{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,7,28]],"date-time":"2026-07-28T20:21:26Z","timestamp":1785270086522,"version":"3.55.0"},"reference-count":143,"publisher":"Wiley","issue":"9","license":[{"start":{"date-parts":[[2024,8,27]],"date-time":"2024-08-27T00:00:00Z","timestamp":1724716800000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/"}],"content-domain":{"domain":["onlinelibrary.wiley.com"],"crossmark-restriction":true},"short-container-title":["Journal of Morphology"],"published-print":{"date-parts":[[2024,9]]},"abstract":"<jats:title>Abstract<\/jats:title>\n                  <jats:p>\n                    The male genitalia of insects are among the most variable, complex, and informative character systems for evolutionary analysis and taxonomic purposes. Because of these general properties, many generations of systematists have struggled to develop a theory of homology and alignment of parts. This struggle continues to the present day, where fundamentally different models and nomenclatures for the male genitalia of Hymenoptera, for example, are applied. Here, we take a multimodal approach to digitalize and comprehensively document the genital skeletomuscular anatomy of the bullet ant (\n                    <jats:italic>Paraponera clavata<\/jats:italic>\n                    ; Hymenoptera: Formicidae), including hand dissection, synchrotron radiation microcomputed tomography, microphotography, scanning electron microscopy,\u00a0confocal laser scanning microscopy,\u00a0and 3D\u2010printing. Through this work, we generate several new concepts for the structure and form of the male genitalia of Hymenoptera, such as for the endophallic sclerite (=fibula ducti), which we were able to evaluate in detail for the first time for any species. Based on this phenomic anatomical study and comparison with other Holometabola and Hexapoda, we reconsider the homologies of insect genitalia more broadly, and propose a series of clarifications in support of the penis\u2010gonopod theory of male genital identity. Specifically, we use the male genitalia of\n                    <jats:italic>Paraponera<\/jats:italic>\n                    and insects more broadly as an empirical case for hierarchical homology by applying and refining the 5\u2010category classification of serial homologs from DiFrisco\u00a0et al.\u00a0(2023) (DLW23) to all of our formalized concepts. Through this, we find that: (1) geometry is a critical attribute to account for in ontology, especially as all individually identifiable attributes are positionally indexed hence can be recognized as homomorphic; (2) the definition of \u201cstructure\u201d proposed by DLW23 is difficult to apply, and likely heterogeneous; and (3) formative elements, or spatially defined foldings or in\u2010 or evaginations of the epidermis and cuticle, are an important yet overlooked class of homomorphs. We propose a morphogenetic model for male and female insect genitalia, and a model analogous to gene\u2010tree species\u2010tree mappings for the hierarchical homology of male genitalia specifically. For all of the structures evaluated in the present study, we provide 3D\u2010printable models \u2013 with and without musculature, and in various states of digital dissection \u2013 to facilitate the development of a tactile understanding. Our treatment of the male genitalia of\n                    <jats:italic>P. clavata<\/jats:italic>\n                    serves as a basic template for future phenomic studies of male insect genitalia, which will be substantially improved with the development of automation and collections\u2010based data processing pipelines, that is,\u00a0collectomics. The Hymenoptera Anatomy Ontology will be a critical resource to include in this effort, and in best practice concepts should be linked.\n                  <\/jats:p>","DOI":"10.1002\/jmor.21757","type":"journal-article","created":{"date-parts":[[2024,8,28]],"date-time":"2024-08-28T00:38:42Z","timestamp":1724805522000},"update-policy":"https:\/\/doi.org\/10.1002\/crossmark_policy","source":"Crossref","is-referenced-by-count":8,"title":["Male genitalia, hierarchical homology, and the anatomy of the bullet ant (\n                    <i>Paraponera clavata<\/i>\n                    ; Hymenoptera, Formicidae)"],"prefix":"10.1002","volume":"285","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-4588-0430","authenticated-orcid":false,"given":"Brendon E.","family":"Boudinot","sequence":"first","affiliation":[{"name":"Department of Terrestrial Zoology, Entomology II Senckenberg Research Institute and Natural History Museum Frankfurt am Main Germany"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-7390-1318","authenticated-orcid":false,"given":"Thomas","family":"van de Kamp","sequence":"additional","affiliation":[{"name":"Institute for Photon Science and Synchrotron Radiation (IPS), Karlsruhe Institute of Technology (KIT) Karlsruhe Germany"},{"name":"Laboratory for Applications of Synchrotron Radiation (LAS) Karlsruhe Institute of Technology (KIT) Karlsruhe Germany"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Patricia","family":"Peters","sequence":"additional","affiliation":[{"name":"Department of Terrestrial Zoology, Entomology II Senckenberg Research Institute and Natural History Museum Frankfurt am Main Germany"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0003-4490-4300","authenticated-orcid":false,"given":"Katja","family":"Kn\u00f6llinger","sequence":"additional","affiliation":[{"name":"Department of Terrestrial Zoology, Entomology II Senckenberg Research Institute and Natural History Museum Frankfurt am Main Germany"},{"name":"Zurich University of the Arts Zurich Switzerland"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"311","published-online":{"date-parts":[[2024,8,27]]},"reference":[{"key":"e_1_2_10_2_1","doi-asserted-by":"publisher","DOI":"10.1093\/isd\/ixac002"},{"key":"e_1_2_10_3_1","doi-asserted-by":"publisher","DOI":"10.3390\/toxins12050324"},{"key":"e_1_2_10_4_1","doi-asserted-by":"publisher","DOI":"10.1016\/S0074-7696(08)62427-4"},{"key":"e_1_2_10_5_1","doi-asserted-by":"publisher","DOI":"10.1002\/jemt.23242"},{"key":"e_1_2_10_6_1","unstructured":"AntWeb. 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