{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2025,2,21]],"date-time":"2025-02-21T14:09:02Z","timestamp":1740146942002,"version":"3.37.3"},"reference-count":60,"publisher":"Public Library of Science (PLoS)","issue":"11","license":[{"start":{"date-parts":[[2023,11,2]],"date-time":"2023-11-02T00:00:00Z","timestamp":1698883200000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/501100001691","name":"Japan Society for the Promotion of Science","doi-asserted-by":"publisher","award":["22770058"],"award-info":[{"award-number":["22770058"]}],"id":[{"id":"10.13039\/501100001691","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100001691","name":"Japan Society for the Promotion of Science","doi-asserted-by":"publisher","award":["15K07133"],"award-info":[{"award-number":["15K07133"]}],"id":[{"id":"10.13039\/501100001691","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100001691","name":"Japan Society for the Promotion of Science","doi-asserted-by":"publisher","award":["19K06746"],"award-info":[{"award-number":["19K06746"]}],"id":[{"id":"10.13039\/501100001691","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["www.plosone.org"],"crossmark-restriction":false},"short-container-title":["PLoS ONE"],"abstract":"<jats:p>Many animals take advantage of the shaded, humid, and protected environments in subcortical spaces, i.e., thin spaces under the loosened bark of dead trees. Permanent inhabitants of subcortical spaces often show specialized morphologies, such as a miniaturized or dorsoventrally flattened body. However, the evolutionary consequences of these specialized morphologies on behavioral, ecological, and life-history traits have been little studied. We studied the mating biology and anatomy of <jats:italic>Platylabia major<\/jats:italic> (usually placed in the family Anisolabididae), which is an obligate inhabitant of subcortical spaces with a paper-like flattened body, and compared them with those of two thicker, spongiphorid earwigs, <jats:italic>Nesogaster amoenus<\/jats:italic> and <jats:italic>Paralabellula curvicauda<\/jats:italic>. Mating trials in various settings showed that <jats:italic>Pl<\/jats:italic>. <jats:italic>major<\/jats:italic> requires thin spaces sandwiched by two planes to accomplish genital coupling and insemination. In contrast, the thicker species, although also frequently found in subcortical spaces, could mate on a single horizontal plane due to the ability of the male to twist its abdomen through approximately 180\u00b0. Examination by micro-computed tomography and a reagent-based clearing technique revealed no substantive differences in the configuration of mid-abdominal musculature between the species. The dorsal and lateral muscles of <jats:italic>Pl<\/jats:italic>. <jats:italic>major<\/jats:italic>, which are almost parallel to the antero-posterior body axis for accommodation within the thin abdomen, seemed incapable of producing the power to twist the abdomen. The abdominal musculature conforms to a simple pattern in both male and female earwigs, which is repeated in each of the pregenital segments. We conclude that small differences in the range of motion of each abdominal segment can result in large differences in possible mating postures and positions. Surgical experiments also demonstrated that both right and left penises of <jats:italic>Pl<\/jats:italic>. <jats:italic>major<\/jats:italic> are competent and used for insemination with no lateral bias, as in most other earwigs with twin penises studied to date.<\/jats:p>","DOI":"10.1371\/journal.pone.0293701","type":"journal-article","created":{"date-parts":[[2023,11,2]],"date-time":"2023-11-02T17:50:37Z","timestamp":1698947437000},"page":"e0293701","update-policy":"https:\/\/doi.org\/10.1371\/journal.pone.corrections_policy","source":"Crossref","is-referenced-by-count":0,"title":["Subcortical life, evolution of flattened body, and constrained mating posture in the earwig Platylabia major (Insecta: Dermaptera: \u201cAnisolabididae\u201d)"],"prefix":"10.1371","volume":"18","author":[{"ORCID":"https:\/\/orcid.org\/0000-0003-2882-491X","authenticated-orcid":true,"given":"Yoshitaka","family":"Kamimura","sequence":"first","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-8279-4104","authenticated-orcid":true,"given":"Chow-Yang","family":"Lee","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]}],"member":"340","published-online":{"date-parts":[[2023,11,2]]},"reference":[{"issue":"3","key":"pone.0293701.ref001","first-page":"81","article-title":"A review of the anti-predator devices of spiders.","volume":"10","author":"JL Cloudsley-Thompson","year":"1995","journal-title":"Bull Br Arachnol Soc"},{"key":"pone.0293701.ref002","doi-asserted-by":"crossref","first-page":"333","DOI":"10.1111\/j.1095-8312.2000.tb01209.x","article-title":"Trends in the ecological strategies and evolution of millipedes (Diplopoda).","volume":"69","author":"RD Kime","year":"2000","journal-title":"Biological Journal of the Linnean Society"},{"key":"pone.0293701.ref003","first-page":"35","article-title":"The biology of Raphidioptera: A review of present knowledge","volume":"48","author":"H. Asp\u00f6ck","year":"2002","journal-title":"Acta Zoologica Academiae Scientiarum Hungaricae"},{"key":"pone.0293701.ref004","first-page":"272","volume-title":"Staphylinidae Latreille, 1802. American Beetles: Archostemata, Myxophaga, Adephaga, Polyphaga: Staphyliniformia.","author":"AF Newton","year":"2000"},{"key":"pone.0293701.ref005","doi-asserted-by":"crossref","first-page":"423","DOI":"10.1007\/978-94-017-9861-7_15","article-title":"Flat bugs (Aradidae).","author":"C Coscar\u00f3n M del","year":"2015","journal-title":"True Bugs (Heteroptera) of the Neotropics."},{"key":"pone.0293701.ref006","doi-asserted-by":"crossref","first-page":"1957","DOI":"10.1111\/1365-2435.12676","article-title":"Faunal community consequence of interspecific bark trait dissimilarity in early-stage decomposing logs","volume":"30","author":"J Zuo","year":"2016","journal-title":"Funct Ecol"},{"key":"pone.0293701.ref007","doi-asserted-by":"crossref","DOI":"10.3389\/fevo.2022.821335","article-title":"Afraid of the Dark? The Influence of Natural and Artificial Light at Night on the Behavioral Activity of a Nocturnal Gecko.","volume":"10","author":"EJ Nordberg","year":"2022","journal-title":"Front Ecol Evol."},{"key":"pone.0293701.ref008","doi-asserted-by":"crossref","first-page":"394","DOI":"10.1111\/j.1096-0031.2002.tb00158.x","article-title":"The phylogeny of the Histeroidea (Coleoptera: Staphyliniformia).","volume":"18","author":"M. Caterino","year":"2002","journal-title":"Cladistics"},{"key":"pone.0293701.ref009","doi-asserted-by":"crossref","first-page":"259","DOI":"10.11646\/zootaxa.3734.2.9","article-title":"Morphology and life history of Brachypeplus glaber LeConte (Coleoptera: Nitidulidae), with a discussion of multiple life stage data for phylogenetic analyses.","volume":"3734","author":"AR Cline","year":"2013","journal-title":"Zootaxa"},{"key":"pone.0293701.ref010","first-page":"1279","article-title":"The evolution of body shape in response to habitat: Is reproductive output reduced in flat lizards?","volume":"63","author":"BA Goodman","year":"2009","journal-title":"Evolution (N Y)."},{"key":"pone.0293701.ref011","doi-asserted-by":"crossref","first-page":"677","DOI":"10.1111\/aec.12486","article-title":"Extreme specialization to rocky habitats in Tropidurus lizards from Brazil: Trade-offs between a fitted ecomorph and autoecology in a harsh environment.","volume":"42","author":"N Pelegrin","year":"2017","journal-title":"Austral Ecol"},{"key":"pone.0293701.ref012","doi-asserted-by":"crossref","first-page":"233","DOI":"10.11646\/zootaxa.4084.2.4","article-title":"The earwig fauna (Insecta: Dermaptera) of Penang Island, Malaysia, with descriptions of two new species.","volume":"4084","author":"Y Kamimura","year":"2016","journal-title":"Zootaxa"},{"key":"pone.0293701.ref013","doi-asserted-by":"crossref","first-page":"51","DOI":"10.3897\/zookeys.636.10592","article-title":"A new earwig of the genus Echinosoma from Penang Island, Peninsular Malaysia, with notes on the taxonomic and nomenclatural problems of the genus Cranopygia (Insecta, dermaptera, pygidicranidae).","volume":"2016","author":"Y Kamimura","year":"2016","journal-title":"Zookeys."},{"key":"pone.0293701.ref014","first-page":"1","volume":"106","author":"H Steinmann","year":"1990","journal-title":"Dermaptera. Eudermaptera I. Das Tierreich."},{"key":"pone.0293701.ref015","doi-asserted-by":"crossref","DOI":"10.5479\/si.00810282.555","article-title":"The functional morphology of male cerci and associated characters in 13 species of tropical earwigs (Dermaptera: Forficulidae, Labiidae, Carcinophoridae, Pygidicranidae).","author":"RD Brice\u00f1o","year":"1995","journal-title":"Smithson Contrib Zool."},{"key":"pone.0293701.ref016","first-page":"1","article-title":"Dermaptera, Catadermaptera II.","volume":"105","author":"H Steinmann","year":"1989","journal-title":"Das Tierreich"},{"key":"pone.0293701.ref017","doi-asserted-by":"crossref","first-page":"75","DOI":"10.3897\/asp.69.e31739","article-title":"Notes on mating and oviposition of a primitive representative of the higher Forficulina, Apachyus chartaceus (de Haan) (Insecta: Dermaptera: Apachyidae).","volume":"69","author":"S Shimizu","year":"2011","journal-title":"Arthropod Syst Phylogeny"},{"key":"pone.0293701.ref018","first-page":"319","article-title":"Notes on a collection of Dermaptera from India.","volume":"45","author":"GK Srivastava","year":"1970","journal-title":"Eos: revista espa\u00f1ola de entomologia."},{"volume-title":"Fauna of India and adjacent countries, Dermaptera part II: Anisolaboidea. Zoological Survey of India.","year":"2003","author":"GK Srivastava","key":"pone.0293701.ref019"},{"key":"pone.0293701.ref020","doi-asserted-by":"crossref","first-page":"139","DOI":"10.1111\/ens.12058","article-title":"Pre- and postcopulatory sexual selection and the evolution of sexually dimorphic traits in earwigs (Dermaptera).","volume":"17","author":"Y. Kamimura","year":"2014","journal-title":"Entomol Sci."},{"key":"pone.0293701.ref021","first-page":"551","article-title":"Description of Chelisoches chongqingensis sp. nov. (Dermaptera: Chelisochidae) and redescription of Platylabia major Dohrn, 1867 from China.","volume":"4790","author":"XH Ye","year":"2020","journal-title":"Zootaxa"},{"key":"pone.0293701.ref022","doi-asserted-by":"crossref","first-page":"196","DOI":"10.1093\/ilar.52.2.196","article-title":"Anesthesia, Analgesia, and Euthanasia of Invertebrates.","volume":"52","author":"JE Cooper","year":"2011","journal-title":"ILAR J"},{"key":"pone.0293701.ref023","doi-asserted-by":"crossref","first-page":"I","DOI":"10.1016\/j.anbehav.2022.09.006","article-title":"Guidelines for the ethical treatment of nonhuman animals in behavioural research and teaching.","volume":"195","author":"ASAB Ethical Committee\/ABS Animal Care Committee.","year":"2023","journal-title":"Anim Behav"},{"key":"pone.0293701.ref024","doi-asserted-by":"crossref","first-page":"37","DOI":"10.1017\/S0952836903004370","article-title":"Mating behaviour and insemination in Diplatys flavicollis, an earwig with double-barrelled penises and a variable number of female sperm-storage organs","volume":"262","author":"Y. Kamimura","year":"2004","journal-title":"J Zool"},{"key":"pone.0293701.ref025","doi-asserted-by":"crossref","first-page":"1381","DOI":"10.1002\/jmor.10484","article-title":"Right-handed penises of the earwig Labidura riparia (Insecta, Dermaptera, Labiduridae): Evolutionary relationships between structural and behavioral asymmetries","volume":"267","author":"Y. Kamimura","year":"2006","journal-title":"J Morphol"},{"key":"pone.0293701.ref026","doi-asserted-by":"crossref","first-page":"11","DOI":"10.3897\/asp.72.e31779","article-title":"Mating and genital coupling in the primitive earwig species Echinosoma denticulatum (Pygidicranidae): Implications for genital evolution in dermapteran phylogeny","volume":"72","author":"Y Kamimura","year":"2014","journal-title":"Arthropod Syst Phylogeny"},{"key":"pone.0293701.ref027","doi-asserted-by":"crossref","first-page":"844","DOI":"10.1111\/jeb.13486","article-title":"Fitness advantages of the biased use of paired laterally symmetrical penises in an insect","volume":"32","author":"Y Kamimura","year":"2019","journal-title":"J Evol Biol"},{"key":"pone.0293701.ref028","doi-asserted-by":"crossref","first-page":"716","DOI":"10.1093\/biolinnean\/blab111","article-title":"Random or handedness? Use of laterally paired penises in Nala earwigs (Insecta: Dermaptera: Labiduridae).","volume":"134","author":"Y Kamimura","year":"2021","journal-title":"Biological Journal of the Linnean Society"},{"key":"pone.0293701.ref029","doi-asserted-by":"crossref","first-page":"399","DOI":"10.1111\/j.1479-8298.2011.00467.x","article-title":"Comparative copulation anatomy of the Drosophila melanogaster species complex (Diptera: Drosophilidae).","volume":"14","author":"Y Kamimura","year":"2011","journal-title":"Entomol Sci"},{"key":"pone.0293701.ref030","doi-asserted-by":"crossref","first-page":"12825","DOI":"10.1038\/s41598-017-12941-z","article-title":"Escape and surveillance asymmetries in locusts exposed to a Guinea fowl-mimicking robot predator.","volume":"7","author":"D Romano","year":"2017","journal-title":"Sci Rep."},{"key":"pone.0293701.ref031","doi-asserted-by":"crossref","first-page":"4480","DOI":"10.1038\/s41598-020-61410-7","article-title":"Individual, but not population asymmetries, are modulated by social environment and genotype in Drosophila melanogaster.","volume":"10","author":"E Versace","year":"2020","journal-title":"Sci Rep"},{"key":"pone.0293701.ref032","first-page":"103","article-title":"Notes on the genus Platylabia Dohrn (Carcinophoridae: Platylabiinae) with the description of two new species from India.","volume":"4","author":"GK Srivastava","year":"1981","journal-title":"Bulletin of the Zoological Survey of India"},{"key":"pone.0293701.ref033","first-page":"1","article-title":"A new proposed classification of the Dermaptera with special reference to the check list of the Dermaptera of the world.","volume":"20","author":"S. Sakai","year":"1982","journal-title":"Bulletin of Daito Bunka University"},{"journal-title":"R: A Language and Environment for Statistical Computing","year":"2020","author":"R Core Team.","key":"pone.0293701.ref034"},{"key":"pone.0293701.ref035","doi-asserted-by":"crossref","first-page":"289","DOI":"10.1111\/j.2517-6161.1995.tb02031.x","article-title":"Controlling the False Discovery Rate: A Practical and Powerful Approach to Multiple Testing","volume":"57","author":"Y Benjamini","year":"1995","journal-title":"Journal of the Royal Statistical Society Series B (Methodological)."},{"key":"pone.0293701.ref036","first-page":"1","author":"CF Dormann","year":"2016","journal-title":"Overdispersion, and how to deal with it in R and JAGS (requires R-packages AER, coda, lme4, R2jags, DHARMa\/devtools)."},{"key":"pone.0293701.ref037","doi-asserted-by":"crossref","first-page":"753","DOI":"10.1046\/j.1420-9101.1996.9060753.x","article-title":"Dimorphisms and fluctuating asymmetry in the forceps of male earwigs","volume":"9","author":"JL Tomkins","year":"1996","journal-title":"J Evol Biol"},{"key":"pone.0293701.ref038","doi-asserted-by":"crossref","first-page":"475","DOI":"10.1023\/A:1020962606724","article-title":"Forceps size does not determine fighting success in European earwigs","volume":"12","author":"JD Styrsky","year":"1999","journal-title":"J Insect Behav"},{"key":"pone.0293701.ref039","first-page":"207","article-title":"A comparative study of the abdominal musculature of orthopteroid insects.","volume":"14","author":"N. Ford","year":"1923","journal-title":"Trans R Can Inst"},{"key":"pone.0293701.ref040","first-page":"1","article-title":"Studies on the Thoracic Musculature of Insects.","volume":"24","author":"T. Maki","year":"1938","journal-title":"Memories of the Faculty of Science and Agriculture"},{"key":"pone.0293701.ref041","doi-asserted-by":"crossref","first-page":"251","DOI":"10.1111\/j.1096-3642.2001.tb02239.x","article-title":"The female abdomen of the viviparous earwig Hemimerus vosseleri (Insecta: Dermaptera: Hemimeridae), with a discussion of the postgenital abdomen of Insecta.","volume":"131","author":"K-D Klass","year":"2001","journal-title":"Zool J Linn Soc"},{"key":"pone.0293701.ref042","doi-asserted-by":"crossref","first-page":"73","DOI":"10.3897\/asp.65.e31666","article-title":"The potential value of the mid-abdominal musculature and nervous system in the reconstruction of interordinal relationships in lower Neoptera.","volume":"65","author":"R Klug","year":"2007","journal-title":"Arthropod Syst Phylogeny"},{"key":"pone.0293701.ref043","doi-asserted-by":"crossref","first-page":"251","DOI":"10.1111\/j.1469-7998.1959.tb05563.x","article-title":"The Anatomy in Relation To Feeding Habits of Forficula auricularia L. and Other Dermaptera","volume":"133","author":"EJ Popham","year":"1959","journal-title":"Proceedings of the Zoological Society of London"},{"key":"pone.0293701.ref044","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1206\/0003-0082(2007)539[1:FNFED]2.0.CO;2","article-title":"Family-group names for earwigs (Dermaptera).","author":"MS Engel","year":"2007","journal-title":"Am Mus Novit"},{"key":"pone.0293701.ref045","doi-asserted-by":"crossref","first-page":"211","DOI":"10.21805\/bzn.v67i3.a8","article-title":"Case 3522 palicinae Burr, 1910 (Dermaptera, spongiphoridae): proposed emendation of the current spelling to palexinae to remove homonymy with palicidae Bouvier, 1898 (Crustacea, Decapoda).","volume":"67","author":"P Ko\u010d\u00e1rek","year":"2010","journal-title":"Bull Zool Nomencl"},{"key":"pone.0293701.ref046","doi-asserted-by":"crossref","first-page":"120","DOI":"10.21805\/bzn.v74.a030","article-title":"Opinion 2400 (Case 3522)\u2014Palicinae Burr, 1910 (Dermaptera, Spongiphoridae): proposed emendation of spelling to Palexinae to remove homonymy with Palicidae Bouvier, 1898 (Crustacea, Decapoda) not granted.","volume":"74","author":"International Commission on Zoological Nomenclature.","year":"2017","journal-title":"Bull Zool Nomencl."},{"key":"pone.0293701.ref047","first-page":"126","article-title":"A key to Dermapteran subfamilies.","volume":"98","author":"EJ Popham","year":"1965","journal-title":"Entomologist"},{"key":"pone.0293701.ref048","first-page":"241","article-title":"Genera and species of the Dermaptera. 2.","volume":"99","author":"EJ Popham","year":"1966","journal-title":"Brachylabinae and Platylabinae. Entomologist"},{"volume-title":"Dermapterorum Catalogus XIX-XX: Iconographia IV-V. Chelisochidae and Anisolabididae.","year":"1987","author":"S. Sakai","key":"pone.0293701.ref049"},{"key":"pone.0293701.ref050","first-page":"1","article-title":"World catalogue of Dermaptera","volume":"43","author":"H. Steinmann","year":"1989","journal-title":"Series Entomologica (Dordrecht)."},{"key":"pone.0293701.ref051","doi-asserted-by":"crossref","first-page":"73","DOI":"10.26515\/rzsi\/v97\/i1\/1999\/160257","article-title":"On the Higher Classification of Anisolabididae (Insecta: Dermaptera) with a Check-List of Genera and Species.","volume":"97","author":"GK Srivastava","year":"1999","journal-title":"Rec Zool Surv India"},{"key":"pone.0293701.ref052","doi-asserted-by":"crossref","first-page":"331","DOI":"10.3897\/asp.72.e31793","article-title":"Genital morphology and mating behaviour of Allostethus (Dermaptera), an earwig genus of enigmatic phylogenetic position.","volume":"72","author":"Y Kamimura","year":"2014","journal-title":"Arthropod Syst Phylogeny."},{"key":"pone.0293701.ref053","doi-asserted-by":"crossref","first-page":"115","DOI":"10.3897\/zookeys.1146.98248","article-title":"Identification and reproductive isolation of Euborellia species (Insecta, Dermaptera, Anisolabididae) from East and Southeast Asia.","volume":"1146","author":"Y Kamimura","year":"2023","journal-title":"Zookeys"},{"key":"pone.0293701.ref054","first-page":"78","article-title":"The evolution of mating behavior in arthropods","volume":"2","author":"RD Alexander","year":"1964","journal-title":"Royal entomological Society London, Symposia."},{"key":"pone.0293701.ref055","doi-asserted-by":"crossref","first-page":"647","DOI":"10.1111\/j.1469-185X.2007.00029.x","article-title":"The evolution of asymmetric genitalia in spiders and insects","volume":"82","author":"BA Huber","year":"2007","journal-title":"Biological Reviews"},{"key":"pone.0293701.ref056","doi-asserted-by":"crossref","first-page":"157","DOI":"10.3733\/hilg.v41n07p157","article-title":"Biology of the African earwig, Euborellia cincticollis (Gerstaecker) in California and comparative notes on Euborellia annulipes (Lucas).","volume":"41","author":"JJ Knabke","year":"1971","journal-title":"Hilgardia"},{"key":"pone.0293701.ref057","first-page":"303","volume-title":"Dermaptera. Insects of Australia.","author":"ET Giles","year":"1970"},{"key":"pone.0293701.ref058","first-page":"1","article-title":"Dermaptera: Catadermaptera I.","volume":"102","author":"H. Steinmann","year":"1986","journal-title":"Das Tierreich"},{"key":"pone.0293701.ref059","doi-asserted-by":"crossref","first-page":"468","DOI":"10.1007\/s001140100259","article-title":"A \u201cspare\u201d compensates for the risk of destruction of the elongated penis of earwigs (Insecta: Dermaptera).","volume":"88","author":"Y Kamimura","year":"2001","journal-title":"Naturwissenschaften"},{"key":"pone.0293701.ref060","doi-asserted-by":"crossref","first-page":"445","DOI":"10.14411\/eje.2001.065","article-title":"Dermaptera hindwing structure and folding: New evidence for familial, ordinal and superordinal relationships within Neoptera (Insecta).","volume":"98","author":"F Haas","year":"2001","journal-title":"Eur J Entomol"}],"container-title":["PLOS ONE"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/dx.plos.org\/10.1371\/journal.pone.0293701","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2023,11,2]],"date-time":"2023-11-02T17:51:13Z","timestamp":1698947473000},"score":1,"resource":{"primary":{"URL":"https:\/\/dx.plos.org\/10.1371\/journal.pone.0293701"}},"subtitle":[],"editor":[{"given":"Ignacio","family":"Escalante","sequence":"first","affiliation":[],"role":[{"role":"editor","vocabulary":"crossref"}]}],"short-title":[],"issued":{"date-parts":[[2023,11,2]]},"references-count":60,"journal-issue":{"issue":"11","published-online":{"date-parts":[[2023,11,2]]}},"URL":"https:\/\/doi.org\/10.1371\/journal.pone.0293701","relation":{},"ISSN":["1932-6203"],"issn-type":[{"type":"electronic","value":"1932-6203"}],"subject":[],"published":{"date-parts":[[2023,11,2]]}}}