{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,20]],"date-time":"2026-10-20T17:00:18Z","timestamp":1792515618510,"version":"4.5.0"},"reference-count":93,"publisher":"Wiley","issue":"2","license":[{"start":{"date-parts":[[2023,9,20]],"date-time":"2023-09-20T00:00:00Z","timestamp":1695168000000},"content-version":"am","delay-in-days":342,"URL":"http:\/\/onlinelibrary.wiley.com\/termsAndConditions#am"},{"start":{"date-parts":[[2022,10,13]],"date-time":"2022-10-13T00:00:00Z","timestamp":1665619200000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/onlinelibrary.wiley.com\/termsAndConditions#vor"}],"funder":[{"DOI":"10.13039\/100000001","name":"National Science Foundation","doi-asserted-by":"publisher","award":["DBI 1759637"],"award-info":[{"award-number":["DBI 1759637"]}],"id":[{"id":"10.13039\/100000001","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000001","name":"National Science Foundation","doi-asserted-by":"publisher","award":["PRFB 1712015"],"award-info":[{"award-number":["PRFB 1712015"]}],"id":[{"id":"10.13039\/100000001","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["anatomypubs.onlinelibrary.wiley.com"],"crossmark-restriction":true},"short-container-title":["The Anatomical Record"],"published-print":{"date-parts":[[2023,2]]},"abstract":"<jats:title>Abstract<\/jats:title><jats:p>Complex prey processing requires the repositioning of food between the teeth, as modulated by a soft tissue appendage like a tongue or lips. In this study, we trace the evolution of lips and ligaments, which are used during prey capture and prey processing in an herbivorous group of fishes. Pacus (Serrasalmidae) are Neotropical freshwater fishes that feed on leaves, fruits, and seeds. These prey are hard or tough, require high forces to fracture, contain abrasive or caustic elements, or deform considerably before failure. Pacus are gape\u2010limited and do not have the pharyngeal jaws many bony fishes use to dismantle and\/or transport prey. Despite their gape limitation, pacus feed on prey larger than their mouths, relying on robust teeth and a hypertrophied lower lip for manipulation and breakdown of food. We used histology to compare the lip morphology across 14 species of pacus and piranhas to better understand this soft tissue. We found that frugivorous pacus have larger, more complex lips which are innervated and folded at their surface, while grazing species have callused, mucus\u2010covered lips. Unlike mammalian lips or tongues, pacu lips lack any intrinsic skeletal or smooth muscle. This implies that pacu lips lack dexterity; however, we found a novel connection to the primordial ligament which suggests that the lips are actuated by the jaw adductors. We propose that pacus combine hydraulic repositioning of prey inside the buccal cavity with direct oral manipulation, the latter using a combination of a morphologically heterodont dentition and compliant lips for reorienting food.<\/jats:p>","DOI":"10.1002\/ar.25075","type":"journal-article","created":{"date-parts":[[2022,9,21]],"date-time":"2022-09-21T01:53:23Z","timestamp":1663725203000},"page":"326-342","update-policy":"https:\/\/doi.org\/10.1002\/crossmark_policy","source":"Crossref","is-referenced-by-count":12,"title":["Lip service: Histological phenotypes correlate with diet and feeding ecology in herbivorous pacus"],"prefix":"10.1002","volume":"306","author":[{"ORCID":"https:\/\/orcid.org\/0000-0001-6556-5414","authenticated-orcid":false,"given":"Karly E.","family":"Cohen","sequence":"first","affiliation":[{"name":"Biology Department University of Washington  Seattle Washington USA"},{"name":"Friday Harbor Laboratories University of Washington  Friday Harbor USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Oliver","family":"Lucanus","sequence":"additional","affiliation":[{"name":"BelowWater, Inc.  Montreal Quebec Canada"},{"name":"Applied Remote Sensing Lab, Department of Geography McGill University  Montreal Quebec Canada"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-1930-9748","authenticated-orcid":false,"given":"Adam P.","family":"Summers","sequence":"additional","affiliation":[{"name":"Biology Department University of Washington  Seattle Washington USA"},{"name":"Friday Harbor Laboratories University of Washington  Friday Harbor USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-9748-2066","authenticated-orcid":false,"given":"Matthew A.","family":"Kolmann","sequence":"additional","affiliation":[{"name":"Museum of Paleontology University of Michigan  Ann Arbor Michigan USA"},{"name":"Dept. of Biology University of Louisville  Louisville Kentucky USA"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"311","published-online":{"date-parts":[[2022,10,13]]},"reference":[{"key":"e_1_2_7_2_1","doi-asserted-by":"publisher","DOI":"10.2307\/1374678"},{"key":"e_1_2_7_3_1","doi-asserted-by":"publisher","DOI":"10.1111\/j.1096-3642.1964.tb00493.x"},{"key":"e_1_2_7_4_1","doi-asserted-by":"crossref","unstructured":"Alexander R. McN. (1967). The functions and mechanisms of the protrusible upper jaws of some acanthopterygian fish.Journal of Zoology 151(1) 43\u201364.https:\/\/doi.org\/10.1111\/j.1469-7998.1967.tb02865.x","DOI":"10.1111\/j.1469-7998.1967.tb02865.x"},{"key":"e_1_2_7_5_1","doi-asserted-by":"publisher","DOI":"10.1111\/jfb.12868"},{"key":"e_1_2_7_6_1","doi-asserted-by":"publisher","DOI":"10.1007\/s10750-018-3838-y"},{"key":"e_1_2_7_7_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.envpol.2018.10.088"},{"key":"e_1_2_7_8_1","doi-asserted-by":"publisher","DOI":"10.1111\/j.1096-3642.1974.tb00030.x"},{"key":"e_1_2_7_9_1","volume-title":"The Red Devil\u2010Midas\u2010Arrow Cichlid Species Complex in Nicaragua","author":"Barlow G. W.","year":"1976"},{"key":"e_1_2_7_10_1","doi-asserted-by":"publisher","DOI":"10.1111\/bij.12502"},{"key":"e_1_2_7_11_1","doi-asserted-by":"publisher","DOI":"10.2307\/1382464"},{"key":"e_1_2_7_12_1","doi-asserted-by":"publisher","DOI":"10.1111\/j.1096-3642.1986.tb00638.x"},{"key":"e_1_2_7_13_1","doi-asserted-by":"publisher","DOI":"10.1016\/0003-9969(75)90152-1"},{"key":"e_1_2_7_14_1","doi-asserted-by":"publisher","DOI":"10.1111\/evo.13649"},{"key":"e_1_2_7_15_1","doi-asserted-by":"publisher","DOI":"10.1007\/s10750-014-1960-z"},{"key":"e_1_2_7_16_1","doi-asserted-by":"publisher","DOI":"10.1093\/iob\/obaa009"},{"key":"e_1_2_7_17_1","doi-asserted-by":"publisher","DOI":"10.1111\/btp.13022"},{"key":"e_1_2_7_18_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.zool.2019.02.001"},{"key":"e_1_2_7_19_1","doi-asserted-by":"publisher","DOI":"10.1111\/joa.13227"},{"key":"e_1_2_7_20_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.biocon.2015.06.019"},{"key":"e_1_2_7_21_1","doi-asserted-by":"publisher","DOI":"10.1890\/13-0393.1"},{"key":"e_1_2_7_22_1","doi-asserted-by":"publisher","DOI":"10.1641\/B570907"},{"key":"e_1_2_7_23_1","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pone.0053274"},{"key":"e_1_2_7_24_1","doi-asserted-by":"publisher","DOI":"10.1590\/1982-0224-20160173"},{"key":"e_1_2_7_25_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.zool.2011.09.008"},{"key":"e_1_2_7_26_1","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pone.0060846"},{"key":"e_1_2_7_27_1","doi-asserted-by":"publisher","DOI":"10.1002\/jmor.20286"},{"key":"e_1_2_7_28_1","doi-asserted-by":"publisher","DOI":"10.1098\/rsbl.2005.0319"},{"key":"e_1_2_7_29_1","doi-asserted-by":"publisher","DOI":"10.1002\/jmor.10966"},{"key":"e_1_2_7_30_1","doi-asserted-by":"publisher","DOI":"10.1590\/S2179-975X2014000400006"},{"key":"e_1_2_7_31_1","doi-asserted-by":"publisher","DOI":"10.1098\/rsbl.2021.0431"},{"key":"e_1_2_7_32_1","doi-asserted-by":"publisher","DOI":"10.1007\/978-0-387-73906-9_1"},{"key":"e_1_2_7_33_1","unstructured":"Froese R.&Pauly D.(2014).Fish Base. World Wide Web Electronic Publication.http:\/\/www.fishbase.org"},{"key":"e_1_2_7_34_1","first-page":"47","article-title":"A head with a suckermouth: A functional\u2010morphological study of the head of the suckermouth armoured catfish Ancistrus cf. triradiatus (Loricariidae, Siluriformes)","volume":"137","author":"Geerinckx T.","year":"2007","journal-title":"Belgian Journal of Zoology"},{"key":"e_1_2_7_35_1","doi-asserted-by":"publisher","DOI":"10.1242\/jeb.041012"},{"key":"e_1_2_7_36_1","first-page":"329","article-title":"Aspects of the diet and feeding ecologies of fish from nine floodplain lakes of the lower Caura, Venezuelan Guayana","volume":"12","author":"Gonz\u00e1lez N.","year":"2003","journal-title":"Scientia Guaianae"},{"key":"e_1_2_7_37_1","doi-asserted-by":"publisher","DOI":"10.2307\/1444953"},{"key":"e_1_2_7_38_1","doi-asserted-by":"publisher","DOI":"10.1525\/9780520316133"},{"key":"e_1_2_7_39_1","doi-asserted-by":"publisher","DOI":"10.1590\/S0101-81751982000200001"},{"key":"e_1_2_7_40_1","doi-asserted-by":"publisher","DOI":"10.1038\/srep01009"},{"key":"e_1_2_7_41_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.ppees.2007.01.001"},{"key":"e_1_2_7_42_1","doi-asserted-by":"publisher","DOI":"10.1016\/0006-3207(74)90026-3"},{"key":"e_1_2_7_43_1","doi-asserted-by":"publisher","DOI":"10.1177\/154411130301400604"},{"key":"e_1_2_7_44_1","doi-asserted-by":"publisher","DOI":"10.1590\/1809-43921974042047"},{"key":"e_1_2_7_45_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.cub.2017.04.056"},{"key":"e_1_2_7_46_1","doi-asserted-by":"publisher","DOI":"10.1635\/053.166.0116"},{"key":"e_1_2_7_47_1","volume-title":"Analysis of design in the feeding apparatus of snakes and fishes. [Doctoral dissertation, Harvard University, Cambridge Massachusetts]","author":"Irish F. J.","year":"1987"},{"key":"e_1_2_7_48_1","doi-asserted-by":"publisher","DOI":"10.1111\/j.1096-3642.1985.tb01178.x"},{"key":"e_1_2_7_49_1","first-page":"e12306","article-title":"Tooth and consequences: Heterodonty and dental replacement in piranhas and pacus (Serrasalmidae)","volume":"20","author":"Kolmann M. A.","year":"2019","journal-title":"Evolution & Development"},{"key":"e_1_2_7_50_1","doi-asserted-by":"publisher","DOI":"10.1093\/sysbio\/syaa065"},{"key":"e_1_2_7_51_1","doi-asserted-by":"publisher","DOI":"10.1098\/rsos.171581"},{"key":"e_1_2_7_52_1","doi-asserted-by":"publisher","DOI":"10.1098\/rspb.2016.1392"},{"key":"e_1_2_7_53_1","doi-asserted-by":"publisher","DOI":"10.1890\/0012-9615(1999)069[0001:DBRATM]2.0.CO;2"},{"key":"e_1_2_7_54_1","first-page":"353","article-title":"Food habits of two species of Acnodon (Characiformes, Serrasalmidae) and scale\u2010eating habits of Acnodon normani","volume":"14","author":"Leite R. G.","year":"1990","journal-title":"Cybium"},{"key":"e_1_2_7_55_1","doi-asserted-by":"publisher","DOI":"10.1007\/978-3-642-81073-2"},{"key":"e_1_2_7_56_1","first-page":"E143","volume-title":"Integrative and comparative biology","author":"Lomax J. J.","year":"2020"},{"key":"e_1_2_7_57_1","doi-asserted-by":"publisher","DOI":"10.1111\/j.1744-7429.2008.00415.x"},{"key":"e_1_2_7_58_1","doi-asserted-by":"publisher","DOI":"10.1006\/anbo.2000.1261"},{"key":"e_1_2_7_59_1","doi-asserted-by":"publisher","DOI":"10.1002\/jmor.11003"},{"key":"e_1_2_7_60_1","doi-asserted-by":"publisher","DOI":"10.1016\/0300-5712(77)90117-8"},{"key":"e_1_2_7_61_1","first-page":"19","article-title":"Studies on the subfamily Serrasalminae, part III. On the generic status and phylogenetic relationships of the genera Pygopristis, Pygocentrus, Pristobrycon, and Serrasalmus (Teleostei\u2010Characidae\u2010Serrasalminae)","volume":"12","author":"Machado\u2010Allison A.","year":"1985","journal-title":"Acta Biologica Venezuela"},{"key":"e_1_2_7_62_1","doi-asserted-by":"publisher","DOI":"10.1111\/evo.12367"},{"key":"e_1_2_7_63_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.zool.2011.05.005"},{"key":"e_1_2_7_64_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.ympev.2020.106945"},{"key":"e_1_2_7_65_1","doi-asserted-by":"publisher","DOI":"10.1023\/A:1011033025706"},{"key":"e_1_2_7_66_1","doi-asserted-by":"publisher","DOI":"10.1007\/s00357-014-9161-z"},{"key":"e_1_2_7_67_1","unstructured":"NicoLG.1991.Trophic ecology of piranhas (Characidae: Serrasalminae) from savanna and forest regions in the Orinoco River basin of Venezuela. Ph.D. dissertation Univ. of Florida Gainesville."},{"key":"e_1_2_7_68_1","doi-asserted-by":"publisher","DOI":"10.2307\/2388321"},{"key":"e_1_2_7_69_1","doi-asserted-by":"publisher","DOI":"10.1242\/jeb.239509"},{"key":"e_1_2_7_70_1","doi-asserted-by":"publisher","DOI":"10.1163\/002829669X00134"},{"key":"e_1_2_7_71_1","doi-asserted-by":"publisher","DOI":"10.1111\/j.1439-0310.1975.tb02010.x"},{"key":"e_1_2_7_72_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.actao.2004.08.004"},{"key":"e_1_2_7_73_1","doi-asserted-by":"publisher","DOI":"10.1093\/sysbio\/syac001"},{"key":"e_1_2_7_74_1","doi-asserted-by":"publisher","DOI":"10.1590\/S1679-62252007000300019"},{"key":"e_1_2_7_75_1","doi-asserted-by":"publisher","DOI":"10.1590\/1809-4392201600123"},{"key":"e_1_2_7_76_1","doi-asserted-by":"publisher","DOI":"10.1590\/1676-0611-bn-2017-0471"},{"key":"e_1_2_7_77_1","doi-asserted-by":"publisher","DOI":"10.1111\/j.2041-210X.2011.00169.x"},{"key":"e_1_2_7_78_1","doi-asserted-by":"publisher","DOI":"10.1093\/icb\/icm044"},{"key":"e_1_2_7_79_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.zool.2005.12.005"},{"key":"e_1_2_7_80_1","doi-asserted-by":"publisher","DOI":"10.1111\/j.1095-8649.1986.tb04926.x"},{"key":"e_1_2_7_81_1","doi-asserted-by":"publisher","DOI":"10.2307\/2413111"},{"key":"e_1_2_7_82_1","doi-asserted-by":"publisher","DOI":"10.1242\/jeb.220749"},{"key":"e_1_2_7_83_1","doi-asserted-by":"publisher","DOI":"10.1111\/j.1469-7998.1976.tb04664.x"},{"key":"e_1_2_7_84_1","doi-asserted-by":"publisher","DOI":"10.1016\/S0169-5347(98)01491-8"},{"key":"e_1_2_7_85_1","doi-asserted-by":"publisher","DOI":"10.1007\/978-1-60761-950-5_2"},{"key":"e_1_2_7_86_1","doi-asserted-by":"publisher","DOI":"10.1590\/S1984-46702014000100004"},{"key":"e_1_2_7_87_1","doi-asserted-by":"publisher","DOI":"10.1093\/bioinformatics\/btl117"},{"key":"e_1_2_7_88_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.ympev.2014.06.018"},{"key":"e_1_2_7_89_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.tice.2010.04.005"},{"key":"e_1_2_7_90_1","doi-asserted-by":"publisher","DOI":"10.1007\/s004420050349"},{"key":"e_1_2_7_91_1","doi-asserted-by":"publisher","DOI":"10.1590\/1809-43921979091143"},{"key":"e_1_2_7_92_1","doi-asserted-by":"publisher","DOI":"10.1088\/1748-3190\/ab9418"},{"key":"e_1_2_7_93_1","doi-asserted-by":"publisher","DOI":"10.2307\/1445434"},{"key":"e_1_2_7_94_1","doi-asserted-by":"publisher","DOI":"10.1111\/j.1469-7998.1999.tb00763.x"}],"container-title":["The Anatomical Record"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/onlinelibrary.wiley.com\/doi\/pdf\/10.1002\/ar.25075","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/onlinelibrary.wiley.com\/doi\/full-xml\/10.1002\/ar.25075","content-type":"application\/xml","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/anatomypubs.onlinelibrary.wiley.com\/doi\/am-pdf\/10.1002\/ar.25075","content-type":"application\/pdf","content-version":"am","intended-application":"syndication"},{"URL":"https:\/\/anatomypubs.onlinelibrary.wiley.com\/doi\/pdf\/10.1002\/ar.25075","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,10,8]],"date-time":"2025-10-08T12:33:25Z","timestamp":1759926805000},"score":1,"resource":{"primary":{"URL":"https:\/\/anatomypubs.onlinelibrary.wiley.com\/doi\/10.1002\/ar.25075"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2022,10,13]]},"references-count":93,"journal-issue":{"issue":"2","published-print":{"date-parts":[[2023,2]]}},"alternative-id":["10.1002\/ar.25075"],"URL":"https:\/\/doi.org\/10.1002\/ar.25075","archive":["Portico"],"relation":{},"ISSN":["1932-8486","1932-8494"],"issn-type":[{"value":"1932-8486","type":"print"},{"value":"1932-8494","type":"electronic"}],"subject":[],"published":{"date-parts":[[2022,10,13]]},"assertion":[{"value":"2022-02-11","order":0,"name":"received","label":"Received","group":{"name":"publication_history","label":"Publication History"}},{"value":"2022-08-24","order":2,"name":"accepted","label":"Accepted","group":{"name":"publication_history","label":"Publication History"}},{"value":"2022-10-13","order":3,"name":"published","label":"Published","group":{"name":"publication_history","label":"Publication History"}}]}}