mannelqvist A, Panara V, Smith K, Tiemann S (2018) Embryonic expression patterns and phylogenetic analysis of panarthropod sox genes: insight into nervous system development, segmentation and gonadogenesis. BMC Evol Biol 18(1):88","journal-title":"BMC Evol Biol"},{"issue":"7-8","key":"707_CR34","doi-asserted-by":"publisher","first-page":"451","DOI":"10.1007\/s00427-006-0092-5","volume":"216","author":"R Janssen","year":"2006","unstructured":"Janssen R, Damen WG (2006) The ten Hox genes of the millipede Glomeris marginata. Dev Genes Evol 216(7-8):451\u2013465","journal-title":"Dev Genes Evol"},{"issue":"2","key":"707_CR35","doi-asserted-by":"publisher","first-page":"155","DOI":"10.1111\/j.1525-142X.2008.00223.x","volume":"10","author":"R Janssen","year":"2008","unstructured":"Janssen R, Damen WG (2008) Diverged and conserved aspects of heart formation in a spider. Evol Dev. 10(2):155\u2013165","journal-title":"Evol Dev."},{"issue":"1","key":"707_CR36","doi-asserted-by":"publisher","first-page":"22","DOI":"10.1186\/1742-9994-11-22","volume":"11","author":"R Janssen","year":"2014","unstructured":"Janssen R, Eriksson BJ, Tait NN, Budd GE (2014) Onychophoran Hox genes and the evolution of arthropod Hox gene expression. Front Zool 11(1):22","journal-title":"Front Zool"},{"key":"707_CR37","doi-asserted-by":"publisher","first-page":"96","DOI":"10.3389\/fevo.2015.00096","volume":"3","author":"R Janssen","year":"2015","unstructured":"Janssen R, Sch\u00f6nauer A, Weber M, Turetzek N, Hogvall M, Goss GE, Patel NH, McGregor AP, Hilbrant M (2015) The evolution and expression of panarthropod frizzled genes. Front Ecol Evol 3:96","journal-title":"Front Ecol Evol"},{"issue":"3","key":"707_CR38","doi-asserted-by":"publisher","first-page":"533","DOI":"10.1093\/icb\/icx063","volume":"57","author":"EL Jockusch","year":"2017","unstructured":"Jockusch EL (2017) Developmental and evolutionary perspectives on the origin and diversification of arthropod appendages. Integr Comp Biol 57(3):533\u2013545","journal-title":"Integr Comp Biol"},{"issue":"1","key":"707_CR39","doi-asserted-by":"publisher","first-page":"293","DOI":"10.1242\/dev.117.1.293","volume":"117","author":"R Kelsh","year":"1993","unstructured":"Kelsh R, Dawson I, Akam M (1993) An analysis of Abdominal-B expression in the locust Schistocerca gregaria. Development 117(1):293\u2013305","journal-title":"Development"},{"issue":"13","key":"707_CR40","doi-asserted-by":"publisher","first-page":"4921","DOI":"10.1073\/pnas.1116421109","volume":"109","author":"S Khadjeh","year":"2012","unstructured":"Khadjeh S, Turetzek N, Pechmann M, Schwager EE, Wimmer EA, Damen WG, Prpic NM (2012) Divergent role of the Hox gene Antennapedia in spiders is responsible for the convergent evolution of abdominal limb repression. Proc Natl Acad Sci U S A 109(13):4921\u20134926","journal-title":"Proc Natl Acad Sci U S A"},{"key":"707_CR41","doi-asserted-by":"publisher","first-page":"311","DOI":"10.1007\/978-3-642-59828-9_19","volume-title":"Development, Genetics, Epigenetics and Environmental Regulation","author":"M Klingler","year":"1999","unstructured":"Klingler M, Tautz D (1999) Formation of embryonic axes and blastoderm pattern in Drosophila. In: Russo VEA, Cove DJ, Edgar LG, Jaenisch R, Salamini F (eds) Development, Genetics, Epigenetics and Environmental Regulation. Springer, Berlin, pp 311\u2013330"},{"issue":"6","key":"707_CR42","doi-asserted-by":"publisher","first-page":"389","DOI":"10.1007\/s00427-017-0595-2","volume":"227","author":"T K\u00f6nigsmann","year":"2017","unstructured":"K\u00f6nigsmann T, Turetzek N, Pechmann M, Prpic NM (2017) Expression and function of the zinc finger transcription factor Sp6-9 in the spider Parasteatoda tepidariorum. Dev Genes Evol 227(6):389\u2013400","journal-title":"Dev Genes Evol"},{"issue":"1","key":"707_CR43","doi-asserted-by":"publisher","first-page":"2","DOI":"10.1186\/2041-9139-5-2","volume":"5","author":"B Konopova","year":"2014","unstructured":"Konopova B, Akam M (2014) The Hox genes Ultrabithorax and abdominal-A specify three different types of abdominal appendage in the springtail Orchesella cincta (Collembola). Evodevo 5(1):2","journal-title":"Evodevo"},{"issue":"11","key":"707_CR44","doi-asserted-by":"publisher","first-page":"2037","DOI":"10.1002\/j.1460-2075.1983.tb01697.x","volume":"2","author":"M Levine","year":"1983","unstructured":"Levine M, Hafen E, Garber RL, Gehring WJ (1983) Spatial distribution of Antennapedia transcripts during Drosophila development. EMBO J 2(11):2037\u20132046","journal-title":"EMBO J"},{"issue":"5688","key":"707_CR45","doi-asserted-by":"publisher","first-page":"565","DOI":"10.1038\/276565a0","volume":"276","author":"EB Lewis","year":"1978","unstructured":"Lewis EB (1978) A gene complex controlling segmentation in Drosophila. Nature 276(5688):565\u2013570","journal-title":"Nature"},{"issue":"Pt A","key":"707_CR46","first-page":"269","volume":"85","author":"EB Lewis","year":"1982","unstructured":"Lewis EB (1982) Control of body segment differentiation in Drosophila by the bithorax gene complex. Prog Clin Biol Res 85(Pt A):269\u2013288","journal-title":"Prog Clin Biol Res"},{"issue":"33","key":"707_CR47","doi-asserted-by":"publisher","first-page":"13892","DOI":"10.1073\/pnas.0903105106","volume":"106","author":"DM Liubicich","year":"2009","unstructured":"Liubicich DM, Serano JM, Pavlopoulos A, Kontarakis Z, Protas ME, Kwan E, Chatterjee S, Tran KD, Averof M, Patel NH (2009) Knockdown of Parhyale Ultrabithorax recapitulates evolutionary changes in crustacean appendage morphology. Proc Natl Acad Sci U S A 106(33):13892\u201313896","journal-title":"Proc Natl Acad Sci U S A"},{"issue":"9","key":"707_CR48","doi-asserted-by":"publisher","first-page":"e866","DOI":"10.1371\/journal.pone.0000866","volume":"2","author":"N Mahfooz","year":"2007","unstructured":"Mahfooz N, Turchyn N, Mihajlovic M, Hrycaj S, Popadi\u0107 A (2007) Ubx regulates differential enlargement and diversification of insect hind legs. PLoS One 2(9):e866","journal-title":"PLoS One"},{"key":"707_CR49","doi-asserted-by":"publisher","first-page":"109","DOI":"10.3389\/fevo.2020.00109","volume":"8","author":"P Mariano-Martins","year":"2020","unstructured":"Mariano-Martins P, Lo-Man-Hung N, Torres TT (2020) Evolution of spiders and silk spinning: mini review of the morphology, evolution, and development of spiders\u2019 spinnerets. Front. Ecol. Evol 8:109","journal-title":"Front. Ecol. Evol"},{"issue":"1","key":"707_CR50","doi-asserted-by":"publisher","first-page":"14","DOI":"10.1016\/j.cub.2015.11.021","volume":"26","author":"A Martin","year":"2016","unstructured":"Martin A, Serano JM, Jarvis E, Bruce HS, Wang J, Ray S, Barker CA, O'Connell LC, Patel NH (2016) CRISPR\/Cas9 mutagenesis reveals versatile roles of Hox genes in crustacean limb specification and evolution. Curr Biol 26(1):14\u201326","journal-title":"Curr Biol"},{"issue":"5","key":"707_CR51","doi-asserted-by":"publisher","first-page":"487","DOI":"10.1002\/bies.20744","volume":"30","author":"AP McGregor","year":"2008","unstructured":"McGregor AP, Hilbrant M, Pechmann M, Schwager EE, Prpic NM, Damen WG (2008) Cupiennius salei and Achaearanea tepidariorum: spider models for investigating evolution and development. Bioessays 30(5):487\u2013498","journal-title":"Bioessays"},{"key":"707_CR52","first-page":"82","volume":"188","author":"J Mendoza","year":"2020","unstructured":"Mendoza J, Francke O (2020) Systematic revision of Mexican threatened tarantulas Brachypelma (Araneae: Theraphosidae: Theraphosinae), with a description of a new genus, and implications on the conservation. Zool J Linnean Soc 188:82\u2013147","journal-title":"Zool J Linnean Soc"},{"issue":"5785","key":"707_CR53","doi-asserted-by":"publisher","first-page":"795","DOI":"10.1038\/287795a0","volume":"287","author":"C N\u00fcsslein-Volhard","year":"1980","unstructured":"N\u00fcsslein-Volhard C, Wieschaus E (1980) Mutations affecting segment number and polarity in Drosophila. Nature 287(5785):795\u2013801","journal-title":"Nature"},{"key":"707_CR54","doi-asserted-by":"publisher","first-page":"6","DOI":"10.1186\/s13227-020-00152-z","volume":"11","author":"H Oda","year":"2020","unstructured":"Oda H, Akiyama-Oda Y (2020) The common house spider Parasteatoda tepidariorum. Evodevo 11:6","journal-title":"Evodevo"},{"issue":"3","key":"707_CR55","doi-asserted-by":"publisher","first-page":"354","DOI":"10.1016\/j.asd.2016.10.011","volume":"46","author":"J Ortega-Hern\u00e1ndez","year":"2017","unstructured":"Ortega-Hern\u00e1ndez J, Janssen R, Budd GE (2017) Origin and evolution of the panarthropod head - a palaeobiological and developmental perspective. Arthropod Struct Dev 46(3):354\u2013379","journal-title":"Arthropod Struct Dev"},{"key":"707_CR56","doi-asserted-by":"publisher","first-page":"23","DOI":"10.1186\/s12983-019-0322-0","volume":"16","author":"V Panara","year":"2019","unstructured":"Panara V, Budd GE, Janssen R (2019) Phylogenetic analysis and embryonic expression of panarthropod Dmrt genes. Front Zool 16:23","journal-title":"Front Zool"},{"issue":"33","key":"707_CR57","doi-asserted-by":"publisher","first-page":"13897","DOI":"10.1073\/pnas.0902804106","volume":"106","author":"A Pavlopoulos","year":"2009","unstructured":"Pavlopoulos A, Kontarakis Z, Liubicich DM, Serano JM, Akam M, Patel NH, Averof M (2009) Probing the evolution of appendage specialization by Hox gene misexpression in an emerging model crustacean. Proc Natl Acad Sci U S A 106(33):13897\u201313902","journal-title":"Proc Natl Acad Sci U S A"},{"issue":"2","key":"707_CR58","doi-asserted-by":"publisher","first-page":"75","DOI":"10.1007\/s00427-020-00653-w","volume":"230","author":"M Pechmann","year":"2020","unstructured":"Pechmann M (2020) Embryonic development and secondary axis induction in the Brazilian white knee tarantula Acanthoscurria geniculata, C. L. Koch, 1841 (Araneae; Mygalomorphae; Theraphosidae). Dev Genes Evol 230(2):75\u201394","journal-title":"Dev Genes Evol"},{"issue":"6","key":"707_CR59","doi-asserted-by":"publisher","first-page":"453","DOI":"10.1016\/j.asd.2010.07.007","volume":"39","author":"M Pechmann","year":"2010","unstructured":"Pechmann M, Khadjeh S, Sprenger F, Prpic NM (2010) Patterning mechanisms and morphological diversity of spider appendages and their importance for spider evolution. Arthropod Struct Dev 39(6):453\u2013467","journal-title":"Arthropod Struct Dev"},{"issue":"4","key":"707_CR60","doi-asserted-by":"publisher","first-page":"189","DOI":"10.1007\/s00427-009-0279-7","volume":"219","author":"M Pechmann","year":"2009","unstructured":"Pechmann M, Prpic NM (2009) Appendage patterning in the South American bird spider Acanthoscurria geniculata (Araneae: Mygalomorphae). Dev Genes Evol 219(4):189\u2013198","journal-title":"Dev Genes Evol"},{"issue":"3","key":"707_CR61","doi-asserted-by":"publisher","first-page":"535","DOI":"10.1007\/s00412-015-0553-6","volume":"125","author":"L Pick","year":"2016","unstructured":"Pick L (2016) Hox genes, evo-devo, and the case of the ftz gene. Chromosoma 125(3):535\u2013551","journal-title":"Chromosoma"},{"issue":"1865","key":"707_CR62","doi-asserted-by":"publisher","first-page":"20210269","DOI":"10.1098\/rstb.2021.0269","volume":"377","author":"NM Prpic","year":"2022","unstructured":"Prpic NM, Pechmann M (2022) Extraembryonic tissue in chelicerates: a review and outlook. Philos Trans R Soc Lond B Biol Sci 377(1865):20210269","journal-title":"Philos Trans R Soc Lond B Biol Sci"},{"key":"707_CR63","doi-asserted-by":"publisher","first-page":"pdb.prot5067","DOI":"10.1101\/pdb.prot5067","volume":"2008","author":"NM Prpic","year":"2008","unstructured":"Prpic NM, Schoppmeier M, Damen WG (2008) Collection and fixation of spider embryos. CSH Protoc 2008:pdb.prot5067. https:\/\/doi.org\/10.1101\/pdb.prot5067","journal-title":"CSH Protoc"},{"issue":"7","key":"707_CR64","doi-asserted-by":"publisher","first-page":"901","DOI":"10.1101\/gad.2.7.901","volume":"2","author":"MA Pultz","year":"1988","unstructured":"Pultz MA, Diederich RJ, Cribbs DL, Kaufman TC (1988) The proboscipedia locus of the Antennapedia complex: a molecular and genetic analysis. Genes Dev 2(7):901\u2013920","journal-title":"Genes Dev"},{"issue":"2","key":"707_CR65","doi-asserted-by":"publisher","first-page":"441","DOI":"10.1016\/j.ydbio.2014.05.021","volume":"392","author":"PN Refki","year":"2014","unstructured":"Refki PN, Armis\u00e9n D, Crumi\u00e8re AJ, Viala S, Khila A (2014) Emergence of tissue sensitivity to Hox protein levels underlies the evolution of an adaptive morphological trait. Dev Biol 392(2):441\u2013453","journal-title":"Dev Biol"},{"key":"707_CR66","first-page":"1","volume-title":"Insect Development","author":"LK Robertson","year":"2009","unstructured":"Robertson LK, Mahaffey JW (2009) Insect homeotic complex genes and development, lessons from Drosophila and beyond. In: Gilbert LI (ed) Insect Development. Elsevier, San Diego, pp 1\u201357"},{"key":"707_CR67","doi-asserted-by":"publisher","first-page":"16","DOI":"10.1186\/s13227-015-0010-x","volume":"6","author":"L Samadi","year":"2015","unstructured":"Samadi L, Schmid A, Eriksson BJ (2015) Differential expression of retinal determination genes in the principal and secondary eyes of Cupiennius salei Keyserling (1877). Evodevo. 6:16","journal-title":"Evodevo."},{"issue":"5998","key":"707_CR68","doi-asserted-by":"publisher","first-page":"108","DOI":"10.1038\/313108a0","volume":"313","author":"E S\u00e1nchez-Herrero","year":"1985","unstructured":"S\u00e1nchez-Herrero E, Vern\u00f3s I, Marco R, Morata G (1985) Genetic organization of Drosophila bithorax complex. Nature 313(5998):108\u2013113","journal-title":"Nature"},{"issue":"2","key":"707_CR69","doi-asserted-by":"publisher","first-page":"76","DOI":"10.1007\/s004270000121","volume":"211","author":"M Schoppmeier","year":"2001","unstructured":"Schoppmeier M, Damen WG (2001) Double-stranded RNA interference in the spider Cupiennius salei: the role of Distal-less is evolutionarily conserved in arthropod appendage formation. Dev Genes Evol 211(2):76\u201382","journal-title":"Dev Genes Evol"},{"key":"707_CR70","doi-asserted-by":"publisher","first-page":"10","DOI":"10.1186\/1742-9994-4-10","volume":"4","author":"EE Schwager","year":"2007","unstructured":"Schwager EE, Schoppmeier M, Pechmann M, Damen WG (2007) Duplicated Hox genes in the spider Cupiennius salei. Front Zool 4:10","journal-title":"Front Zool"},{"issue":"1","key":"707_CR71","doi-asserted-by":"publisher","first-page":"62","DOI":"10.1186\/s12915-017-0399-x","volume":"15","author":"EE Schwager","year":"2017","unstructured":"Schwager EE, Sharma PP, Clarke T, Leite DJ, Wierschin T, Pechmann M, Akiyama-Oda Y, Esposito L, Bechsgaard J, Bilde T, Buffry AD, Chao H, Dinh H, Doddapaneni H, Dugan S, Eibner C, Extavour CG, Funch P, Garb J et al (2017) The house spider genome reveals an ancient whole-genome duplication during arachnid evolution. BMC Biol 15(1):62","journal-title":"BMC Biol"},{"issue":"1","key":"707_CR72","doi-asserted-by":"publisher","first-page":"297","DOI":"10.1016\/j.ydbio.2015.10.029","volume":"409","author":"JM Serano","year":"2016","unstructured":"Serano JM, Martin A, Liubicich DM, Jarvis E, Bruce HS, La K, Browne WE, Grimwood J, Patel NH (2016) Comprehensive analysis of Hox gene expression in the amphipod crustacean Parhyale hawaiensis. Dev Biol 409(1):297\u2013309","journal-title":"Dev Biol"},{"issue":"3","key":"707_CR73","doi-asserted-by":"publisher","first-page":"510","DOI":"10.1093\/icb\/icx078","volume":"57","author":"PP Sharma","year":"2017","unstructured":"Sharma PP (2017) Chelicerates and the conquest of land: a view of arachnid origins through an Evo-Devo spyglass. Integr Comp Biol 57(3):510\u2013522","journal-title":"Integr Comp Biol"},{"issue":"11","key":"707_CR74","doi-asserted-by":"publisher","first-page":"2963","DOI":"10.1093\/molbev\/msu235","volume":"31","author":"PP Sharma","year":"2014","unstructured":"Sharma PP, Kaluziak ST, P\u00e9rez-Porro AR, Gonz\u00e1lez VL, Hormiga G, Wheeler WC, Giribet G (2014b) Phylogenomic interrogation of arachnida reveals systemic conflicts in phylogenetic signal. Mol Biol Evol 31(11):2963\u20132984","journal-title":"Mol Biol Evol"},{"issue":"5","key":"707_CR75","doi-asserted-by":"publisher","first-page":"450","DOI":"10.1111\/j.1525-142X.2012.00565.x","volume":"14","author":"PP Sharma","year":"2012","unstructured":"Sharma PP, Schwager EE, Extavour CG, Giribet G (2012) Hox gene expression in the harvestman Phalangium opilio reveals divergent patterning of the chelicerate opisthosoma. Evol Dev 14(5):450\u2013463","journal-title":"Evol Dev"},{"issue":"1792","key":"707_CR76","first-page":"20140661","volume":"281","author":"PP Sharma","year":"2014","unstructured":"Sharma PP, Schwager EE, Extavour CG, Wheeler WC (2014a) Hox gene duplications correlate with posterior heteronomy in scorpions. Proc Biol Sci 281(1792):20140661","journal-title":"Proc Biol Sci"},{"issue":"4","key":"707_CR77","doi-asserted-by":"publisher","first-page":"228","DOI":"10.1111\/ede.12029","volume":"15","author":"PP Sharma","year":"2013","unstructured":"Sharma PP, Schwager EE, Giribet G, Jockusch EL, Extavour CG (2013) Distal-less and dachshund pattern both plesiomorphic and apomorphic structures in chelicerates: RNA interference in the harvestman Phalangium opilio (Opiliones). Evol Dev 15(4):228\u2013242","journal-title":"Evol Dev"},{"issue":"2","key":"707_CR78","doi-asserted-by":"publisher","first-page":"224","DOI":"10.1016\/j.cub.2015.11.059","volume":"26","author":"FW Smith","year":"2016","unstructured":"Smith FW, Boothby TC, Giovannini I, Rebecchi L, Jockusch EL, Goldstein B (2016) The compact body plan of tardigrades evolved by the loss of a large body region. Curr Biol 26(2):224\u2013229","journal-title":"Curr Biol"},{"key":"707_CR79","doi-asserted-by":"publisher","first-page":"615","DOI":"10.1146\/annurev.genet.38.072902.092831","volume":"38","author":"JS Taylor","year":"2004","unstructured":"Taylor JS, Raes J (2004) Duplication and divergence: the evolution of new genes and old ideas. Annu Rev Genet 38:615\u2013643","journal-title":"Annu Rev Genet"},{"issue":"18","key":"707_CR80","doi-asserted-by":"publisher","first-page":"10671","DOI":"10.1073\/pnas.95.18.10671","volume":"95","author":"MJ Telford","year":"1998","unstructured":"Telford MJ, Thomas RH (1998) Expression of homeobox genes shows chelicerate arthropods retain their deutocerebral segment. Proc Natl Acad Sci U S A 95(18):10671\u201310675","journal-title":"Proc Natl Acad Sci U S A"},{"key":"707_CR81","volume-title":"On the role of Hox genes and co-factors in body plan specification in spiders","author":"N Turetzek","year":"2011","unstructured":"Turetzek N (2011) On the role of Hox genes and co-factors in body plan specification in spiders. Master Thesis. Georg-August University, G\u00f6ttingen, Germany"},{"key":"707_CR82","doi-asserted-by":"publisher","unstructured":"Turetzek N, Pechmann M, Janssen R, Prpic NM (2022) Hox genes in spiders: their significance for development and evolution. Semin Cell Dev Biol. https:\/\/doi.org\/10.1016\/j.semcdb.2022.11.014","DOI":"10.1016\/j.semcdb.2022.11.014"},{"issue":"6","key":"707_CR83","doi-asserted-by":"publisher","first-page":"413","DOI":"10.1007\/s00427-016-0562-3","volume":"226","author":"N Turetzek","year":"2016","unstructured":"Turetzek N, Prpic NM (2016) Observations on germ band development in the cellar spider Pholcus phalangioides. Dev Genes Evol 226(6):413\u2013422","journal-title":"Dev Genes Evol"},{"issue":"3","key":"707_CR84","doi-asserted-by":"publisher","first-page":"437","DOI":"10.1016\/0092-8674(92)90513-C","volume":"71","author":"G Vachon","year":"1992","unstructured":"Vachon G, Cohen B, Pfeifle C, McGuffin ME, Botas J, Cohen SM (1992) Homeotic genes of the Bithorax complex repress limb development in the abdomen of the Drosophila embryo through the target gene Distal-less. Cell 71(3):437\u2013450","journal-title":"Cell"},{"issue":"1","key":"707_CR85","doi-asserted-by":"publisher","first-page":"15","DOI":"10.1186\/1742-9994-8-15","volume":"8","author":"C Wolff","year":"2011","unstructured":"Wolff C, Hilbrant M (2011) The embryonic development of the central American wandering spider Cupiennius salei. Front Zool 8(1):15","journal-title":"Front Zool"},{"issue":"1","key":"707_CR86","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1002\/arch.20393","volume":"76","author":"H Xiang","year":"2011","unstructured":"Xiang H, Li MW, Guo JH, Jiang JH, Huang YP (2011) Influence of RNAi knockdown for E-complex genes on the silkworm proleg development. Arch Insect Biochem Physiol 76(1):1\u201311","journal-title":"Arch Insect Biochem Physiol"}],"container-title":["Development Genes and Evolution"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/link.springer.com\/content\/pdf\/10.1007\/s00427-023-00707-9.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/link.springer.com\/article\/10.1007\/s00427-023-00707-9\/fulltext.html","content-type":"text\/html","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/link.springer.com\/content\/pdf\/10.1007\/s00427-023-00707-9.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2023,12,22]],"date-time":"2023-12-22T14:04:39Z","timestamp":1703253879000},"score":1,"resource":{"primary":{"URL":"https:\/\/link.springer.com\/10.1007\/s00427-023-00707-9"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2023,7,27]]},"references-count":86,"journal-issue":{"issue":"2","published-print":{"date-parts":[[2023,12]]}},"alternative-id":["707"],"URL":"https:\/\/doi.org\/10.1007\/s00427-023-00707-9","relation":{},"ISSN":["0949-944X","1432-041X"],"issn-type":[{"value":"0949-944X","type":"print"},{"value":"1432-041X","type":"electronic"}],"subject":[],"published":{"date-parts":[[2023,7,27]]},"assertion":[{"value":"29 May 2023","order":1,"name":"received","label":"Received","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"11 July 2023","order":2,"name":"accepted","label":"Accepted","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"27 July 2023","order":3,"name":"first_online","label":"First Online","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"23 August 2023","order":4,"name":"change_date","label":"Change Date","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"Update","order":5,"name":"change_type","label":"Change Type","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"Editorial responsibility was changed from \"Responsible editor\" to \"Communicated by\"","order":6,"name":"change_details","label":"Change Details","group":{"name":"ArticleHistory","label":"Article History"}},{"order":1,"name":"Ethics","group":{"name":"EthicsHeading","label":"Declarations"}},{"value":"Not applicable.","order":2,"name":"Ethics","group":{"name":"EthicsHeading","label":"Ethics approval and consent to participate"}},{"value":"Not applicable.","order":3,"name":"Ethics","group":{"name":"EthicsHeading","label":"Consent for publication"}},{"value":"The authors declare no competing interests.","order":4,"name":"Ethics","group":{"name":"EthicsHeading","label":"Competing interests"}}]}}