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Historical maps confirm the accuracy of zero?inflated model predictions of ancient tree abundance in English wood?pastures

Nolan, Victoria; Reader, Tom; Gilbert, Francis; Atkinson, Nick


Victoria Nolan

Associate Professor

Nick Atkinson


Ancient trees have important ecological, historical and social connections, and are a key source of dead and decaying wood, a globally declining resource. Wood-pastures, which combine livestock grazing, open spaces and scattered trees, are significant reservoirs of ancient trees, yet information about their true abundance within wood-pastures is limited. England has extensive databases of both ancient trees and wood-pasture habitat, providing a unique opportunity for the first large-scale, national case study to address this knowledge gap. We investigated the relationship between the abundance of ancient trees in a large sample of English wood-pastures (5,571) and various unique environmental, historical and anthropogenic predictors, to identify wood-pastures with high numbers of undiscovered ancient trees. A major challenge in many modelling studies is obtaining independent data for model verification: here we introduce a novel model verification step using series of historic maps with detailed records of trees to validate our model predictions. This desk-based method enables rapid verification of model predictions using completely independent data across a large geographical area, without the need for, or limitations associated with, extensive field surveys. Historic map verification estimates correlated well with model predictions of tree abundance. Model predictions suggest there are ~101,400 undiscovered ancient trees in all wood-pastures in England, around 10 times the total current number of ancient tree records. Important predictors of ancient tree abundance included wood-pasture area, distance to several features including cities, commons, historic Royal forests and Tudor deer parks, and different types of soil and land classes. Synthesis and applications. Historical maps and statistical models can be used in combination to produce accurate predictions of ancient tree abundance in wood-pastures, and inform future targeted surveys of wood-pasture habitat, with a focus on those deemed to have undiscovered ancient trees. This study provides support for improvements to conservation policy and protection measures for ancient trees and wood-pastures.


Nolan, V., Reader, T., Gilbert, F., & Atkinson, N. (2021). Historical maps confirm the accuracy of zero?inflated model predictions of ancient tree abundance in English wood?pastures. Journal of Applied Ecology, 58(11), 2661-2672.

Journal Article Type Article
Acceptance Date Aug 3, 2021
Online Publication Date Sep 4, 2021
Publication Date 2021-11
Deposit Date Sep 13, 2021
Publicly Available Date Sep 5, 2022
Journal Journal of Applied Ecology
Print ISSN 0021-8901
Electronic ISSN 1365-2664
Publisher Wiley
Peer Reviewed Peer Reviewed
Volume 58
Issue 11
Pages 2661-2672
Keywords Ecology
Public URL
Publisher URL
Additional Information This is the peer reviewed version of the following article: Nolan, V., Reader, T., Gilbert, F., & Atkinson, N. (2021). Historical maps confirm the accuracy of zero-inflated model predictions of ancient tree abundance in English wood-pastures. Journal of Applied Ecology, 58, 2661– 2672., which has been published in final form at This article may be used for non-commercial purposes in accordance with Wiley Terms and Conditions for Use of Self-Archived Versions. This article may not be enhanced, enriched or otherwise transformed into a derivative work, without express permission from Wiley or by statutory rights under applicable legislation. Copyright notices must not be removed, obscured or modified. The article must be linked to Wiley’s version of record on Wiley Online Library and any embedding, framing or otherwise making available the article or pages thereof by third parties from platforms, services and websites other than Wiley Online Library must be prohibited.


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