{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,25]],"date-time":"2026-09-25T09:00:44Z","timestamp":1790326844049,"version":"4.1.0"},"reference-count":66,"publisher":"Wiley","issue":"9","license":[{"start":{"date-parts":[[2026,9,14]],"date-time":"2026-09-14T00:00:00Z","timestamp":1789344000000},"content-version":"vor","delay-in-days":13,"URL":"http:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/"},{"start":{"date-parts":[[2026,9,1]],"date-time":"2026-09-01T00:00:00Z","timestamp":1788220800000},"content-version":"tdm","delay-in-days":0,"URL":"http:\/\/doi.wiley.com\/10.1002\/tdm_license_1.1"}],"funder":[{"DOI":"10.13039\/501100001554","name":"Massey University","doi-asserted-by":"publisher","id":[{"id":"10.13039\/501100001554","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["onlinelibrary.wiley.com"],"crossmark-restriction":true},"short-container-title":["Journal of Morphology"],"published-print":{"date-parts":[[2026,9]]},"abstract":"<jats:title>ABSTRACT<\/jats:title>\n                  <jats:p>\n                    Shorebirds within Charadriiformes show substantial variation in where, what, and how they eat. Foraging is aided by a complex feeding apparatus that may include a bill\u2010tip organ with sensory pits and micro tongue tip spines. The morphology of these structures varies among species, but their evolutionary patterns have not been previously mapped. Here we present a comparative morphological analysis of 20 shorebird species across five families, using morphometric data from digital photographs and scanning electron microscopy to reconstruct the evolutionary history of bill\u2010tip organ morphology and micro tongue tip spines. While sensory pits occurred in all species, well\u2010developed bill\u2010tip organs for remote touch were characteristic of Scolopacidae, with pit number and bill length varying among species. Sensory pit numbers were best explained by a Brownian motion model of trait evolution, suggesting increases and decreases of pit numbers across time and high pit numbers in the most recent common ancestor of this clade. Species possessing tongue spines were generally smaller and exhibited a shorter distance between the tongue tip and mandible tip (e.g., the black\u2010fronted dotterel,\n                    <jats:italic>Thinornis melanops<\/jats:italic>\n                    ), supporting a functional role in biofilm grazing. The presence of micro tongue tip spines was best explained by an equal\u2010rates model of trait evolution, with the trait evolving twice with no evidence of subsequent loss: once in the most recent common ancestor shared by calidrid sandpipers (\n                    <jats:italic>Calidris<\/jats:italic>\n                    spp.), shanks (\n                    <jats:italic>Tringa<\/jats:italic>\n                    spp.), the terek sandpiper (\n                    <jats:italic>Xenus cinereus<\/jats:italic>\n                    ), and the ruddy turnstone (\n                    <jats:italic>Arenaria interpres<\/jats:italic>\n                    ), and again in a plover lineage. The phylogenetic distribution of sensory pits and tongue spines showed functional adaptations to diverse foraging strategies. These findings provide new insights into the evolutionary diversification of the shorebird feeding apparatus.\n                  <\/jats:p>","DOI":"10.1002\/jmor.70175","type":"journal-article","created":{"date-parts":[[2026,9,14]],"date-time":"2026-09-14T14:21:17Z","timestamp":1789395677000},"update-policy":"https:\/\/doi.org\/10.1002\/crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["Functional Variation in the Feeding Apparatus of Charadriiform Shorebirds With Emphasis on Bill\u2010Tip Organs and Micro Tongue Tip Spines"],"prefix":"10.1002","volume":"287","author":[{"ORCID":"https:\/\/orcid.org\/0009-0005-9335-2033","authenticated-orcid":false,"given":"Chieh\u2010Ting","family":"Lin","sequence":"first","affiliation":[{"name":"Zoology and Ecology Group, School of Food Technology and Natural Sciences Massey University Palmerston North Manawat\u016b\u2010Whanganui New Zealand"},{"name":"Biodiversity, Genetics and Evolution, School of Environment and Science Griffith University Nathan Queensland Australia"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-7679-0991","authenticated-orcid":false,"given":"Daniel B.","family":"Thomas","sequence":"additional","affiliation":[{"name":"School of Environmental and Animal Sciences MIT\u2010Unitec Institute of Technology Auckland New Zealand"},{"name":"School of Biological Sciences University of Auckland Auckland New Zealand"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-8590-8098","authenticated-orcid":false,"given":"Phil F.","family":"Battley","sequence":"additional","affiliation":[{"name":"Zoology and Ecology Group, School of Food Technology and Natural Sciences Massey University Palmerston North Manawat\u016b\u2010Whanganui New Zealand"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"311","published-online":{"date-parts":[[2026,9,14]]},"reference":[{"issue":"2","key":"e_1_2_11_2_1","first-page":"76","article-title":"A Review of Bone Preparation Techniques for Anatomical Studies","volume":"3","author":"Ajayi A.","year":"2016","journal-title":"Malaya Journal of Biosciences"},{"key":"e_1_2_11_3_1","doi-asserted-by":"publisher","DOI":"10.2307\/3544520"},{"key":"e_1_2_11_4_1","doi-asserted-by":"publisher","DOI":"10.1675\/1524-4695(2003)026[0026:SFBWIS]2.0.CO;2"},{"key":"e_1_2_11_5_1","doi-asserted-by":"publisher","DOI":"10.1002\/ecs2.3133"},{"key":"e_1_2_11_6_1","volume-title":"Model Selection and Multimodel Inference: A Practical Information\u2010Theoretic Approach","author":"Burnham K. P.","year":"2002"},{"key":"e_1_2_11_7_1","unstructured":"Burton P. J. K.1974. Feeding and the Feeding Apparatus in Waders: A Study of Anatomy And Adaptations in the Charadrii: British Museum (Natural History). Publication; 719."},{"issue":"3","key":"e_1_2_11_8_1","first-page":"233","article-title":"Phylogenetic Analysis. Models and Estimation Procedures","volume":"19","author":"Cavalli\u2010Sforza L. L.","year":"1967","journal-title":"American Journal of Human Genetics"},{"key":"e_1_2_11_9_1","doi-asserted-by":"publisher","DOI":"10.1071\/ZO9920291"},{"key":"e_1_2_11_10_1","doi-asserted-by":"publisher","DOI":"10.1525\/9780520947962"},{"key":"e_1_2_11_11_1","doi-asserted-by":"publisher","DOI":"10.1525\/auk.2009.09117"},{"key":"e_1_2_11_12_1","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pone.0080036"},{"key":"e_1_2_11_13_1","doi-asserted-by":"publisher","DOI":"10.1080\/00063658609476896"},{"key":"e_1_2_11_14_1","unstructured":"Dowding J. E.2022. Variable Oystercatcher | T\u014drea Pango. New Zealand Birds Online.https:\/\/nzbirdsonline.org.nz\/species\/variable-oystercatcher."},{"key":"e_1_2_11_15_1","doi-asserted-by":"publisher","DOI":"10.1080\/00063657.2013.836153"},{"key":"e_1_2_11_16_1","first-page":"52","volume-title":"Theory, Practice, & Procedures. Facility for Advance Instrumentation","author":"Dunlap M.","year":"1997"},{"key":"e_1_2_11_17_1","doi-asserted-by":"publisher","DOI":"10.1007\/s00227-004-1521-5"},{"key":"e_1_2_11_18_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.aanat.2013.09.005"},{"key":"e_1_2_11_19_1","doi-asserted-by":"publisher","DOI":"10.1111\/azo.12075"},{"key":"e_1_2_11_20_1","doi-asserted-by":"publisher","DOI":"10.1080\/00063659409477205"},{"key":"e_1_2_11_21_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.ympev.2012.03.008"},{"key":"e_1_2_11_22_1","volume-title":"Shorebirds: An Identification Guide to the Waders of the World","author":"Hayman P.","year":"1986"},{"key":"e_1_2_11_23_1","doi-asserted-by":"publisher","DOI":"10.1111\/j.1469-7998.1970.tb02053.x"},{"key":"e_1_2_11_24_1","volume-title":"Waders of Southern Africa","author":"Hockey P. A. R.","year":"1995"},{"issue":"1","key":"e_1_2_11_25_1","first-page":"89","article-title":"The Oystercatcher Haematopus ostralegus as a Predator of the Bivalve Macoma balthica in the Dutch Wadden Sea","volume":"55","author":"Hulscher J. B.","year":"1982","journal-title":"Ardea"},{"key":"e_1_2_11_26_1","doi-asserted-by":"publisher","DOI":"10.1038\/nature11631"},{"key":"e_1_2_11_27_1","unstructured":"Jetz W. G. H.Thomas J. B.Joy K.Hartmann andA. O.Mooers.2014. A Global Phylogeny of Birds.https:\/\/birdtree.org\/."},{"key":"e_1_2_11_28_1","volume-title":"The Avian Tongue","author":"Johnston N. E.","year":"2014"},{"key":"e_1_2_11_29_1","volume-title":"Shorebirds: An Illustrated Behavioural Ecology","author":"Kam J.","year":"2004"},{"key":"e_1_2_11_30_1","doi-asserted-by":"publisher","DOI":"10.1093\/molbev\/msaa191"},{"key":"e_1_2_11_31_1","doi-asserted-by":"publisher","DOI":"10.1890\/07-1442.1"},{"key":"e_1_2_11_32_1","doi-asserted-by":"publisher","DOI":"10.1111\/j.1461-0248.2012.01744.x"},{"key":"e_1_2_11_33_1","doi-asserted-by":"publisher","DOI":"10.1007\/s12237-015-0029-1"},{"key":"e_1_2_11_34_1","doi-asserted-by":"publisher","DOI":"10.1098\/rspb.2008.1110"},{"key":"e_1_2_11_35_1","doi-asserted-by":"publisher","DOI":"10.4081\/rio.2026.950"},{"key":"e_1_2_11_36_1","doi-asserted-by":"publisher","DOI":"10.1675\/063.033.0305"},{"key":"e_1_2_11_37_1","doi-asserted-by":"publisher","DOI":"10.1111\/j.1096-3642.2010.00654.x"},{"key":"e_1_2_11_38_1","doi-asserted-by":"publisher","DOI":"10.2307\/4089332"},{"key":"e_1_2_11_39_1","doi-asserted-by":"publisher","DOI":"10.1111\/j.2041-210X.2012.00196.x"},{"key":"e_1_2_11_40_1","doi-asserted-by":"publisher","DOI":"10.1163\/1570756054472818"},{"key":"e_1_2_11_41_1","first-page":"35","article-title":"The Evolution of Sexual Bill\u2010Size Dimorphism in Shorebirds: A Morphometric Test of the Resource Partitioning Hypothesis","volume":"13","author":"Nebel S.","year":"2011","journal-title":"Evolutionary Ecology Research"},{"key":"e_1_2_11_42_1","doi-asserted-by":"publisher","DOI":"10.1038\/44766"},{"key":"e_1_2_11_43_1","doi-asserted-by":"publisher","DOI":"10.1093\/bioinformatics\/bty633"},{"key":"e_1_2_11_44_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.ecss.2014.11.013"},{"key":"e_1_2_11_45_1","doi-asserted-by":"publisher","DOI":"10.1093\/bioinformatics\/btu181"},{"key":"e_1_2_11_46_1","doi-asserted-by":"publisher","DOI":"10.1080\/03014223.1985.10428298"},{"key":"e_1_2_11_47_1","doi-asserted-by":"publisher","DOI":"10.1098\/rspb.1998.0445"},{"key":"e_1_2_11_48_1","doi-asserted-by":"publisher","DOI":"10.1111\/j.2041-210X.2011.00169.x"},{"key":"e_1_2_11_49_1","doi-asserted-by":"publisher","DOI":"10.1007\/978-3-030-13739-7_17"},{"key":"e_1_2_11_50_1","doi-asserted-by":"publisher","DOI":"10.1159\/000006580"},{"key":"e_1_2_11_51_1","volume-title":"The Roles of Morphology, Individuality and Arrival From Migration in the Foraging Ecology of Bar\u2010tailed Godwits at the Manawat\u016b River Estuary: A Thesis Presented in Partial Fulfilment of the Requirements for the Degree of Master of Science in Zoology at Massey University, Manawat\u016b","author":"Ross T. A.","year":"2018"},{"key":"e_1_2_11_52_1","doi-asserted-by":"publisher","DOI":"10.2307\/2986146"},{"key":"e_1_2_11_53_1","doi-asserted-by":"publisher","DOI":"10.1093\/auk\/110.2.169"},{"key":"e_1_2_11_54_1","doi-asserted-by":"publisher","DOI":"10.18476\/sbna.v9.a16"},{"key":"e_1_2_11_55_1","doi-asserted-by":"publisher","DOI":"10.1038\/nmeth.2089"},{"key":"e_1_2_11_56_1","unstructured":"Steed C.2017. Grey\u2010Tailed Tattler. New Zealand Birds Online.https:\/\/nzbirdsonline.org.nz\/species\/grey-tailed-tattler."},{"key":"e_1_2_11_57_1","doi-asserted-by":"publisher","DOI":"10.1007\/s002270000406"},{"key":"e_1_2_11_58_1","doi-asserted-by":"publisher","DOI":"10.1111\/j.2041-210X.2011.00132.x"},{"key":"e_1_2_11_59_1","doi-asserted-by":"publisher","DOI":"10.1111\/j.1365-2435.2006.01073.x"},{"key":"e_1_2_11_60_1","unstructured":"du Toit C. J.2022. \u201cMechanosensory Structures in the Beaks of Probe\u2010Foraging Birds in Relation to Their Foraging Ecology.\u201d (PhD). FitzPatrick Institute of African Ornithology Department of Biological Sciences Faculty of Science University of Cape Town.http:\/\/hdl.handle.net\/11427\/36755."},{"key":"e_1_2_11_61_1","doi-asserted-by":"publisher","DOI":"10.1098\/rspb.2020.2322"},{"key":"e_1_2_11_62_1","doi-asserted-by":"publisher","DOI":"10.1111\/joa.13734"},{"key":"e_1_2_11_63_1","doi-asserted-by":"publisher","DOI":"10.1111\/jav.03243"},{"key":"e_1_2_11_64_1","volume-title":"Exploratory Data Analysis","author":"Tukey J. W.","year":"1977"},{"key":"e_1_2_11_65_1","doi-asserted-by":"publisher","DOI":"10.1111\/j.1474-919X.1993.tb02827.x"},{"key":"e_1_2_11_66_1","doi-asserted-by":"publisher","DOI":"10.2307\/5168"},{"key":"e_1_2_11_67_1","unstructured":"Withington R. J.2015. The Foraging Ecology of Non\u2010Breeding Wrybills (Anarhynchus frontalis) in the Firth of Thames: A Thesis presented in Partial Fulfilment of the Requirements for the Degree of Master of Science in Ecology at Massey University Palmerston North New Zealand.http:\/\/hdl.handle.net\/10179\/7898."}],"container-title":["Journal of Morphology"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/onlinelibrary.wiley.com\/doi\/pdf\/10.1002\/jmor.70175","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/onlinelibrary.wiley.com\/doi\/full-xml\/10.1002\/jmor.70175","content-type":"application\/xml","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/onlinelibrary.wiley.com\/doi\/pdf\/10.1002\/jmor.70175","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,9,25]],"date-time":"2026-09-25T08:05:35Z","timestamp":1790323535000},"score":1,"resource":{"primary":{"URL":"https:\/\/onlinelibrary.wiley.com\/doi\/10.1002\/jmor.70175"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,9]]},"references-count":66,"journal-issue":{"issue":"9","published-print":{"date-parts":[[2026,9]]}},"alternative-id":["10.1002\/jmor.70175"],"URL":"https:\/\/doi.org\/10.1002\/jmor.70175","archive":["Portico"],"relation":{},"ISSN":["0362-2525","1097-4687"],"issn-type":[{"value":"0362-2525","type":"print"},{"value":"1097-4687","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,9]]},"assertion":[{"value":"2026-07-02","order":0,"name":"received","label":"Received","group":{"name":"publication_history","label":"Publication History"}},{"value":"2026-09-05","order":2,"name":"accepted","label":"Accepted","group":{"name":"publication_history","label":"Publication History"}},{"value":"2026-09-14","order":3,"name":"published","label":"Published","group":{"name":"publication_history","label":"Publication History"}}],"article-number":"e70175"}}