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Temperature Thresholds of Physically Dormant Seeds and Plant Functional Response to Fire: Variation Among Species and Relative Impact of Climate Change

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Temperature Thresholds of Physically Dormant Seeds and Plant Functional Response to Fire: Variation Among Species and Relative Impact of Climate Change University of Wollongong Research Online Faculty of Science, Medicine and Health - Papers: part A Faculty of Science, Medicine and Health 1-1-2014 Temperature thresholds of physically dormant seeds and plant functional response to fire: variation among species and relative impact of climate change Mark K. J Ooi University of Wollongong, [email protected] Andrew J. Denham University of Wollongong, [email protected] Victor M. Santana University of Liverpool Tony D. Auld University of Wollongong, [email protected] Follow this and additional works at: https://ro.uow.edu.au/smhpapers Part of the Medicine and Health Sciences Commons, and the Social and Behavioral Sciences Commons Recommended Citation Ooi, Mark K. J; Denham, Andrew J.; Santana, Victor M.; and Auld, Tony D., "Temperature thresholds of physically dormant seeds and plant functional response to fire: variation among species and relative impact of climate change" (2014). Faculty of Science, Medicine and Health - Papers: part A. 1601. https://ro.uow.edu.au/smhpapers/1601 Research Online is the open access institutional repository for the University of Wollongong. For further information contact the UOW Library: [email protected] Temperature thresholds of physically dormant seeds and plant functional response to fire: variation among species and relative impact of climate change Abstract Variation in dormancy thresholds among species is rarely studied but may provide a basis to better understand the mechanisms controlling population persistence. Incorporating dormancy-breaking temperature thresholds into existing trait frameworks could improve predictions regarding seed bank persistence, and subsequently species resilience in response to fire, climate change and anthropogenic management. A key ecological strategy for many species from fire-prone ecosystems is the possession of a long-lived seed bank, ensuring recovery after fire. Physical dormancy is dominant in these ecosystems and maintaining this dormancy is directly linked to seed bank persistence. We identified a suite of seed-related factors relevant to maintaining populations in fire-prone regions for 14 co-occurring physically dormant species. We measured variation in initial levels of dormancy and then applied experimental heating treatments, based on current seasonal temperatures and those occurring during fires, to seeds of all study species. Additionally, higher seasonal temperature treatments were applied to assess response of seeds to temperatures projected under future climate scenarios. Levels of germination response and mortality were determined to assess how tightly germination response was bound to either fire or seasonal cues. Six species were found to have dormancy cues bound to temperatures that only occur during fires (80°C and above) and were grouped as having obligate pyrogenic dormancy release. The remaining species, classified as having facultative pyrogenic dormancy, had lower temperature dormancy thresholds and committed at least 30% of seeds to germinate after summer-temperature treatments. Evidence from this study supports including dormancy-breaking temperature thresholds as an attribute for identifying functional types. High temperature thresholds for breaking dormancy, found in our obligate pyrogenic group, appear to be a fire-adapted trait, while we predict that species in the facultative group are most at risk to increased seed bank decay resulting from elevated soil temperatures under projected climate change. Keywords Bet-hedging, Fabaceae, fire adaptation, fire management, germination, hard-seeded, heat shock, physical dormancy, seed bank, seedling emergence Disciplines Medicine and Health Sciences | Social and Behavioral Sciences Publication Details Ooi, M. K. J., Denham, A. J., Santana, V. M. & Auld, T. D. (2014). Temperature thresholds of physically dormant seeds and plant functional response to fire: variation among species and relative impact of climate change. Ecology and Evolution, 4 (5), 656-671. This journal article is available at Research Online: https://ro.uow.edu.au/smhpapers/1601 Temperature thresholds of physically dormant seeds and plant functional response to fire: variation among species and relative impact of climate change Mark K. J. Ooi1, Andrew J. Denham1,2, Victor M. Santana3,4 & Tony D. Auld1,2 1Institute for Conservation Biology & Environmental Management, School of Biological Sciences, University of Wollongong, Wollongong, New South Wales 2522, Australia 2Office of Environment & Heritage, P.O. Box 1967, Hurstville, New South Wales 2220, Australia 3School of Environmental Sciences, University of Liverpool, Liverpool L69 3GP, U.K. 4Fundacio´ n de la Generalitat Valenciana Centro de Estudios Ambientales del Mediterraneo (CEAM), Parque Tecnolo´ gico Paterna. C/ Charles Darwin, 14, E-46980 Paterna, Valencia, Spain Keywords Abstract Bet-hedging, Fabaceae, fire adaptation, fire management, germination, hard-seeded, Variation in dormancy thresholds among species is rarely studied but may pro- heat shock, physical dormancy, seed bank, vide a basis to better understand the mechanisms controlling population persis- seedling emergence. tence. Incorporating dormancy-breaking temperature thresholds into existing trait frameworks could improve predictions regarding seed bank persistence, Correspondence and subsequently species resilience in response to fire, climate change and Mark Ooi, Institute for Conservation Biology, anthropogenic management. A key ecological strategy for many species from University of Wollongong, Wollongong, NSW fire-prone ecosystems is the possession of a long-lived seed bank, ensuring 2522, Australia. Tel: +61 (0)2 4252 8209; Fax: +61 (0)2 4221 4135; recovery after fire. Physical dormancy is dominant in these ecosystems and E-mail: [email protected] maintaining this dormancy is directly linked to seed bank persistence. We iden- tified a suite of seed-related factors relevant to maintaining populations in fire- Funding Information prone regions for 14 co-occurring physically dormant species. We measured An Australian Research Council (ARC) variation in initial levels of dormancy and then applied experimental heating Linkage Grant (LP110100527) between the treatments, based on current seasonal temperatures and those occurring during University of Wollongong and the NSW fires, to seeds of all study species. Additionally, higher seasonal temperature Office of Environment and Heritage provided the funding for this research. treatments were applied to assess response of seeds to temperatures projected under future climate scenarios. Levels of germination response and mortality Received: 12 December 2013; Revised: 14 were determined to assess how tightly germination response was bound to January 2014; Accepted: 14 January 2014 either fire or seasonal cues. Six species were found to have dormancy cues bound to temperatures that only occur during fires (80°C and above) and were Ecology and Evolution 2014; 4(5): grouped as having obligate pyrogenic dormancy release. The remaining species, 656–671 classified as having facultative pyrogenic dormancy, had lower temperature dor- doi: 10.1002/ece3.973 mancy thresholds and committed at least 30% of seeds to germinate after sum- mer-temperature treatments. Evidence from this study supports including dormancy-breaking temperature thresholds as an attribute for identifying functional types. High temperature thresholds for breaking dormancy, found in our obligate pyrogenic group, appear to be a fire-adapted trait, while we predict that species in the facultative group are most at risk to increased seed bank decay resulting from elevated soil temperatures under projected climate change. Introduction ecosystem over time (Harper 1977; Noble and Slatyer 1980; Woodward 1987). For plant communities from fire- The interaction between critical life-history stages and the prone regions, species can persist using two broad func- environment can determine the relative success, distribu- tional approaches: resprouting and recruitment from seed tion, and level of persistence of species within an (Pausas et al. 2004). For the latter group in particular, 656 ª 2014 The Authors. Ecology and Evolution published by John Wiley & Sons Ltd. This is an open access article under the terms of the Creative Commons Attribution License, which permits use, distribution and reproduction in any medium, provided the original work is properly cited. M. K. J. Ooi et al. Physical Dormancy and Functional Response the establishment phase after disturbance is important very likely. Additionally, measuring only maximal germi- due to the small window of opportunity for successful nation may not be a robust assessment because many spe- recruitment to take place and the need to replace individ- cies respond to seasonal temperatures by committing only uals killed by fire (Whelan 1995). Thus, a critical ecologi- a small proportion of seeds to germinate (e.g., Auld 1995; cal strategy for many species from such ecosystems is the Ooi et al. 2009). Determining the extent of variation to a possession of a long-lived seed bank. Seed banks permit range of seasonal and fire temperatures could provide key long-term persistence for a species when environmental information for understanding how seed banks function or biotic conditions are unfavorable for other life stages and how species coexist. (Grime 1989; Fenner and Thompson 2005), as well as a Grouping species by functional traits allows predictions bet-hedging capacity (Philippi 1993; Ooi et al. 2009). for large numbers of
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