{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,12]],"date-time":"2026-10-12T03:34:51Z","timestamp":1791776091062,"version":"4.3.4"},"reference-count":182,"publisher":"Wiley","license":[{"start":{"date-parts":[[2026,9,20]],"date-time":"2026-09-20T00:00:00Z","timestamp":1789862400000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/"},{"start":{"date-parts":[[2026,9,20]],"date-time":"2026-09-20T00:00:00Z","timestamp":1789862400000},"content-version":"tdm","delay-in-days":0,"URL":"http:\/\/doi.wiley.com\/10.1002\/tdm_license_1.1"}],"content-domain":{"domain":["onlinelibrary.wiley.com"],"crossmark-restriction":true},"short-container-title":["Biological Reviews"],"abstract":"<jats:title>ABSTRACT<\/jats:title>\n                  <jats:p>\n                    For millennia, human activities have transformed ecosystems globally. Determining which landscapes are predominantly anthropogenic is crucial to conservation and habitat restoration. Madagascar, a critically\u2010threatened biodiversity hotspot, is a prominent example. Currently 80% of non\u2010cropland habitat is tree\u2010limited savanna falling under the tropical grassy biome (savanna\/TGB). Savanna\/TGB is particularly prevalent in the Dry Deciduous Forest (DDF) in the north\/northwest and below the treeline in the Central Highlands (CH). Relative savanna\/TGB cover when agropastoralism arrived &lt;1300\u2009years ago (ya) has been intensely debated. Two competing hypotheses have emerged. The Mixed Mosaic Perspective argues current savanna\/TGB is mostly anthropogenic, and differs from endemic, tree\u2010rich savannas, because it has been produced by frequent agropastoralist fires, grazing by 7\u201310 million zebu cattle, and habitat clearance. These novel disturbances replaced a mosaic of savanna\/TGB, comprised of C\n                    <jats:sub>3<\/jats:sub>\n                    and C\n                    <jats:sub>4<\/jats:sub>\n                    graminoids (both grasses and sedges), forest, heathland and wetland, with non\u2010endemic fire\u2010 and grazing\u2010adapted pantropical C\n                    <jats:sub>4<\/jats:sub>\n                    grasses. This perspective is founded on multiple lines of evidence: &gt;90% of endemic vertebrates are forest\u2010adapted, Malagasy people named many locations currently in savanna\/TGB after forest, and only 1000 ya, abundant tree\u2010adapted fauna with predominantly C\n                    <jats:sub>3<\/jats:sub>\n                    diets inhabited the CH and DDF. The Vast Grassland Perspective (VGP) contends that late Holocene aridity favoured vast savanna\/TGB, and that cattle and human\u2010lit fires maintain savanna\/TGB previously grazed by now\u2010extinct megafauna. This view contends that ecological models predict \u2018treeless grassland\u2019 across the CH and DDF, with savanna\/TGB woodland naturally limited to &lt;5%, and it questions palaeoecological evidence as spatially\u2010limited. Clarifying which viewpoint is correct is fundamental to conservation; if human\u2010lit fires and cattle grazing are not surrogates for past ecological processes, they may be detrimental to ecosystem function. We address this debate by applying ecological models that predict grassy habitat likelihood based on precipitation, and, for the first time, map these likelihoods across the entire CH and DDF. We review all available literature and apply novel interpolations of palaeo\u2010data to highlight existing spatial coverage and broad spatiotemporal trends. We then assess if predicted habitats from our ecological models concur with reconstructed habitat at the onset of pastoralism. Ecological models align with palaeo\u2010data at &gt;90% of sites but diverge markedly from current conditions. Today, savanna\/TGB is three to seven\u00a0times\u00a0more extensive than predicted, with &gt;183,000\u2009km\n                    <jats:sup>2<\/jats:sup>\n                    less forest than models suggest should exist, given current precipitation. Multiple lines of evidence indicate widespread anthropogenic transformation started with the arrival of agropastoralism ~1300 ya, and began shifting biomes to tree\u2010limited savanna by ~1100\u20131000 ya. Charcoal records show increased fire activity preceded decline of forest, tree\u2010rich savanna woodlands, heathland, and populations of large, now\u2010extinct, endemic vertebrates. Paradoxically, this happened when DDF climate was relatively wet, which should have promoted, not reduced, forest, and limited fire. Expansion of savanna\/TGB dominated by C\n                    <jats:sub>4<\/jats:sub>\n                    grasses followed the increase in fire coinciding with the onset of pastoralism. In the context of a biome transition that is still unfolding, we present a critical pre\u2010agropastoralist baseline that can guide ecological restoration efforts to reinstate lost ecological functions and preserve Madagascar's remaining extraordinary endemic biota.\n                  <\/jats:p>","DOI":"10.1002\/brv.70211","type":"journal-article","created":{"date-parts":[[2026,9,21]],"date-time":"2026-09-21T00:36:30Z","timestamp":1789950990000},"update-policy":"https:\/\/doi.org\/10.1002\/crossmark_policy","source":"Crossref","is-referenced-by-count":1,"title":["Resolving the biome debate: ecological and palaeo\u2010data concur that the extensive tree\u2010limited grassy biomes of central and north\u2010western Madagascar are largely anthropogenic"],"prefix":"10.1002","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-8731-9828","authenticated-orcid":false,"given":"Grant S.","family":"Joseph","sequence":"first","affiliation":[{"name":"FitzPatrick Institute of African Ornithology, DSI\u2010NRF Centre of Excellence, Department of Biological Sciences University of Cape Town  Rondebosch 7701 South Africa"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-6147-7118","authenticated-orcid":false,"given":"Sean","family":"Hixon","sequence":"additional","affiliation":[{"name":"Department of Integrative Biology Oregon State University  Corvallis OR 97322 USA"},{"name":"Max Planck Institute of Geoanthropology  Kahlaische Strasse 10 07745 Jena Germany"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-6729-2576","authenticated-orcid":false,"given":"Colleen L.","family":"Seymour","sequence":"additional","affiliation":[{"name":"South African National Biodiversity Institute, Kirstenbosch Research Centre  Private Bag X7 Claremont 7735 South Africa"},{"name":"Department of Conservation Ecology and Entomology Stellenbosch University  Stellenbosch 7602 South Africa"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-9997-0207","authenticated-orcid":false,"given":"Laurie R.","family":"Godfrey","sequence":"additional","affiliation":[{"name":"Department of Anthropology University of Massachusetts  Amherst MA 01003 USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Miguel","family":"Pedrono","sequence":"additional","affiliation":[{"name":"CIRAD  Ampandrianomby B.P. 853 Antananarivo 101 Madagascar"},{"name":"UAR PatriNat, Mus\u00e9um National d'Histoire Naturelle  Paris 75005 France"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-3597-9215","authenticated-orcid":false,"given":"Karen E.","family":"Samonds","sequence":"additional","affiliation":[{"name":"Department of Biological Sciences Northern Illinois University  DeKalb 60115 USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-9318-7465","authenticated-orcid":false,"given":"Andrinajoro R.","family":"Rakotoarivelo","sequence":"additional","affiliation":[{"name":"Afro\u2010mountain Research Unit University of the Free State, Qwaqwa  Private Bag X13 Phuthaditjhaba 9866 South Africa"},{"name":"Natiora Ahy  Lot IIU57K Bis, Ampahibe Antananarivo 101 Madagascar"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-8462-6806","authenticated-orcid":false,"given":"Brooke E.","family":"Crowley","sequence":"additional","affiliation":[{"name":"Departments of Geosciences and Anthropology University of Cincinnati  Cincinnati OH 45221 USA"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"311","published-online":{"date-parts":[[2026,9,20]]},"reference":[{"key":"e_1_2_11_3_1","doi-asserted-by":"crossref","first-page":"623","DOI":"10.1006\/jasc.1996.0146","article-title":"Preliminary vegetation maps of the world since the last glacial maximum: an aid to archaeological understanding","volume":"24","author":"Adams J. 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