{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,2,7]],"date-time":"2026-02-07T04:04:21Z","timestamp":1770437061287,"version":"3.49.0"},"reference-count":76,"publisher":"Wiley","issue":"7","license":[{"start":{"date-parts":[[2025,7,22]],"date-time":"2025-07-22T00:00:00Z","timestamp":1753142400000},"content-version":"vor","delay-in-days":21,"URL":"http:\/\/creativecommons.org\/licenses\/by\/4.0\/"},{"start":{"date-parts":[[2025,7,1]],"date-time":"2025-07-01T00:00:00Z","timestamp":1751328000000},"content-version":"tdm","delay-in-days":0,"URL":"http:\/\/doi.wiley.com\/10.1002\/tdm_license_1.1"}],"content-domain":{"domain":["onlinelibrary.wiley.com"],"crossmark-restriction":true},"short-container-title":["Journal of Morphology"],"published-print":{"date-parts":[[2025,7]]},"abstract":"<jats:title>ABSTRACT<\/jats:title>\n                  <jats:p>\n                    The study of ant morphology is advancing through parallel insights provided by phylogenomics\u2014which provides a statistically robust basis for comparison and evolutionary inference\u2014and phenomics via the application of microcomputed tomography (\u00b5\u2010CT) for the efficient and precise documentation of anatomy. The information provided by \u00b5\u2010CT is complex and rich, allowing for the quantification of geometry and biomechanically relevant variables, as well as comparative morphology via 3D rendering. Recently, the complete musculature of the thorax, propodeum, and legs was documented for the first time in an ant (\n                    <jats:italic>Formica rufa<\/jats:italic>\n                    L.). Here, we provide a detailed comparison of those findings for\n                    <jats:italic>Protanilla lini<\/jats:italic>\n                    Terayama, 2009 (Leptanillinae), representing the Leptanillomorpha, a clade putatively sister to all other living ants. Using \u00b5\u2010CT, computer\u2010based reconstruction, and scanning electron microscopy (SEM), we observe a novel series of morphological features that are plausibly part of the groundplan of the Formicidae. For several of these features, we provide new anatomical concepts and terms, with special discussion of the pronotum, promesothoracic articulation, and metapleural gland region. We also observe characters that are likely correlated with specialized subterranean habits, including the increased flexibility of the promesothoracic articulation, the slender shape of the mesosoma, a simplified vestiture of short setae, and depigmentation. Mesosomal skeletomusculature in\n                    <jats:italic>P. lini<\/jats:italic>\n                    appears to be plesiomorphic relative to other Leptanillomorpha, resembling the putative ancestral condition for the Formicidae. An exception lies in the lack of cervical muscle (Idlm1) in\n                    <jats:italic>Protanilla<\/jats:italic>\n                    that is present in almost all other insects for which this character has been sampled. With this study of\n                    <jats:italic>P. lini<\/jats:italic>\n                    , we are one step closer to realizing the complete set of defining features and variation of the ant mesosoma.\n                  <\/jats:p>","DOI":"10.1002\/jmor.70064","type":"journal-article","created":{"date-parts":[[2025,7,22]],"date-time":"2025-07-22T08:10:04Z","timestamp":1753171804000},"update-policy":"https:\/\/doi.org\/10.1002\/crossmark_policy","source":"Crossref","is-referenced-by-count":2,"title":["The Mesosoma of\n                    <i>Protanilla<\/i>\n                    (Leptanillinae) and the Groundplan of the Formicidae (Hymenoptera)"],"prefix":"10.1002","volume":"286","author":[{"given":"Lazzat","family":"Aibekova","sequence":"first","affiliation":[{"name":"Okinawa Institute of Science and Technology (OIST) Okinawa Japan"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Adrian","family":"Richter","sequence":"additional","affiliation":[{"name":"Okinawa Institute of Science and Technology (OIST) Okinawa Japan"},{"name":"Division Messel Research &amp; Mammalogy, Evolution of Sensory Systems Section Senckenberg Research Institute and Natural History Museum Frankfurt am Main Germany"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-0433-7626","authenticated-orcid":false,"given":"Rolf G.","family":"Beutel","sequence":"additional","affiliation":[{"name":"Institut f\u00fcr Zoologie und Evolutionsforschung FSU Jena Jena Germany"},{"name":"Division of Terrestrial Zoology, Section Entomology II Senckenberg Research Institute and Natural History Museum Frankfurt am Main Germany"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Thomas","family":"van de Kamp","sequence":"additional","affiliation":[{"name":"Institute for Photon Science and Synchrotron Radiation (IPS) Karlsruhe Institute of Technology (KIT) Eggenstein\u2010Leopoldshafen Germany"},{"name":"Laboratory for Applications of Synchrotron Radiation (LAS) Karlsruhe Institute of Technology (KIT) Karlsruhe Germany"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Evan P.","family":"Economo","sequence":"additional","affiliation":[{"name":"Okinawa Institute of Science and Technology (OIST) Okinawa Japan"},{"name":"Department of Entomology University of Maryland College Park Maryland USA"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Zachary","family":"Griebenow","sequence":"additional","affiliation":[{"name":"College of Agricultural Sciences, Agricultural Biology Colorado State University Fort Collins Colorado USA"},{"name":"Systematic Entomology Laboratory, United States Department of Agriculture Agricultural Research Service c\/o National Museum of Natural History Washington District of Columbia USA"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Brendon E.","family":"Boudinot","sequence":"additional","affiliation":[{"name":"Division of Terrestrial Zoology, Section Entomology II Senckenberg Research Institute and Natural History Museum Frankfurt am Main Germany"}],"role":[{"role":"author","vocabulary":"crossref"}]}],"member":"311","published-online":{"date-parts":[[2025,7,22]]},"reference":[{"key":"e_1_2_9_2_1","doi-asserted-by":"publisher","DOI":"10.1093\/isd\/ixac002"},{"key":"e_1_2_9_3_1","doi-asserted-by":"publisher","DOI":"10.1093\/iob\/obad026"},{"key":"e_1_2_9_4_1","doi-asserted-by":"publisher","DOI":"10.3897\/asp.64.e31640"},{"key":"e_1_2_9_5_1","volume-title":"Insect Morphology and Phylogeny","author":"Beutel R. 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E.Lieberman. in press. \u201cAnt Phylogeny Is Not Resolved by the Application of Site\u2010Heterogeneous Models.\u201dCommunications Biology."},{"key":"e_1_2_9_20_1","doi-asserted-by":"publisher","DOI":"10.1093\/zoolinnean\/zlab097"},{"key":"e_1_2_9_21_1","doi-asserted-by":"publisher","DOI":"10.1002\/jmor.21757"},{"key":"e_1_2_9_22_1","doi-asserted-by":"publisher","DOI":"10.1590\/S0031-10492010002600001"},{"key":"e_1_2_9_23_1","doi-asserted-by":"publisher","DOI":"10.1111\/2041-210X.12742"},{"key":"e_1_2_9_24_1","doi-asserted-by":"publisher","DOI":"10.1038\/s42003-024-05793-7"},{"key":"e_1_2_9_25_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.nima.2010.10.150"},{"key":"e_1_2_9_26_1","doi-asserted-by":"publisher","DOI":"10.1071\/IS12063"},{"key":"e_1_2_9_27_1","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.1308650111"},{"key":"e_1_2_9_28_1","doi-asserted-by":"publisher","DOI":"10.1088\/1748-0221\/7\/09\/P09016"},{"key":"e_1_2_9_29_1","first-page":"3","volume-title":"A Contribution to the Biology of North American Vespine Wasps","author":"Duncan D.","year":"1939"},{"key":"e_1_2_9_30_1","first-page":"103","article-title":"Intorno al Torace delle Formiche e Particolarmente dei Neutri [About the Thorax of Ants and Especially of Neuters]","volume":"32","author":"Emery C.","year":"1900","journal-title":"Bollettino della Societa Entomologica Italiana"},{"key":"e_1_2_9_31_1","doi-asserted-by":"publisher","DOI":"10.1107\/S160057752200282X"},{"key":"e_1_2_9_32_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.asd.2007.04.003"},{"key":"e_1_2_9_33_1","first-page":"229","article-title":"Delimitation of Tribes in the Subfamily Leptanillinae (Hymenoptera: Formicidae), With a Description of the Male of Protanilla lini Terayama, 2009","volume":"30","author":"Griebenow Z.","year":"2020","journal-title":"Myrmecological News"},{"key":"e_1_2_9_34_1","doi-asserted-by":"publisher","DOI":"10.3897\/zookeys.1189.107506"},{"key":"e_1_2_9_35_1","doi-asserted-by":"publisher","DOI":"10.1071\/IS22035"},{"key":"e_1_2_9_36_1","doi-asserted-by":"publisher","DOI":"10.5852\/ejt.2025.987.2867"},{"key":"e_1_2_9_37_1","first-page":"945","article-title":"Comparative Morphology of Male Genital Skeletomusculature in the Leptanillinae (Hymenoptera: Formicidae), With a Standardized Muscular Terminology for the Male Genitalia of Hymenoptera","volume":"81","author":"Griebenow Z. 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