Articles
Ancient woodland indicator species: can old herbarium specimens supplement recent records to inform ecological management?
Kelly Hemmings
Published 2022-12-15
Keywords
- Ancient forest,
- connectivity,
- continuity forest,
- herbaria,
- inventory
- natural history,
- restoration,
- old growth forest ...More
How to Cite
Hemmings, K. (2022). Ancient woodland indicator species: can old herbarium specimens supplement recent records to inform ecological management?. Webbia, 77(2), 327–336. https://doi.org/10.36253/jopt-13400
Abstract
Old herbarium specimens have become increasingly well-recognised as a rich source of ecological baseline data. For long-continuity plant communities, such as ancient woodland, these records may be particularly important for present day ecological management. To evaluate this potential, searches for pre-1950 Ancient Woodland Indicator (AWI) herbarium specimens collected in East Gloucestershire, UK, were conducted using digital open access sources and the physical Royal Agricultural University herbarium. In total 305 specimens were retrieved from twelve herbaria, with small regional collections being particularly important sources. The earliest specimen dated to 1834. There was a significant association between old specimen availability and year of collection, due to a peak in the late-1800s and early-1900s. Over half of the AWI species for the region were represented, although some taxonomic bias was evident. To determine if old AWI specimens contributed any new location records, 246 unique specimens with detailed georeferences were mapped and compared to the locations of 1950-1999 and 2000-2021 biological records. One third of the pre-1950 specimens had not been recorded in the same locality since collection of the old specimen, indicating either a gap in recent records or floristic change. However, length of time since specimen collection was not a predictor of a 1950-2000 or 2000-2021 record in the same locality. Overall, it is highly recommended that policy-makers, land managers, and field surveyors consult old AWI herbarium records for ancient woodland identification, management, and restoration.
References
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Albani Rocchetti G, Armstrong CG, Abeli T, Orsenigo S, Jasper C., Joly S, Bruneau A. Zytaruk M, Vamosi, JC. 2021. Reversing extinction trends: new uses of (old) herbarium specimens to accelerate conservation action on threatened species. New Phytol. 230:433–45. https://doi.org/10.1111/nph.17133
Applequist WL, Mcglinn DJ, Miller M, Long QG, Miller JS. 2007. How well do herbarium data predict the locationof present populations? A test using Echinacea species in Missouri. Biodivers Conserv. 16(5): 1397-1407. https://doi.org/10.1007/s10531-006-6737-x
Baldini RM, Cristofolini G, Aedo, C. 2022. The extant herbaria from the Sixteenth Century: a synopsis. Webbia. 77(1): 23-33. https://doi.org/10.36253/jopt-13038
Bergès L, Dupouey JL. 2021. Historical ecology and ancient forests: Progress, conservation issues and scientific prospects, with some examples from the French case. J Veg Sci, 32(1), e12846. https://doi.org/10.1111/jvs.12846
Botanical Society of Britain and Ireland Distribution Database. 2000- . BSBI [accessed 2022 Apr 25]. https://database.bsbi.org/
British Geological Survey. Apr 2022. Data from: geology of Britain viewer. [accessed 2002 Apr 25]. https://mapapps.bgs.ac.uk/geologyofbritain/home.html
Brown ND, Curtis T, Adams EC. 2015. Effects of clear-felling versus gradual removal of conifer trees on the survival of understorey plants during the restoration of ancient woodlands. Forest Ecol Manag. 348: 15-22. https://doi.org/10.1016/j.foreco.2015.03.030
Broughton RK, Bullock JM, George C, Hill RA, Hinsley SA, Maziarz M, Melin M, Mountford JO, Sparks TH, Pywell RF. 2021. Long-term woodland restoration on lowland farmland through passive rewilding. Plos one. 16(6), e0252466. https://doi.org/10.1371/journal.pone.0252466
Colombo B, Kaehier M, Calvente A. 2016. An inventory of the Bignoniaceae from the Brazilian state of Rio Grande do Norte highlights the importance of small herbaria to biodiversity studies. Phytotaxa. 278(1): 19-28. https://doi.org/10.11646/phytotaxa.278.1.2
Cornwell WK, Pearse WD, Dalrymple RL, Zanne AE. 2019. What we (don't) know about global plant diversity. Ecography. 42: 1819– 1831. https://doi.org/10.1111/ecog.04481
Craig M, Buckley P, Howell R. 2015. Responses of an ancient woodland field layer to soil translocation: methods and timing. Appl Veg Sci. 18(4): 579-590. https://doi.org/10.1111/avsc.12170
Daru BH, Park DS, Primack RB, Willis CG, Barrington DS, Whitfeld TJ, Seidler TG, Sweeney PW, Foster DR, Ellison AM, et al. (2018). Widespread sampling biases in herbaria revealed from large‐scale digitization. New Phytol. 217(2): 939-955. https://doi.org/10.1111/nph.14855
Dyderski MK, Tyborski J, Jagodziński AM. 2017. The utility of ancient forest indicator species in urban environments: A case study from Poznań, Poland. Urban For Urban Gree. 27: 76-83. https://doi.org/10.1016/j.ufug.2017.06.016
Global Biodiversity Information Facility database. 2001- .GBIF [accessed 2022 Apr 25]. https/www.GBIF.org.
Glaves P, Handley C , Birkbeck J, Rotherham ID, Wright B. 2009. A survey of the coverage, use and application of ancient woodland indicator lists in the UK. Hallam Environmental Consultants Ltd., Biodiversity and Landscape History Research Institute/Geography, Tourism and Environment Change Research Unit, Sheffield Hallam University, Sheffield. Available from: https://core.ac.uk/download/pdf/4149223.pdf
Greve M, Lykke AM, Fagg CW, Gereau RE, Lewis GP, Marchant R, Marshall AR, Ndaysishimiye J, Bogaert J, Svenning JC. 2016. Realising the potential of herbarium records for conservation biology. S Afr J Bot. 105: 317-323. https://doi.org/10.1016/j.sajb.2016.03.017
Heberling, JM. (2022). Herbaria as big data sources of plant traits. Int J Plant Sci. 183(2): 87-118. https://doi.org/10.1086/717623
Hedrick BP, Heberling JM, Meineke EK, Turner KG, Grassa CJ, Park DS, Kennedy J, Clarke JA, Cook JA, Blackburn DC. 2020. Digitization and the future of natural history collections. BioScience. 70(3): 243-251. https://doi.org/10.1093/biosci/biz163
Herbaria United database. 2006 - . Botanical Society of Britain and Ireland and Botanical Collection Managers Group. [accessed 2022 Apr 25]. http://herbariaunited.org/
Hermy M, Honnay O, Firbank L, Grashof-Bokdam C, Lawesson JE. 1999. An ecological comparison between ancient and other forest plant species of Europe, and the implications for forest conservation. Biol Conserv. 91(1): 9-22. https://doi.org/10.1016/S0006-3207(99)00045-2
HM Government. 2018. A green future: our 25 year plan to improve the environment. London: Department for Environment, Fisheries and Rural Affairs. Available from: https://www. gov. uk/government/publications/25-year-environment-plan.
James SA, Soltis PS, Belbin L, Chapman AD, Nelson G, Paul DL, Collins M. (2018). Herbarium data: Global biodiversity and societal botanical needs for novel research. App Plant Sci, 6(2), e1024. https://doi.org/10.1002/aps3.1024
Kelemen K, Kriván A, Standovár T. 2014. Effects of land‐use history and current management on ancient woodland herbs in Western Hungary. J Veg Sci. 25(1): 172-183. https://doi.org/10.1111/jvs.12046
Kirby K. 2004. Table of ancient woodland indicator plants. In: Rose F. O'Reilly C, Smith D.P. and Collings. M, editors. The Wild Flower Key: How to Identify Wild Flowers, Trees and Shrubs in Britain and Ireland. London, UK: Warne; p. 558-561
Kricsfalusy VV, Trevisan N. (2014). Prioritizing regionally rare plant species for conservation using herbarium data. Biodivers Conserv. 23(1): 39-61. https://doi.org/10.1007/s10531-013-0583-4
Lang PL, Willems FM, Scheepens JF, Burbano HA, Bossdorf, O. 2019. Using herbaria to study global environmental change. New Phytol. 221(1): 110-122. https://doi.org/10.1111/nph.15401
Lavoie C. 2013. Biological collections in an ever changing world: Herbaria as tools for biogeographical and environmental studies. Perspect Plant Ecol. 15(1), 68-76. https://doi.org/10.1016/j.ppees.2012.10.002
Lienert J, Fischer M, Diemer M. 2002. Local extinctions of the wetland specialist Swertia perennis L.(Gentianaceae) in Switzerland: a revisitation study based on herbarium records. Biol Conserv. 103(1): 65-76. https://doi.org/10.1016/S0006-3207(01)00121-5
Le Bras G, Pignal M, Jeanson ML, Muller S, Aupic C, Carré B, Flament G, Gaudeul M, Gonçalves C, Invernón VR et al. 2017) The French Muséum national d’histoire naturelle vascular plant herbarium collection dataset. Sci data. 4(1): 1-16. https://doi.org/10.1038/sdata.2017.16
Marsico TD, Krimmel ER, Carter JR, Gillespie EL, Lowe PD, McCauley R, Morris AB, Nelson G, Smith M, Soteropoulos D, et al. 2020. Small herbaria contribute unique biogeographic records to county, locality, and temporal scales. Am J Bot. 107(11): 1577-1587. https://doi.org/10.1002/ajb2.1563
Martine CT, Ward ME. 2013. Establishment of regional herbarium leads to more than 200 new flora atlas records for New York State. J Torrey Bot Soc. 140(1): 125-131. https://doi.org/10.3159/TORREY-D-12-00036.1
McMullin RT, Wiersma YF. 2019. Out with OLD growth, in with ecological continNEWity: new perspectives on forest conservation. Front Ecol Environ. 17(3): 176-181. https://doi.org/10.1002/fee.2016
Meineke EK, Davis CC, Davies TJ. 2018. The unrealized potential of herbaria for global change biology. Ecol Monogr. 88(4): 505-525. https://doi.org/10.1002/ecm.1307
Ministry of Housing, Communities and Local Government. 2019. National planning policy framework. London: Ministry of Housing, Communities and Local Government. [accessed 2022 Apr 25]. https://assets.publishing.service.gov.uk/government/uploads/system/uploads/attachment_data/file/1005759/NPPF_July_2021.pdf
Natural England (2022). Ancient woodland inventory. Open government licence. [accessed 2022 Apr 25]. https://data.gov.uk/dataset/9461f463-c363-4309-ae77-fdcd7e9df7d3/ancient-woodland-england
Palo A, Ivask M, Liira J. 2013. Biodiversity composition reflects the history of ancient semi-natural woodland and forest habitats—compilation of an indicator complex for restoration practice. Ecol Indic. 34: 336-344. https://doi.org/10.1016/j.ecolind.2013.05.020
Pescott OL, Humphrey TA, Walker KJ. 2018. A short guide to using British and Irish plant occurrence data for research. Wallingford, NERC/Centre for Ecology & Hydrology. Available from: http://nora.nerc.ac.uk/id/eprint/520609/
Peterken GF. 1974. A method for assessing woodland flora for conservation using indicator species. Biol Conserv. 6(4): 239-245. https://doi.org/10.1016/0006-3207(74)90001-9
Rackham O. 2008. Ancient woodlands: modern threats. New Phytol. 180(3): 571-586. http://www.jstor.org/stable/25150608.
Razzaque J, Lester C. 2021. Why Protect Ancient Woodland in the UK? Rethinking the Ecosystem Approach. Transnatl Env La. 10(1): 135-158. https://doi.org/10.1017/S2047102520000333
Reid C, Hornigold K, McHenry E, Nichols C, Townsend M, Lewthwaite K, Elliot M, Pullinger R, Hotchkiss A, Gilmartin E, et al. 2021. State of the UK's Woods and Trees 2021.Grantham: The Woodland Trust.
Rose F. 1999. Indicators of ancient woodland-the use of vascular plants in evaluating ancient woods for nature conservation. Br Wild. 10: 241-251.
Rotherham ID. 2011. A landscape history approach to the assessment of ancient woodlands. In, Wallace EB, editor. Woodlands: ecology, management and conservation,. New York: Nova Science; p. 161-184.
Stace C. 2019. New Flora of the British Isles (4th ed). UK: C&M Floristics.
Swallow KA, Wood MJ, Goodenough AE. 2020. Relative contribution of ancient woodland indicator and non‐indicator species to herb layer distinctiveness in ancient semi‐natural, ancient replanted, and recent woodland. Appl Veg Sci. 23(4): 471-481.
https://doi.org/10.1111/avsc.12501
Troudet J, Grandcolas P, Blin A, Vignes-Lebbe R, Legendre F. 2017. Taxonomic bias in biodiversity data and societal preferences. Sci Rep. 7(1): 1-14.
https://doi.org/10.1038/s41598-017-09084-6
Walker K, Pearman D, Ellis B, McIntosh J, Lockton A. 2010. Recording the British and Irish flora 2010-2020. London: Botanical Society Britain and Ireland.
Willis KJ, Araújo MB, Bennett KD, Figueroa-Rangel B, Froyd CA, Myers N. 2007. How can a knowledge of the past help to conserve the future? Biodiversity conservation and the relevance of long-term ecological studies. Philos Trans R Soc Lond B Bio Sci. 362(1478): 175-187. https://doi.org/10.1098/rstb.2006.1977
Webb JC, Goodenough AE. 2018. Questioning the reliability of “ancient” woodland indicators: Resilience to interruptions and persistence following deforestation. Ecol Indic. 84: 354-363. https://doi.org/10.1016/j.ecolind.2017.09.010
Wulf M. 1997. Plant species as indicators of ancient woodland in northwestern Germany. J Veg Sci. 8(5): 635-642. https://doi.org/10.2307/3237367
Albani Rocchetti G, Armstrong CG, Abeli T, Orsenigo S, Jasper C., Joly S, Bruneau A. Zytaruk M, Vamosi, JC. 2021. Reversing extinction trends: new uses of (old) herbarium specimens to accelerate conservation action on threatened species. New Phytol. 230:433–45. https://doi.org/10.1111/nph.17133
Applequist WL, Mcglinn DJ, Miller M, Long QG, Miller JS. 2007. How well do herbarium data predict the locationof present populations? A test using Echinacea species in Missouri. Biodivers Conserv. 16(5): 1397-1407. https://doi.org/10.1007/s10531-006-6737-x
Baldini RM, Cristofolini G, Aedo, C. 2022. The extant herbaria from the Sixteenth Century: a synopsis. Webbia. 77(1): 23-33. https://doi.org/10.36253/jopt-13038
Bergès L, Dupouey JL. 2021. Historical ecology and ancient forests: Progress, conservation issues and scientific prospects, with some examples from the French case. J Veg Sci, 32(1), e12846. https://doi.org/10.1111/jvs.12846
Botanical Society of Britain and Ireland Distribution Database. 2000- . BSBI [accessed 2022 Apr 25]. https://database.bsbi.org/
British Geological Survey. Apr 2022. Data from: geology of Britain viewer. [accessed 2002 Apr 25]. https://mapapps.bgs.ac.uk/geologyofbritain/home.html
Brown ND, Curtis T, Adams EC. 2015. Effects of clear-felling versus gradual removal of conifer trees on the survival of understorey plants during the restoration of ancient woodlands. Forest Ecol Manag. 348: 15-22. https://doi.org/10.1016/j.foreco.2015.03.030
Broughton RK, Bullock JM, George C, Hill RA, Hinsley SA, Maziarz M, Melin M, Mountford JO, Sparks TH, Pywell RF. 2021. Long-term woodland restoration on lowland farmland through passive rewilding. Plos one. 16(6), e0252466. https://doi.org/10.1371/journal.pone.0252466
Colombo B, Kaehier M, Calvente A. 2016. An inventory of the Bignoniaceae from the Brazilian state of Rio Grande do Norte highlights the importance of small herbaria to biodiversity studies. Phytotaxa. 278(1): 19-28. https://doi.org/10.11646/phytotaxa.278.1.2
Cornwell WK, Pearse WD, Dalrymple RL, Zanne AE. 2019. What we (don't) know about global plant diversity. Ecography. 42: 1819– 1831. https://doi.org/10.1111/ecog.04481
Craig M, Buckley P, Howell R. 2015. Responses of an ancient woodland field layer to soil translocation: methods and timing. Appl Veg Sci. 18(4): 579-590. https://doi.org/10.1111/avsc.12170
Daru BH, Park DS, Primack RB, Willis CG, Barrington DS, Whitfeld TJ, Seidler TG, Sweeney PW, Foster DR, Ellison AM, et al. (2018). Widespread sampling biases in herbaria revealed from large‐scale digitization. New Phytol. 217(2): 939-955. https://doi.org/10.1111/nph.14855
Dyderski MK, Tyborski J, Jagodziński AM. 2017. The utility of ancient forest indicator species in urban environments: A case study from Poznań, Poland. Urban For Urban Gree. 27: 76-83. https://doi.org/10.1016/j.ufug.2017.06.016
Global Biodiversity Information Facility database. 2001- .GBIF [accessed 2022 Apr 25]. https/www.GBIF.org.
Glaves P, Handley C , Birkbeck J, Rotherham ID, Wright B. 2009. A survey of the coverage, use and application of ancient woodland indicator lists in the UK. Hallam Environmental Consultants Ltd., Biodiversity and Landscape History Research Institute/Geography, Tourism and Environment Change Research Unit, Sheffield Hallam University, Sheffield. Available from: https://core.ac.uk/download/pdf/4149223.pdf
Greve M, Lykke AM, Fagg CW, Gereau RE, Lewis GP, Marchant R, Marshall AR, Ndaysishimiye J, Bogaert J, Svenning JC. 2016. Realising the potential of herbarium records for conservation biology. S Afr J Bot. 105: 317-323. https://doi.org/10.1016/j.sajb.2016.03.017
Heberling, JM. (2022). Herbaria as big data sources of plant traits. Int J Plant Sci. 183(2): 87-118. https://doi.org/10.1086/717623
Hedrick BP, Heberling JM, Meineke EK, Turner KG, Grassa CJ, Park DS, Kennedy J, Clarke JA, Cook JA, Blackburn DC. 2020. Digitization and the future of natural history collections. BioScience. 70(3): 243-251. https://doi.org/10.1093/biosci/biz163
Herbaria United database. 2006 - . Botanical Society of Britain and Ireland and Botanical Collection Managers Group. [accessed 2022 Apr 25]. http://herbariaunited.org/
Hermy M, Honnay O, Firbank L, Grashof-Bokdam C, Lawesson JE. 1999. An ecological comparison between ancient and other forest plant species of Europe, and the implications for forest conservation. Biol Conserv. 91(1): 9-22. https://doi.org/10.1016/S0006-3207(99)00045-2
HM Government. 2018. A green future: our 25 year plan to improve the environment. London: Department for Environment, Fisheries and Rural Affairs. Available from: https://www. gov. uk/government/publications/25-year-environment-plan.
James SA, Soltis PS, Belbin L, Chapman AD, Nelson G, Paul DL, Collins M. (2018). Herbarium data: Global biodiversity and societal botanical needs for novel research. App Plant Sci, 6(2), e1024. https://doi.org/10.1002/aps3.1024
Kelemen K, Kriván A, Standovár T. 2014. Effects of land‐use history and current management on ancient woodland herbs in Western Hungary. J Veg Sci. 25(1): 172-183. https://doi.org/10.1111/jvs.12046
Kirby K. 2004. Table of ancient woodland indicator plants. In: Rose F. O'Reilly C, Smith D.P. and Collings. M, editors. The Wild Flower Key: How to Identify Wild Flowers, Trees and Shrubs in Britain and Ireland. London, UK: Warne; p. 558-561
Kricsfalusy VV, Trevisan N. (2014). Prioritizing regionally rare plant species for conservation using herbarium data. Biodivers Conserv. 23(1): 39-61. https://doi.org/10.1007/s10531-013-0583-4
Lang PL, Willems FM, Scheepens JF, Burbano HA, Bossdorf, O. 2019. Using herbaria to study global environmental change. New Phytol. 221(1): 110-122. https://doi.org/10.1111/nph.15401
Lavoie C. 2013. Biological collections in an ever changing world: Herbaria as tools for biogeographical and environmental studies. Perspect Plant Ecol. 15(1), 68-76. https://doi.org/10.1016/j.ppees.2012.10.002
Lienert J, Fischer M, Diemer M. 2002. Local extinctions of the wetland specialist Swertia perennis L.(Gentianaceae) in Switzerland: a revisitation study based on herbarium records. Biol Conserv. 103(1): 65-76. https://doi.org/10.1016/S0006-3207(01)00121-5
Le Bras G, Pignal M, Jeanson ML, Muller S, Aupic C, Carré B, Flament G, Gaudeul M, Gonçalves C, Invernón VR et al. 2017) The French Muséum national d’histoire naturelle vascular plant herbarium collection dataset. Sci data. 4(1): 1-16. https://doi.org/10.1038/sdata.2017.16
Marsico TD, Krimmel ER, Carter JR, Gillespie EL, Lowe PD, McCauley R, Morris AB, Nelson G, Smith M, Soteropoulos D, et al. 2020. Small herbaria contribute unique biogeographic records to county, locality, and temporal scales. Am J Bot. 107(11): 1577-1587. https://doi.org/10.1002/ajb2.1563
Martine CT, Ward ME. 2013. Establishment of regional herbarium leads to more than 200 new flora atlas records for New York State. J Torrey Bot Soc. 140(1): 125-131. https://doi.org/10.3159/TORREY-D-12-00036.1
McMullin RT, Wiersma YF. 2019. Out with OLD growth, in with ecological continNEWity: new perspectives on forest conservation. Front Ecol Environ. 17(3): 176-181. https://doi.org/10.1002/fee.2016
Meineke EK, Davis CC, Davies TJ. 2018. The unrealized potential of herbaria for global change biology. Ecol Monogr. 88(4): 505-525. https://doi.org/10.1002/ecm.1307
Ministry of Housing, Communities and Local Government. 2019. National planning policy framework. London: Ministry of Housing, Communities and Local Government. [accessed 2022 Apr 25]. https://assets.publishing.service.gov.uk/government/uploads/system/uploads/attachment_data/file/1005759/NPPF_July_2021.pdf
Natural England (2022). Ancient woodland inventory. Open government licence. [accessed 2022 Apr 25]. https://data.gov.uk/dataset/9461f463-c363-4309-ae77-fdcd7e9df7d3/ancient-woodland-england
Palo A, Ivask M, Liira J. 2013. Biodiversity composition reflects the history of ancient semi-natural woodland and forest habitats—compilation of an indicator complex for restoration practice. Ecol Indic. 34: 336-344. https://doi.org/10.1016/j.ecolind.2013.05.020
Pescott OL, Humphrey TA, Walker KJ. 2018. A short guide to using British and Irish plant occurrence data for research. Wallingford, NERC/Centre for Ecology & Hydrology. Available from: http://nora.nerc.ac.uk/id/eprint/520609/
Peterken GF. 1974. A method for assessing woodland flora for conservation using indicator species. Biol Conserv. 6(4): 239-245. https://doi.org/10.1016/0006-3207(74)90001-9
Rackham O. 2008. Ancient woodlands: modern threats. New Phytol. 180(3): 571-586. http://www.jstor.org/stable/25150608.
Razzaque J, Lester C. 2021. Why Protect Ancient Woodland in the UK? Rethinking the Ecosystem Approach. Transnatl Env La. 10(1): 135-158. https://doi.org/10.1017/S2047102520000333
Reid C, Hornigold K, McHenry E, Nichols C, Townsend M, Lewthwaite K, Elliot M, Pullinger R, Hotchkiss A, Gilmartin E, et al. 2021. State of the UK's Woods and Trees 2021.Grantham: The Woodland Trust.
Rose F. 1999. Indicators of ancient woodland-the use of vascular plants in evaluating ancient woods for nature conservation. Br Wild. 10: 241-251.
Rotherham ID. 2011. A landscape history approach to the assessment of ancient woodlands. In, Wallace EB, editor. Woodlands: ecology, management and conservation,. New York: Nova Science; p. 161-184.
Stace C. 2019. New Flora of the British Isles (4th ed). UK: C&M Floristics.
Swallow KA, Wood MJ, Goodenough AE. 2020. Relative contribution of ancient woodland indicator and non‐indicator species to herb layer distinctiveness in ancient semi‐natural, ancient replanted, and recent woodland. Appl Veg Sci. 23(4): 471-481.
https://doi.org/10.1111/avsc.12501
Troudet J, Grandcolas P, Blin A, Vignes-Lebbe R, Legendre F. 2017. Taxonomic bias in biodiversity data and societal preferences. Sci Rep. 7(1): 1-14.
https://doi.org/10.1038/s41598-017-09084-6
Walker K, Pearman D, Ellis B, McIntosh J, Lockton A. 2010. Recording the British and Irish flora 2010-2020. London: Botanical Society Britain and Ireland.
Willis KJ, Araújo MB, Bennett KD, Figueroa-Rangel B, Froyd CA, Myers N. 2007. How can a knowledge of the past help to conserve the future? Biodiversity conservation and the relevance of long-term ecological studies. Philos Trans R Soc Lond B Bio Sci. 362(1478): 175-187. https://doi.org/10.1098/rstb.2006.1977
Webb JC, Goodenough AE. 2018. Questioning the reliability of “ancient” woodland indicators: Resilience to interruptions and persistence following deforestation. Ecol Indic. 84: 354-363. https://doi.org/10.1016/j.ecolind.2017.09.010
Wulf M. 1997. Plant species as indicators of ancient woodland in northwestern Germany. J Veg Sci. 8(5): 635-642. https://doi.org/10.2307/3237367