{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,1]],"date-time":"2026-09-01T22:03:31Z","timestamp":1788300211885,"version":"build-2803163510"},"reference-count":48,"publisher":"Wiley","issue":"9","license":[{"start":{"date-parts":[[2020,7,18]],"date-time":"2020-07-18T00:00:00Z","timestamp":1595030400000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/onlinelibrary.wiley.com\/termsAndConditions#vor"}],"funder":[{"name":"AEI\/FEDER, UE","award":["CGL2013-48950-C2"],"award-info":[{"award-number":["CGL2013-48950-C2"]}]}],"content-domain":{"domain":["onlinelibrary.wiley.com"],"crossmark-restriction":true},"short-container-title":["Journal of Morphology"],"published-print":{"date-parts":[[2020,9]]},"abstract":"<jats:title>Abstract<\/jats:title><jats:p>Clavigeritae is a group of obligate myrmecophiles of the rove beetle subfamily Pselaphinae (Staphylinidae). Some are blind and wingless, and all are believed to depend on ant hosts through feeding by trophallaxis. Phylogenetic hypotheses suggest that their ancestors, as are most pselaphines today, were free\u2010living predators. Morphological alterations required to transform such beetles into extreme myrmecophiles were poorly understood. By studying the cephalic morphology of <jats:italic>Claviger testaceus<\/jats:italic>, we demonstrate that profound changes in all mouthpart components took place during this process, with a highly unusual connection of the maxillae to the hypopharynx, and formation of a uniquely transformed labium with a vestigial prementum. The primary sensory function of the modified maxillary and labial palps is reduced, and the ventral mouthparts transformed into a licking\/\u2018sponging\u2019 device. Many muscles have been reduced, in relation to the coleopteran groundplan or other staphylinoids. The head capsule contains voluminous glands whose appeasement secretions are crucial for the beetle survival in ant colonies. The brain, in turn, has been shifted into the neck region. The prepharyngeal dilator is composed of an entire series of bundles. However, the pharynx does not show any peculiar adaptations to taking up liquid food. We demonstrate that far\u2010reaching cephalic modifications characterize <jats:italic>C. testaceus<\/jats:italic>, and that the development of appeasement glands and adaptation of the mouthparts to trophallaxis determine the head architecture of this extreme myrmecophile.<\/jats:p>","DOI":"10.1002\/jmor.21232","type":"journal-article","created":{"date-parts":[[2020,7,18]],"date-time":"2020-07-18T15:12:04Z","timestamp":1595085124000},"page":"1072-1085","update-policy":"https:\/\/doi.org\/10.1002\/crossmark_policy","source":"Crossref","is-referenced-by-count":20,"title":["Profound head modifications in <scp><i>Claviger testaceus<\/i><\/scp> (Pselaphinae, Staphylinidae, Coleoptera) facilitate integration into communities of ants"],"prefix":"10.1002","volume":"281","author":[{"ORCID":"https:\/\/orcid.org\/0000-0003-2973-1803","authenticated-orcid":false,"given":"Pawe\u0142","family":"Ja\u0142oszy\u0144ski","sequence":"first","affiliation":[{"name":"Museum of Natural History University of Wroc\u0142aw  Wroc\u0142aw Poland"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-5253-267X","authenticated-orcid":false,"given":"Xiao\u2010Zhu","family":"Luo","sequence":"additional","affiliation":[{"name":"Institut f\u00fcr Zoologie und Evolutionsforschung Friedrich\u2010Schiller\u2010Universit\u00e4t Jena  Jena Germany"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Rolf Georg","family":"Beutel","sequence":"additional","affiliation":[{"name":"Institut f\u00fcr Zoologie und Evolutionsforschung Friedrich\u2010Schiller\u2010Universit\u00e4t Jena  Jena Germany"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"311","published-online":{"date-parts":[[2020,7,18]]},"reference":[{"key":"e_1_2_10_2_1","doi-asserted-by":"publisher","DOI":"10.1007\/s00049-002-8330-4"},{"key":"e_1_2_10_3_1","doi-asserted-by":"publisher","DOI":"10.1078\/0044-5231-00107"},{"key":"e_1_2_10_4_1","doi-asserted-by":"publisher","DOI":"10.14411\/eje.2006.014"},{"key":"e_1_2_10_5_1","first-page":"3","article-title":"The head morphology of Dascillus (L.) (Dascilloidea: Dascillidae) and Glaresis Erichson (Scarabeoidea: Glaresidae) and its phylogenetic implications","volume":"70","author":"Anton E.","year":"2012","journal-title":"Arthropod Structure and Development"},{"key":"e_1_2_10_6_1","doi-asserted-by":"publisher","DOI":"10.1002\/jmor.20524"},{"key":"e_1_2_10_7_1","doi-asserted-by":"publisher","DOI":"10.1007\/s13127-016-0305-3"},{"key":"e_1_2_10_8_1","doi-asserted-by":"publisher","DOI":"10.11646\/zootaxa.3861.2.4"},{"key":"e_1_2_10_9_1","first-page":"1","article-title":"Fossil ants (Hymenoptera: Formicidae): Ancient diversity and the rise of modern lineages","volume":"24","author":"Barden P.","year":"2017","journal-title":"Myrmecological News"},{"key":"e_1_2_10_10_1","first-page":"1","article-title":"Skelet und muskulatur des kopfes und thorax von Noterus laevis Sturm: ein Beitrag zur morphologie und phylogenie der Noteridae (Coleoptera: Adephaga)","volume":"462","author":"Belkaceme T.","year":"1991","journal-title":"Stuttgarter Beitr\u00e4ge Zur Naturkunde Serie A (Biologie)"},{"key":"e_1_2_10_11_1","doi-asserted-by":"publisher","DOI":"10.5962\/bhl.part.79801"},{"key":"e_1_2_10_12_1","doi-asserted-by":"publisher","DOI":"10.1515\/9783110264043"},{"key":"e_1_2_10_13_1","doi-asserted-by":"publisher","DOI":"10.1111\/j.1096-0031.2007.00186.x"},{"key":"e_1_2_10_14_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.asd.2009.05.002"},{"key":"e_1_2_10_15_1","doi-asserted-by":"publisher","DOI":"10.1007\/978-3-030-29654-4_12"},{"key":"e_1_2_10_16_1","doi-asserted-by":"publisher","DOI":"10.1007\/BF00389904"},{"key":"e_1_2_10_17_1","first-page":"155","article-title":"Interactions comportementales entre la Fourmi Lasius flavus (Formicidae) et le Col\u00e9opt\u00e8re myrm\u00e9cophile Claviger testaceus (Pselaphidae). I. Ethogramme et modalit\u00e9s des interactions avec les ouvri\u00e8s","volume":"127","author":"Cammaerts R.","year":"1991","journal-title":"Bulletin et Annales de La Soci\u00e9t\u00e9 Royale Belge d'entomologie"},{"key":"e_1_2_10_18_1","doi-asserted-by":"publisher","DOI":"10.1016\/0376-6357(92)90051-E"},{"key":"e_1_2_10_19_1","doi-asserted-by":"publisher","DOI":"10.1016\/0376-6357(94)00068-R"},{"key":"e_1_2_10_20_1","doi-asserted-by":"publisher","DOI":"10.1016\/S0376-6357(96)00040-X"},{"key":"e_1_2_10_21_1","first-page":"433","article-title":"Transport location patterns of the guest beetle Claviger testaceus (Pselaphidae) and other objects moved by workers of the ant, Lasius flavus (Formicidae)","volume":"34","author":"Cammaerts R.","year":"1999","journal-title":"Sociobiology"},{"key":"e_1_2_10_22_1","volume-title":"Biology, morphology and systematics of the ant\u2010like litter beetle genera of Australia (Coleoptera: Staphylinidae: Pselaphinae). Memoirs on Entomology, International 15","author":"Chandler D. S.","year":"2001"},{"key":"e_1_2_10_23_1","doi-asserted-by":"crossref","first-page":"239","DOI":"10.3897\/asp.68.e31730","article-title":"The morphology and evolution of the adult head of Adephaga (Insecta: Coleoptera)","volume":"68","author":"Dressler C.","year":"2010","journal-title":"Arthropod Systematics & Phylogeny"},{"key":"e_1_2_10_24_1","first-page":"33","volume-title":"Biodiversity research and its perspectives in the East Asia","author":"Elmes G. W.","year":"1996"},{"key":"e_1_2_10_25_1","first-page":"19","article-title":"Observations on the feeding behavior of several pselaphid beetles","volume":"67","author":"Engelmann M. D.","year":"1956","journal-title":"Entomological News"},{"key":"e_1_2_10_26_1","doi-asserted-by":"publisher","DOI":"10.1007\/s00114-007-0271-x"},{"key":"e_1_2_10_27_1","doi-asserted-by":"publisher","DOI":"10.11646\/zootaxa.4105.3.4"},{"key":"e_1_2_10_28_1","doi-asserted-by":"publisher","DOI":"10.11646\/zootaxa.3670.3.2"},{"key":"e_1_2_10_29_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.asd.2012.07.002"},{"key":"e_1_2_10_30_1","doi-asserted-by":"crossref","first-page":"17","DOI":"10.1016\/j.asd.2011.07.003","article-title":"The evolution of \u201cdeformed\u201d brains in ant\u2010like stone beetles (Scydmaeninae, Staphylinidae)","volume":"41","author":"Ja\u0142oszy\u0144ski P.","year":"2012","journal-title":"Arthropod Structure and Development"},{"key":"e_1_2_10_31_1","volume-title":"Entomologisches W\u00f6rterbuch mit besonderer Ber\u016bcksichtigung der morphologischen Terminologie","author":"K\u00e9ler S. V.","year":"1963"},{"key":"e_1_2_10_32_1","first-page":"327","article-title":"Beitr\u00e4ge zur anatomie und biologie des Claviger testaceus Preyss","volume":"95","author":"Kr\u00fcger E.","year":"1910","journal-title":"Zeitschrift f\u00fcr Wissenschaftliche Zoologie"},{"key":"e_1_2_10_33_1","first-page":"215","article-title":"Behavioral observations on the myrmecophile Fustiger knausii (Coleoptera: Pselaphidae: Clavigerinae) with a discussion of grasping notches in myrmecophiles","volume":"102","author":"Leschen R. A. B.","year":"1991","journal-title":"Entomological News"},{"key":"e_1_2_10_34_1","first-page":"219","volume-title":"Protopselaphinae new subfamily for Protopselaphus new genus from Malaysia, with a phylogenetic analysis and review of the Omaliine Group of Staphylinidae including Pselaphidae (Coleoptera). Biology, Phylogeny, and Classification of Coleoptera: Papers Celebrating the 80th Birthday of Roy A. Crowson","author":"Newton A. F.","year":"1995"},{"key":"e_1_2_10_35_1","first-page":"123","article-title":"A taxonomic revision of the Clavigerine genus Cerylambus (Insecta, Coleoptera, Staphylinidae, Pselaphinae)","volume":"34","author":"Nomura S.","year":"2008","journal-title":"Bulletin of the National Museum of Nature and Science, Tokyo Series A"},{"key":"e_1_2_10_36_1","doi-asserted-by":"publisher","DOI":"10.1093\/aesa\/25.1.77"},{"key":"e_1_2_10_37_1","first-page":"45","article-title":"Observations on Batrisodes (Coleoptera: Pselaphidae), with particular reference to the American species east of the Rocky Mountains","volume":"8","author":"Park O.","year":"1947","journal-title":"Bulletin of the Chicago Academy of Sciences"},{"key":"e_1_2_10_38_1","doi-asserted-by":"publisher","DOI":"10.1111\/syen.12034"},{"key":"e_1_2_10_39_1","doi-asserted-by":"publisher","DOI":"10.1111\/syen.12173"},{"key":"e_1_2_10_40_1","first-page":"65","article-title":"Myrmecophily in beetles (Coleoptera): Evolutionary patterns and biological mechanisms","volume":"22","author":"Parker J.","year":"2016","journal-title":"Myrmecological News"},{"key":"e_1_2_10_41_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.cub.2014.08.068"},{"key":"e_1_2_10_42_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.asd.2009.08.005"},{"key":"e_1_2_10_43_1","doi-asserted-by":"publisher","DOI":"10.14411\/eje.2008.117"},{"key":"e_1_2_10_44_1","first-page":"296","volume-title":"Handbuch der zoologie\/handbook of zoology, Vol. IV (Arthropoda: Insecta), part 38 Coleoptera, beetles. Volume 1: Morphology and systematics (Archostemata, Adephaga, Myxophaga, Polyphaga partim)","author":"Thayer M. K.","year":"2005"},{"key":"e_1_2_10_45_1","first-page":"63","article-title":"Zur n\u00e4heren Kenntnis des echten Gastverh\u00e4ltnisses (Symphilie) bei den Ameisen und Termiteng\u00e4sten","volume":"28","author":"Wasmann E.","year":"1903","journal-title":"Biologisches Zentralblatt"},{"key":"e_1_2_10_46_1","doi-asserted-by":"publisher","DOI":"10.1002\/jmor.10773"},{"key":"e_1_2_10_47_1","doi-asserted-by":"publisher","DOI":"10.1111\/j.1365-3113.2010.00556.x"},{"key":"e_1_2_10_48_1","doi-asserted-by":"publisher","DOI":"10.2478\/aemnp-2019-0008"},{"key":"e_1_2_10_49_1","doi-asserted-by":"publisher","DOI":"10.1080\/14772019.2017.1313790"}],"container-title":["Journal of Morphology"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/api.wiley.com\/onlinelibrary\/tdm\/v1\/articles\/10.1002%2Fjmor.21232","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/onlinelibrary.wiley.com\/doi\/pdf\/10.1002\/jmor.21232","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/onlinelibrary.wiley.com\/doi\/full-xml\/10.1002\/jmor.21232","content-type":"application\/xml","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/onlinelibrary.wiley.com\/doi\/pdf\/10.1002\/jmor.21232","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2023,9,4]],"date-time":"2023-09-04T09:12:30Z","timestamp":1693818750000},"score":1,"resource":{"primary":{"URL":"https:\/\/onlinelibrary.wiley.com\/doi\/10.1002\/jmor.21232"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2020,7,18]]},"references-count":48,"journal-issue":{"issue":"9","published-print":{"date-parts":[[2020,9]]}},"alternative-id":["10.1002\/jmor.21232"],"URL":"https:\/\/doi.org\/10.1002\/jmor.21232","archive":["Portico"],"relation":{},"ISSN":["0362-2525","1097-4687"],"issn-type":[{"value":"0362-2525","type":"print"},{"value":"1097-4687","type":"electronic"}],"subject":[],"published":{"date-parts":[[2020,7,18]]},"assertion":[{"value":"2020-04-15","order":0,"name":"received","label":"Received","group":{"name":"publication_history","label":"Publication History"}},{"value":"2020-06-14","order":1,"name":"accepted","label":"Accepted","group":{"name":"publication_history","label":"Publication History"}},{"value":"2020-07-18","order":2,"name":"published","label":"Published","group":{"name":"publication_history","label":"Publication History"}}]}}