{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,6,26]],"date-time":"2026-06-26T03:16:29Z","timestamp":1782443789113,"version":"3.54.5"},"reference-count":24,"publisher":"Wiley","issue":"3","license":[{"start":{"date-parts":[[2023,3,13]],"date-time":"2023-03-13T00:00:00Z","timestamp":1678665600000},"content-version":"am","delay-in-days":12,"URL":"http:\/\/creativecommons.org\/licenses\/by\/4.0\/"},{"start":{"date-parts":[[2023,3,13]],"date-time":"2023-03-13T00:00:00Z","timestamp":1678665600000},"content-version":"vor","delay-in-days":12,"URL":"http:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/100000001","name":"National Science Foundation","doi-asserted-by":"publisher","award":["DEB1936971"],"award-info":[{"award-number":["DEB1936971"]}],"id":[{"id":"10.13039\/100000001","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["esajournals.onlinelibrary.wiley.com"],"crossmark-restriction":true},"short-container-title":["Ecosphere"],"published-print":{"date-parts":[[2023,3]]},"abstract":"<jats:title>Abstract<\/jats:title><jats:p>Temperate deciduous forests by definition include a large proportion of woody species that shed their leaves each autumn and are completely leafless during winter months. Leaf senescence in deciduous trees is an active, complex process typically involving the physiological formation of an abscission layer causing the petiole to mechanically detach from the branch. However, several deciduous species retain all or some senesced leaves on branches through much of winter, a phenomenon called leaf marcescence. Marcescence has long fascinated botanists, including Pehr Kalm as early as 1749. Yet, surprisingly little research has been done to date. Here, we review and explore patterns and mechanisms of leaf marcescence in temperate forests, bringing together six nonmutually exclusive but separately proposed hypotheses: (1) Marcescence has no adaptive function but rather an evolutionary byproduct; (2) Marcescent leaves deter winter browsing herbivores; (3) Leaf retention through winter improves nutrient resorption during autumn senescence; (4) Prolonged leaf shedding into spring minimizes nutrient leaching and promotes decomposition; (5) Marcescent leaves protect overwintering buds from frost or desiccation; and (6) Marcescent canopies provide winter cover for animals (including insects, birds, bats), thereby affecting plant nutrient availability via excrement. No hypothesis has complete support and few tests of multiple hypotheses have been done. It is likely that any adaptive value of marcescence is species and context dependent. Despite increased interest in plant phenology and prevalence of this trait, much remains to be understood on the physiology, evolution, function, and ecological implications of leaf marcescence.<\/jats:p>","DOI":"10.1002\/ecs2.4410","type":"journal-article","created":{"date-parts":[[2023,3,13]],"date-time":"2023-03-13T11:54:36Z","timestamp":1678708476000},"update-policy":"https:\/\/doi.org\/10.1002\/crossmark_policy","source":"Crossref","is-referenced-by-count":16,"title":["Not all temperate deciduous trees are leafless in winter: The curious case of marcescence"],"prefix":"10.1002","volume":"14","author":[{"ORCID":"https:\/\/orcid.org\/0000-0003-0756-5090","authenticated-orcid":false,"given":"J. Mason","family":"Heberling","sequence":"first","affiliation":[{"name":"Section of Botany Carnegie Museum of Natural History  Pittsburgh Pennsylvania USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-6278-2798","authenticated-orcid":false,"given":"Rose\u2010Marie","family":"Muzika","sequence":"additional","affiliation":[{"name":"Section of Botany Carnegie Museum of Natural History  Pittsburgh Pennsylvania USA"},{"name":"Powdermill Nature Reserve Carnegie Museum of Natural History  Rector Pennsylvania USA"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"311","published-online":{"date-parts":[[2023,3,13]]},"reference":[{"key":"e_1_2_5_2_1","doi-asserted-by":"publisher","DOI":"10.1111\/j.1365-3040.1996.tb00403.x"},{"key":"e_1_2_5_3_1","doi-asserted-by":"publisher","DOI":"10.1007\/s11104-017-3318-6"},{"key":"e_1_2_5_4_1","doi-asserted-by":"publisher","DOI":"10.1007\/s11104-022-05341-4"},{"key":"e_1_2_5_5_1","doi-asserted-by":"publisher","DOI":"10.1086\/334178"},{"key":"e_1_2_5_6_1","doi-asserted-by":"publisher","DOI":"10.2307\/3565549"},{"key":"e_1_2_5_7_1","unstructured":"GBIF (Global Biodiversity Information Facility).2022a.\u201cGBIF Occurrence\u201d (9 August 2022).https:\/\/doi.org\/10.15468\/dl.uucms9."},{"key":"e_1_2_5_8_1","unstructured":"GBIF (Global Biodiversity Information Facility).2022b.\u201cGBIF Occurrence\u201d (3 November 2022).https:\/\/doi.org\/10.15468\/dl.g272me."},{"key":"e_1_2_5_9_1","doi-asserted-by":"publisher","DOI":"10.1080\/0028825X.2004.9512933"},{"key":"e_1_2_5_10_1","doi-asserted-by":"publisher","DOI":"10.1111\/j.1469-8137.2011.03663.x"},{"key":"e_1_2_5_11_1","doi-asserted-by":"crossref","unstructured":"Heberling J. M. andR.Muzika.2022.\u201cData for: \u2018Not all Temperate Deciduous Trees Are Leafless in Winter: The Curious Case of Marcescence\u2019.\u201d Zenodo.https:\/\/doi.org\/10.5281\/zenodo.6979970.","DOI":"10.1002\/ecs2.4410"},{"key":"e_1_2_5_12_1","doi-asserted-by":"publisher","DOI":"10.1086\/335558"},{"key":"e_1_2_5_13_1","volume-title":"Travels into North America: Containing Its Natural History, and a Circumstantial Account of Its Plantations and Agriculture in General, with the Civil, Ecclesiastical and Commercial State of the Country, the Manners of the Inhabitants, and Several Curious and Important Remarks on Various Subjects","author":"Kalm P.","year":"1776"},{"key":"e_1_2_5_14_1","doi-asserted-by":"publisher","DOI":"10.1007\/s00442-007-0709-z"},{"key":"e_1_2_5_15_1","doi-asserted-by":"publisher","DOI":"10.1002\/ajb2.1763"},{"key":"e_1_2_5_16_1","doi-asserted-by":"publisher","DOI":"10.1111\/gcb.14904"},{"key":"e_1_2_5_17_1","doi-asserted-by":"publisher","DOI":"10.1016\/B978-0-12-425050-5.50014-5"},{"key":"e_1_2_5_18_1","doi-asserted-by":"publisher","DOI":"10.1111\/j.1600-0706.2008.17293.x"},{"key":"e_1_2_5_19_1","doi-asserted-by":"publisher","DOI":"10.2307\/3544599"},{"key":"e_1_2_5_20_1","doi-asserted-by":"publisher","DOI":"10.2307\/3544134"},{"key":"e_1_2_5_21_1","doi-asserted-by":"publisher","DOI":"10.1007\/s11829-021-09815-7"},{"key":"e_1_2_5_22_1","doi-asserted-by":"publisher","DOI":"10.1111\/ele.13512"},{"key":"e_1_2_5_23_1","doi-asserted-by":"publisher","DOI":"10.2307\/2388171"},{"key":"e_1_2_5_24_1","first-page":"475","article-title":"Effects of Marcescent Leaves on Winter Browsing by Large Herbivores in Northern Temperate Deciduous Forests","volume":"37","author":"Svendsen C. 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