Experimental seed sowing reveals seedling recruitment vulnerability to unseasonal fire.
Banksia woodlands
fire regime
fire season
germination
plant demography
prescribed burning
species functional traits
survival analysis
Journal
Ecological applications : a publication of the Ecological Society of America
ISSN: 1051-0761
Titre abrégé: Ecol Appl
Pays: United States
ID NLM: 9889808
Informations de publication
Date de publication:
10 2021
10 2021
Historique:
revised:
25
02
2021
received:
17
06
2020
accepted:
22
03
2021
pubmed:
14
7
2021
medline:
21
10
2021
entrez:
13
7
2021
Statut:
ppublish
Résumé
Unseasonal fire occurrence is increasing globally, driven by climate change and other human activity. Changed timing of fire can inhibit postfire seedling recruitment through interactions with plant phenology (the timing of key processes, e.g., flower initiation, seed production, dispersal, germination), and therefore threaten the persistence of many plant species. Although empirical evidence from winter-rainfall ecosystems shows that optimal seedling recruitment is expected following summer and autumn (dry season) fires, we sought experimental evidence isolating the mechanisms of poor recruitment following unseasonal (wet season) fire. We implemented a seed-sowing experiment using nine species native to fire-prone, Mediterranean-climate woodlands in southwestern Australia to emulate the timing of postfire recruitment and test key mechanisms of fire seasonality effects. For seeds sown during months when fire is unseasonal (i.e., August-September: end of the wet winter season), seedling recruitment was reduced by up to 99% relative to seeds sown during seasonal fire months (i.e., May-June: end of the dry summer season) because of varying seed persistence, seedling emergence, and seedling survival. We found that up to 70 times more seedlings emerged when seeds were sown during seasonal fire months compared to when seeds were sown during unseasonal fire months. The few seedlings that emerged from unseasonal sowings all died with the onset of the dry season. Of the seeds that failed to germinate from unseasonal sowings, only 2% survived exposure on the soil surface over the ensuing hot and dry summer. Our experimental results demonstrate the potential for unseasonal fire to inhibit seedling recruitment via impacts on pregermination seed persistence and seedling establishment. As ongoing climate change lengthens fire seasons (i.e., unseasonal wildfires become more common) and managed fires are implemented further outside historically typical fire seasons, postfire seedling recruitment may become more vulnerable to failure, causing shifts in plant community composition towards those with fewer species solely dependent on seeds for regeneration.
Banques de données
Dryad
['10.5061/dryad.qrfj6q5fb']
Types de publication
Journal Article
Research Support, Non-U.S. Gov't
Langues
eng
Sous-ensembles de citation
IM
Pagination
e02411Informations de copyright
© 2021 by the Ecological Society of America.
Références
Andrys, J., J. Kala, and T. J. Lyons. 2017. Regional climate projections of mean and extreme climate for the southwest of Western Australia (1970-1999 compared to 2030-2059). Climate Dynamics 48:1723-1747.
Australian Bureau of Meteorology. 2020. Climate data online [Online]. http://www.bom.gov.au/climate/data/
Balch, J. K., B. A. Bradley, J. T. Abatzoglou, R. C. Nagy, E. J. Fusco, and A. L. Mahood. 2017. Human-started wildfires expand the fire niche across the United States. Proceedings of the National Academy of Sciences of the United States of America 114:2946-2951.
Barton, K. 2019. MuMIn: multi-model inference. https://CRAN.R-project.org/package=MuMIn
Bates, B. C., P. Hope, B. Ryan, I. Smith, and S. Charles. 2008. Key findings from the Indian Ocean Climate Initiative and their impact on policy development in Australia. Climatic Change 89:339-354.
Bates, D., M. Maechler, B. Bolker, and S. Walker. 2015. Fitting linear mixed-effects models using lme4. Journal of Statistical Software 67:1-48.
Bond, W. J. 1984. Fire survival of Cape Proteaceae-influence of fire season and seed predators. Vegetatio 56:65-74.
Bond, W. J., J. Vlok, and M. Viviers. 1984. Variation in seedling recruitment of Cape Proteaceae after fire. Journal of Ecology 72:209-221.
Bowman, D., et al. 2011. The human dimension of fire regimes on Earth. Journal of Biogeography 38:2223-2236.
Bradstock, R. A., and M. Bedward. 1992. Simulation of the effect of season of fire on post-fire seedling emergence of two species based on long-term rainfall records. Australian Journal of Botany 40:75-88.
Buma, B., C. D. Brown, D. C. Donato, J. B. Fontaine, and J. F. Johnstone. 2013. The impacts of changing disturbance regimes on serotinous plant populations and communities. BioScience 63:866-876.
Burrows, N., B. Ward, A. Wills, M. Williams, and R. Cranfield. 2019. Fine-scale temporal turnover of jarrah forest understory vegetation assemblages is independent of fire regime. Fire Ecology 15:10.
Céspedes, B., B. Luna, B. Perez, I. R. Urbieta, and J. M. Moreno. 2014. Burning season effects on the short-term post-fire vegetation dynamics of a Mediterranean heathland. Applied Vegetation Science 17:86-96.
Céspedes, B., I. Torres, B. Luna, B. Pérez, and J. M. Moreno. 2012. Soil seed bank, fire season, and temporal patterns of germination in a seeder-dominated Mediterranean shrubland. Plant Ecology 213:383-393.
Clarke, P. J., K. J. E. Knox, and D. Butler. 2010. Fire intensity, serotiny and seed release in 19 woody species: evidence for risk spreading among wind-dispersed and resprouting syndromes. Australian Journal of Botany 58:629-636.
Clarke, P. J., M. J. Lawes, J. J. Midgley, B. Lamont Byron, F. Ojeda, G. E. Burrows, N. J. Enright, and K. E. J. Knox. 2013. Resprouting as a key functional trait: How buds, protection and resources drive persistence after fire. New Phytologist 197:19-35.
Commander, L. E., L. Merino-Martín, C. P. Elliott, B. P. Miller, K. Dixon, and J. Stevens. 2019. Demographic, seed and microsite limitations to seedling recruitment in semi-arid mine site restoration. Plant and Soil 457:113-129.
Cowling, R. M., and B. B. Lamont. 1985a. Seed release in Banksia: the role of wet-dry cycles. Australian Journal of Ecology 10:169-171.
Cowling, R. M., and B. B. Lamont. 1985b. Variation in serotiny of three Banksia species along a climatic gradient. Australian Journal of Ecology 10:345-350.
Cowling, R. M., and B. B. Lamont. 1987. Post-fire recruitment of four co-occurring Banksia species. Journal of Applied Ecology 24:645-658.
Densmore, V. S., and E. S. Clingan. 2019. Prescribed burning in a Mediterranean-climate region mitigates the disturbance by bushfire to a critical food resource for an endangered bird, the Carnaby’s cockatoo. Fire Ecology 15:36.
Dunker, B., C. M. Bull, D. A. Keith, and D. A. Driscoll. 2019. Season of fire influences seed dispersal by wind in a serotinous obligate seeding tree. Plant Ecology 220:405-416.
Enright, N. J., J. B. Fontaine, D. Bowman, R. A. Bradstock, and R. J. Williams. 2015. Interval squeeze: altered fire regimes and demographic responses interact to threaten woody species persistence as climate changes. Frontiers in Ecology and the Environment 13:265-272.
Enright, N. J., J. B. Fontaine, B. B. Lamont, B. P. Miller, and V. C. Westcott. 2014. Resistance and resilience to changing climate and fire regime depend on plant functional traits. Journal of Ecology 102:1572-1581.
Enright, N. J., and B. B. Lamont. 1989. Seed banks, fire season, safe sites and seedling recruitment in fire co-occurring Banksia species. Journal of Ecology 77:1111-1122.
Enright, N. J., and B. B. Lamont. 1992. Survival, growth and water relations of Banksia seedlings on a sand mine rehabilitation site and adjacent scrub-heath sites. Journal of Applied Ecology 29:663-671.
Frazer, J. M., and S. D. Davis. 1988. Differential survival of chaparral seedlings during the first summer drought after wildfire. Oecologia 76:215-221.
Groom, P., and B. Lamont. 2015. Plant life of southwestern Australia. De Gruyter Open, Warsaw, Poland.
Harrison, X. A. 2015. A comparison of observation-level random effect and beta-binomial models for modelling overdispersion in binomial data in ecology & evolution. PeerJ 3:e1114.
Heelemann, S., S. Proches, A. G. Rebelo, B. W. Van Wilgen, S. Porembski, and R. M. Cowling. 2008. Fire season effects on the recruitment of non-sprouting serotinous Proteaceae in the eastern (bimodal rainfall) fynbos biome, South Africa. Austral Ecology 33:119-127.
Hidayati, S. N., D. J. Merritt, S. R. Turner, K. W. Dixon, and J. L. Walck. 2019. Temporal dynamics of seedling emergence among four fire ephemerals: the interplay of after-ripening and embryo growth with smoke. Seed Science Research 29:104-114.
Holden, Z. A., P. Morgan, M. A. Crimmins, R. K. Steinhorst, and A. M. S. Smith. 2007. Fire season precipitation variability influences fire extent and severity in a large southwestern wilderness area, United States. Geophysical Research Letters 34:L16708.
Hothorn, T., F. Bretz, and P. Westfall. 2008. Simultaneous inference in general parametric models. Biometrical Journal 50:346-363.
Hurvich, C. M., and C.-L. Tsai. 1989. Regression and time series model selection in small samples. Biometrika 76:297-307.
Jasinge, N. U., T. Huynh, and A. C. Lawrie. 2018. Consequences of season of prescribed burning on two spring-flowering terrestrial orchids and their endophytic fungi. Australian Journal of Botany 66:298-312.
Kassambara, A., and M. Kosinski. 2018. survminer: drawing survival curves using ‘ggplot2’. https://CRAN.R-project.org/package=survminer
Keeley, J. E., and A. D. Syphard. 2018. Historical patterns of wildfire ignition sources in California ecosystems. International Journal of Wildland Fire 27:781-799.
Keith, D. A., B. Dunker, and D. A. Driscoll. 2020. Dispersal: the eighth fire seasonality effect on plants. Trends in Ecology & Evolution 35:305-307.
Knapp, E. E., J. E. Keeley, E. A. Ballenger, and T. J. Brennan. 2005. Fuel reduction and coarse woody debris dynamics with early season and late season prescribed fire in a Sierra Nevada mixed conifer forest. Forest Ecology and Management 208:383-397.
Knox, K. J. E., and P. J. Clarke. 2006. Fire season and intensity affect shrub recruitment in temperate sclerophyllous woodlands. Oecologia 149:730-739.
Kraaij, T., R. M. Cowling, B. Van Wilgen, D. R. Rikhotso, and M. Difford. 2017. Vegetation responses to season of fire in an aseasonal, fire-prone fynbos shrubland. PeerJ 5;e3591.
Lamont, B. B., E. T. F. Witkowski, and N. J. Enright. 1993. Post-fire litter microsites: safe for seeds, unsafe for seedlings. Ecology 74:501-512.
Le Page, Y., D. Oom, J. M. N. Silva, P. Jönsson, and J. M. C. Pereira. 2010. Seasonality of vegetation fires as modified by human action: observing the deviation from eco-climatic fire regimes. Global Ecology and Biogeography 19:575-588.
Mackenzie, B. D. E., T. D. Auld, D. A. Keith, F. K. C. Hui, and M. K. J. Ooi. 2016. The effect of seasonal ambient temperatures on fire-stimulated germination of species with physiological dormancy: a case study using Boronia (Rutaceae). PLoS One 11:e0156142.
Merino-Martín, L., C. Courtauld, L. Commander, S. Turner, W. Lewandrowski, and J. Stevens. 2017. Interactions between seed functional traits and burial depth regulate germination and seedling emergence under water stress in species from semi-arid environments. Journal of Arid Environments 147:25-33.
Merritt, D. J., S. R. Turner, S. Clarke, and K. W. Dixon. 2007. Seed dormancy and germination stimulation syndromes for Australian temperate species. Australian Journal of Botany 55:336-344.
Midgley, J. J. 1989. Season of burn of serotinous fynbos Proteaceae: a critical review and further data. South African Journal of Botany 55:165-170.
Midgley, J. J., T. Hoekstra, and R. Bartholomew. 1989. The effects of date of planting on field establishment of serotinous cape Proteaceae. Vegetatio 79:185-192.
Miller, R., J. Fontaine, D. Merritt, B. Miller, and N. Enright. 2021. Experimental seed sowing reveals seedling recruitment vulnerability to unseasonal fire. Dryad, data set. https://doi.org/10.5061/dryad.qrfj6q5fb
Miller, R. G., R. Tangney, N. J. Enright, J. B. Fontaine, D. J. Merritt, M. K. J. Ooi, K. X. Ruthrof, and B. P. Miller. 2019. Mechanisms of fire seasonality effects on plant populations. Trends in Ecology & Evolution 34:1104-1117.
Miller, R. G., R. Tangney, N. J. Enright, J. B. Fontaine, D. J. Merritt, M. K. J. Ooi, K. X. Ruthrof, and B. P. Miller. 2020. Fire seasonality mechanisms are fundamental for understanding broader fire regime effects. Trends in Ecology & Evolution 35:869-871.
Mustart, P. J., A. G. Rebelo, J. Juritz, and R. M. Cowling. 2012. Wide variation in post-emergence desiccation tolerance of seedlings of fynbos proteoid shrubs. South African Journal of Botany 80:110-117.
Myers, N., R. A. Mittermeier, C. G. Mittermeier, G. A. B. da Fonseca, and J. Kent. 2000. Biodiversity hotspots for conservation priorities. Nature 403:853-858.
Nakagawa, S., and H. Schielzeth. 2013. A general and simple method for obtaining R2 from generalized linear mixed-effects models. Methods in Ecology and Evolution 4:133-142.
Nield, A. P., N. J. Enright, and P. G. Ladd. 2016. Fire-stimulated reproduction in the resprouting, non-serotinous conifer Podocarpus drouynianus (Podocarpaceae): The impact of a changing fire regime. Population Ecology 58:179-187.
Ooi, M. K. J. 2010. Delayed emergence and post-fire recruitment success: effects of seasonal germination, fire season and dormancy type. Australian Journal of Botany 58:248-256.
Ooi, M. K. J. 2019. The importance of fire season when managing threatened plant species: A long-term case-study of a rare Leucopogon species (Ericaceae). Journal of Environmental Management 236:17-24.
Ooi, M. K. J., T. D. Auld, and R. J. Whelan. 2004. Delayed post-fire seedling emergence linked to season: a case study with Leucopogon species (Epacridaceae). Plant Ecology 174:183-196.
Palmer, H. D., A. J. Denham, and M. K. J. Ooi. 2018. Fire severity drives variation in post-fire recruitment and residual seed bank size of Acacia species. Plant Ecology 219:527-537.
Plucinski, M. P. 2014. The timing of vegetation fire occurrence in a human landscape. Fire Safety Journal 67:42-52.
Potts, J. B., E. Marino, and S. L. Stephens. 2010. Chaparral shrub recovery after fuel reduction: a comparison of prescribed fire and mastication techniques. Plant Ecology 210:303-315.
Pratt, R. B., A. L. Jacobsen, R. Mohla, F. W. Ewers, and S. D. Davis. 2008. Linkage between water stress tolerance and life history type in seedlings of nine chaparral species (Rhamnaceae). Journal of Ecology 96:1252-1265.
R Development Core Team. 2018. R: a language and environment for statistical computing. R Foundation for Statistical Computing, Vienna, Austria. www.R-project.org
Ramalho, C. E., E. Laliberté, P. Poot, and R. J. Hobbs. 2014. Complex effects of fragmentation on remnant woodland plant communities of a rapidly urbanizing biodiversity hotspot. Ecology 95:2466-2478.
Ritchie, A. L., et al. 2021. A threatened ecological community: research advances and priorities for Banksia woodlands. Australian Journal of Botany 69:53-84.
Roche, S., K. W. Dixon, and J. S. Pate. 1998. For everything a season: smoke-induced germination and seedling recruitment in a Western Australian Banksia woodland. Australian Journal of Ecology 23:111-120.
Ryan, K. C., E. E. Knapp, and J. M. Varner. 2013. Prescribed fire in North American forests and woodlands: History, current practice, and challenges. Frontiers in Ecology and the Environment 11:e15-e24.
Tangney, R., D. J. Merritt, J. B. Fontaine, and B. P. Miller. 2019. Seed moisture content as a primary trait regulating the lethal temperature thresholds of seeds. Journal of Ecology 107:1093-1105.
Tangney, R., R. G. Miller, N. J. Enright, J. B. Fontaine, D. J. Merritt, M. K. J. Ooi, K. X. Ruthrof, and B. P. Miller. 2020. Seed dormancy interacts with fire seasonality mechanisms. Trends in Ecology & Evolution 35:1057-1059.
Turner, S. R., B. Pearce, D. P. Rokich, R. R. Dunn, D. J. Merritt, J. D. Majer, and K. W. Dixon. 2006. Influence of polymer seed coatings, soil raking, and time of sowing on seedling performance in post-mining restoration. Restoration Ecology 14:267-277.
Urretavizcaya, M. F., G. E. Defossé, and H. E. Gonda. 2012. Effect of sowing season, plant cover, and climatic variability on seedling emergence and survival in burned Austrocedrus chilensis forests. Restoration Ecology 20:131-140.
Verdú, M., and A. Traveset. 2005. Early emergence enhances plant fitness: a phylogenetically controlled meta-analysis. Ecology 86:1385-1394.
Westerling, A. L., H. G. Hidalgo, D. R. Cayan, and T. W. Swetnam. 2006. Warming and earlier spring increase western U.S. forest wildfire activity. Science 313:940-943.
Whelan, R. J., and J. York. 1998. Post-fire germination of Hakea sericea and Petrophile sessilis after spring burning. Australian Journal of Botany 46:367-376.
Wickham, H. 2009. ggplot2: elegant graphics for data analysis. Springer-Verlag, New York, New York, USA.
Wilson, B. A., J. Kuehs, L. E. Valentine, T. Sonneman, and K. M. Wolfe. 2014. Guidelines for ecological burning regimes in Mediterranean ecosystems: a case study in Banksia woodlands in Western Australia. Pacific Conservation Biology 20:57-74.
Zuur, A. F., E. N. Ieno, and C. S. Elphick. 2010. A protocol for data exploration to avoid common statistical problems. Methods in Ecology and Evolution 1:3-14.