Resprouting plants are more resilient to drought: increased fitness in Anthyllis cytisoides

Resprouting plants are more resilient to drought: increased fitness in Anthyllis cytisoides | Authorea try { document.documentElement.classList.add('js'); } catch (e) { } var _gaq = _gaq || []; _gaq.push(['_setAccount', 'G-8VDV14Y67G']); _gaq.push(['_trackPageview']); (function() { var ga = document.createElement('script'); ga.type = 'text/javascript'; ga.async = true; ga.src = ('https:' == document.location.protocol ? 'https://ssl' : 'http://www') + '.google-analytics.com/ga.js'; var s = document.getElementsByTagName('script')[0]; s.parentNode.insertBefore(ga, s); })(); Skip to main content Preprints Collections Wiley Open Research IET Open Research Ecological Society of Japan All Collections About About Authorea FAQs Contact Us Quick Search anywhere Search for preprint articles, keywords, etc. Search Search ADVANCED SEARCH SCROLL Oikos This is a preprint and has not been peer reviewed. Data may be preliminary. 29 July 2025 V1 Latest version Share on Resprouting plants are more resilient to drought: increased fitness in Anthyllis cytisoides Authors : Jaime Saiz Blanco 0009-0002-2947-1634 , Conchita Alonso 0000-0002-7418-3204 , and Juli Pausas [email protected] Authors Info & Affiliations https://doi.org/10.22541/au.175379050.00325607/v1 282 views 198 downloads Contents Abstract Information & Authors Metrics & Citations View Options References Figures Tables Media Share Abstract Fire and drought increasingly co-occur, exposing plants to greater drought stress during post-fire resprouting. Yet, the effect of this combination of disturbances on plant fitness remains poorly understood. Here, we examine how post-fire resprouting influences reproductive success under drought conditions in the Mediterranean shrub Anthyllis cytisoides. We took advantage of a natural setting where, following a high-intensity wildfire, burned (and resprouting) individuals coexisted with unburned ones. We compared flowering and fruiting performance under a postfire drought in both burned and unburned plants. Our results show that burned individuals were significantly less affected by drought: Compared to their unburned counterparts, they had a higher probability of flowering, produced more flowers, set more fruits and seeds, and produced heavier seeds. We propose that reduced competition and enhanced physiological condition following fire may temporarily boost plant fitness despite environmental stress. These findings suggest that post-fire resprouting may confer a short-term reproductive advantage during drought (“post-fire advantage window”). Understanding this dynamic is essential to predict plant responses under compound disturbance regimes in fire-prone ecosystems. Introduction As the world becomes drier (IPCC, 2023) and fire regimes shift (Pausas & Keeley, 2021), the likelihood that plants will be subject to compound disturbances increases (Batllori et al., 2019; Kane et al., 2017; Nolan et al., 2021). However, how fire and drought interact to affect plants remains unclear. Most studies focus exclusively on trees (Cansler et al., 2024; Hanson et al., 2024; van Mantgem et al., 2016; Valor et al., 2020; Zald et al., 2022) or on the recruitment limitations imposed by postfire drought (Moya et al., 2015; Young et al., 2019), while fewer consider post-fire resprouting (Moore et al., 2019; Parra & Moreno, 2017, 2018; Pratt et al., 2014; Schwilk et al., 2016), which is one of the primary plant adaptations to wildfires (Keeley & Pausas, 2022). While low water availability may limit post-fire resprouting (Nolan et al., 2021), experimental evidence remains mixed. On the one hand, resprouting relies on stored reserves that may take years to replenish (Canadell & López‐Soria, 1998; DeSouza et al., 1986), making resprouts vulnerable to carbon starvation when disturbances occur within a short interval (Canadell & López‐Soria, 1998). Moreover, hydraulic failure might also happen when resprout initiates during a drought event, as the tissues of these new shoots appear to be more vulnerable to cavitation than those of mature plants (Jacobsen et al., 2016; Ramirez et al., 2012; Schwilk et al., 2016). Indeed, in the Californian chaparral, high post-fire mortality rates due to cavitation and carbon starvation have been observed when a wildfire coincided with an extremely severe drought (Pratt et al., 2014). In contrast, under milder conditions such as moderate drought following prescribed burns, resprouting plants have been shown to outperform unburned neighbors and appear unaffected by water stress (Parra & Moreno, 2018; Van Blerk et al., 2021). These contrasting findings suggest that the impact of drought on resprouting plants is context-dependent, varying drastically with the intensity of drought and fire (Pausas et al., 2016), as well as with their temporal occurrence. Of the few studies on drought in resprouting plants, all focus on vegetative traits (survival, growth, or physiological traits). Yet, understanding whether resprouting plants can maintain reproduction under post-fire drought conditions is essential for predicting population persistence under changing disturbance regimes. In this study, we aim to understand the reproductive consequences of drought and fire in a resprouting plant. To do so, we studied a population of the Mediterranean shrub Anthyllis cytisoides , that was partially affected by a wildfire and subsequently experienced a severe drought in the second spring after the fire. This setting allowed us to compare burned (resprouting) and unburned individuals in close proximity. We expect that resprouting individuals will be significantly less affected by drought in terms of flower and fruit production. We propose that the reduced competition and enhanced physiological condition following fire provide resprouting plants with a recovery advantage, enabling them to better withstand future disturbances. Methods Study species Anthyllis cytisoides L. (Fabaceae) is a drought-deciduous shrub distributed across the western Mediterranean region that can reach up to 1.5 meters in height. It thrives in open shrublands and abandoned fields under dry and warm low-altitude environments (de Bolòs & Vigo 1984). It is a facultative resprouter, meaning that, after a fire, it regenerates by resprouting from the root crown and by seedling recruiting from a soil seed bank. Study area The study was conducted at the edge of a high-intensity crown fire that occurred in the summer of 2021 in Azuebar, Castellón province, eastern Spain (365 m elevation; mean annual temperature= 13.6 °C, annual precipitation= 446.4 mm). The area consisted of abandoned fields colonized by Pinus halepensis and shrubs, including A. cytisoides. The edge of the fire formed a mosaic landscape of burned and unburned patches in close proximity (the two furthest studied plants where 150m apart), offering an ideal natural experiment to compare fire-affected (burned and resprouted) and unaffected (unburned) individuals under the same climatic conditions (see Saiz-Blanco et al., under review, for more details on the study area). After fire, the first (2022) and the second (2023) postfire springs were climatically contrasted. Spring 2023 was exceptionally dry (32 mm), and warmer (15.9 °C) than spring 2022 (280 mm, 14.2 °C) and than the historical average (Figure S1, S2). Field sampling In 2022, we selected 31 A. cytisoides individuals within a 0.5 ha plot (16 burned and resprouting, and 15 unburned). We measured a range of phenotypic traits, including flower, seed, and fitness-related traits (Saiz-Blanco et al., under review). Then, in the spring of 2023, we noticed that burned plants showed more flowers than unburned plants, despite not yet being fully recovered from the fire (e.g., in terms of height or aboveground biomass). To quantify these effects, we expanded our sampling to record whether each individual produced flowers, and if so, whether it produced filled fruits (binary traits). We also recorded seed mass and seed set at the within-individual level, using several inflorescences per individual as the sampling unit. Since A. cytisoides fruits are single-seeded, we defined seed set as the proportion of fruits containing a seed. To measure these last two traits, we bagged multiple inflorescences (N ≥ 3) per individual and collected fruits during seed dispersal. Seeds were extracted by manually scrubbing hard-coated fruits with sandpaper and weighed to the nearest mg using a Mettler Toledo AX 205 balance. Additionally, in 2023, we quantified each individual’s total floral display by counting open flowers every seven days from March to May. Considering A. cytisoides flowers last approximately one week (pers. obs.) and that this period spanned the entire flowering season, we can assume that we captured the whole flower production per individual. Statistical analyses We analyzed, at individual level, floral display at peak flowering and the probability of flowering and fruiting, using (generalized) linear models. We fitted “fire” (burned/unburned plant) as the predictor for the year 2023; in 2022 we did not register total flower production and all plants flowered and set fruit. For seed set and seed weight, we used linear mixed models, with the response variable being the trait of interest, while “fire”, “year” and their interaction were included as fixed predictors. Observations were nested within individuals and included as a random factor. Model analyses were conducted using ‘ANOVA’ function from the ‘car’ package (Fox et al., 2024). Predictions and confidence intervals were calculated using the ‘ggeffects’ package (Lüdecke et al., 2025). (Generalized) linear models were fitted with the ‘stats’ package (R Core Team 2024) and mixed models with the ‘lme4’ package (Bates et al., 2025). Model diagnostics were assessed with the ‘DHARMa’ package (Hartig et al., 2024). Results During the drought year, flowering and fruiting rates declined by 28% and 41%, respectively, compared to the previous year in which all individuals produced flowers and filled fruits. These reductions were greater in unburned plants (Figure 1A, C; Table 1). Specifically, 93.7% of burned plants flowered (only one did not), compared to 60% of unburned plants. Burned plants also produced nearly four times more flowers than unburned plants during the drought (Figure 1B; Table 1). Among flowering individuals, 86.7% of burned plants set fruit, versus only 50% of unburned ones (Figure 1C), although this difference was only marginally significant (Table 1). Despite an overall reduction in seed set during the drought, burned plants produced more fruits containing seeds across both years (Figure 1D; Table 2). Finally, seed mass was unaffected by drought, with burned plants producing heavier seeds in both study years (Figure 1E; Table 1). Discussion None of the studied plants died during the two-year monitoring study, consistent with the drought avoidance strategy of A. cytisoides (Haase et al., 2000). Yet, the 2023 drought significantly reduced flower, fruit, and seed production. However, burned plants were less affected by the drought than unburned plants: a greater number of burned plants flowered and fruited, and those that flowered, produced more flowers, had higher seed set and produced heavier seeds. To our knowledge, this fire-mediated reproductive enhancement in resprouting shrubs is an undocumented phenomenon. Considering that burned and unburned plants are located within a 0.5 ha area, it is reasonable to assume that they experienced similar climatic conditions and that genetic differentiation between the two groups was low. Thus, the better performance of burned plants during the drought was likely due to a range of benefits following fire, including micro-environmental changes, reduced competition and increased resource availability (Caon et al., 2014; Parra & Moreno, 2017; Silva et al., 2006), architectural changes (Saiz-Blanco et al., under revision), increased root-shoot ratio (Schafer et al., 2014), and potentially reduced pressure from parasites and invertebrate herbivores (fire’s cleaning effect sensu Álvarez-Ruiz et al., 2021). At our field site, the higher water status of burned plants was evident during 2023: by mid-May, all unburned plants had begun shedding leaves (as response to water stress, Haase et al., 2000) while none of the burned plants did so. After a fire, resprouting plants may experience a vulnerable period during which additional stressors significantly reduce survival ( vulnerability gap , Fig. 2) as reported in previous experiments (Pratt et al., 2014; Twidwell et al., 2016). However, when this vulnerable period is over (i.e., once plant reserves are higher), the optimal post-fire conditions place resprouting plants in a better position to endure further disturbances than undisturbed plants. This leads to a higher growth rates and reproductive output after fire (post-fire advantage window effect; Fig. 2). Similar effects on growth has been detected in pine trees (Marañón-Jiménez et al., 2013; Zald et al., 2022) and in other Mediterranean shrubs (Parra & Moreno, 2017). Our results are consistent with the idea that fire ameliorates drought stress in fire-surviving plants, a mechanism that has been mainly considered in relation to radial tree growth response to prescribed surface fires (Alfaro-Sánchez et al., 2018; van Mantgem et al., 2016). We propose that in certain conditions (Fig. 2) this also occurs under high-intensity crown fires, and not only affects growth or the physiological state (Parra & Moreno, 2017), but also, and more importantly, the reproductive output (Fig. 1). The next step is to investigate the drivers of the magnitude and duration of this advantage window by, for instance, a long-term monitoring of resprouting individuals or by an experimental design addressing the effects of competition and watering in resprouting individuals during the initial post-fire vulnerability gap and subsequent advantage window. For example, to what extent this effect depends on a previous wet spring, or on traits such as storage capacity and the photosynthetic efficiency of resprouts (Goorman et al., 2011; Pausas et al., 2018), remains to be studied. Additionally, individual characteristics such as plant age or micro-environmental conditions need to be integrated, since they strongly influence plant responses to disturbance (Moreira et al., 2012; Zomer et al., 2025). Understanding these patterns will be essential for predicting plant responses in fire-prone ecosystems and improving fire-management under increasingly extreme climate scenarios. In conclusion, we highlight that the intensity and temporal pattern of the two stresses is key to understand plant responses. In particular, burned plants can do better than unburned ones when the drought occurs later, despite the plants aboveground biomass may not have been fully restored yet. That is, fire-induced advantage effect may temporarily enhance drought resilience, and this is not depicted in simulation models aiming to predict the fate of our ecosystems in a world where consecutive stresses are increasing. Bibliography Alfaro-Sánchez, R., Camarero, J. 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J. (2016). Does Prescribed Fire Promote Resistance to Drought in Low Elevation Forests of the Sierra Nevada, California, USA? Fire Ecology , 12 (1), Article 1. https://doi.org/10.4996/fireecology.1201013 Zald, H. S. J., Callahan, C. C., Hurteau, M. D., Goodwin, M. J., & North, M. P. (2022). Tree growth responses to extreme drought after mechanical thinning and prescribed fire in a Sierra Nevada mixed-conifer forest, USA. Forest Ecology and Management , 510 , 120107. https://doi.org/10.1016/j.foreco.2022.120107 Zomer, M.A., Moreira, B., Pausas, J. G. (2025) Pre-disturbance plant condition drives intraspecific resprouting variability in Mediterranean shrubs, Journal of Experimental Botany , eraf246, https://doi.org/10.1093/jxb/eraf246 Tables Table 1. Summary of the models fitted to assess the effects of drought on fitness using fixed effect models (F) and mixed effect models (M) depending on the response trait (first column). For “Fixed” models the only predictor was “Fire” (burned/unburned) for the dry year (2023), while for “Mixed” models, “Year” and its interaction with “Fire” were also included as fixed predictor and individual as random effect. Positive (negative) estimates indicating higher (lower) values of the trait in the burned treatment and the wet year. In all cases, the degree of freedom of the chisq-test was 1. Probability of flowering in 2023 F Fire 2.303 1.160 5.446 0.020 0.163 Total flower production in 2023 F Fire 1311.6 540.5 9677313 0.024 0.175 Probability of fruiting in 2023 F Fire 1.872 1.038 3.532 0.060 0.158 Seed set M Fire 0.124 0.050 7.025 0.008 0.512 Year -0.265 0.078 48.074 <0.001 Year *Fire -0.046 0.093 0.246 0.620 Seed mass M Fire 0.037 0.0250 4.211 0.040 0.126 Year -0.050 0.031 0.787 0.375 Year *Fire 0.048 0.036 1.714 0.191 Figures Figure 1. Model predictions (large solid symbols) and observations (small light-coloured dots) of the data analysed in 2022 and 2023; fire occurred in 2021. For flowering and fruiting, the negative effects of drought are stronger on unburned plants (panels A and C). Burned plants produced nearly four times more flowers than unburned plants (panel B), and had a higher seed set and heavier seeds than unburned plants in both years (panels D and E). For the statistical analyses, see Table 1. Figure 2. Hypothesized effects of post-fire resprouting on plant resistance to drought over time. Following a fire, resprouting plants may initially experience a period of heightened vulnerability to drought and reduced reproduction due to depleted reserves (vulnerability gap). As reserves are replenished and competition levels are not fully restored, resistance to drought temporarily surpasses pre-fire levels (post-fire advantage window); during this period, reproduction is enhanced. Eventually, as community biomass increases, competition recovers and individuals age, the effect of drought returns to pre-disturbance levels. Figure 1 Figure 2. Information & Authors Information Version history V1 Version 1 29 July 2025 Copyright This work is licensed under a Non Exclusive No Reuse License. Collection Oikos Keywords anthyllis cytisoides combined disturbances drought fitness post fire resprouting Authors Affiliations Jaime Saiz Blanco 0009-0002-2947-1634 CIDE-CSIC View all articles by this author Conchita Alonso 0000-0002-7418-3204 Estación Biológica de Doñana, EBD-CSIC View all articles by this author Juli Pausas [email protected] CIDE-CSIC View all articles by this author Metrics & Citations Metrics Article Usage 282 views 198 downloads .FvxKWukQNSOunydq8rnd { width: 100px; } Citations Download citation Jaime Saiz Blanco, Conchita Alonso, Juli Pausas. Resprouting plants are more resilient to drought: increased fitness in Anthyllis cytisoides. Authorea . 29 July 2025. DOI: https://doi.org/10.22541/au.175379050.00325607/v1 If you have the appropriate software installed, you can download article citation data to the citation manager of your choice. Simply select your manager software from the list below and click Download. For more information or tips please see 'Downloading to a citation manager' in the Help menu . Format Please select one from the list RIS (ProCite, Reference Manager) EndNote BibTex Medlars RefWorks Direct import Tips for downloading citations document.getElementById('citMgrHelpLink').addEventListener('click', function() { popupHelp(this.href); return false; }); $(".js__slcInclude").on("change", function(e){ if ($(this).val() == 'refworks') $('#direct').prop("checked", false); $('#direct').prop("disabled", ($(this).val() == 'refworks')); }); Cited by Jaime Saiz‐Blanco, Conchita Alonso, Carlos M. Herrera, Juli G. Pausas, Phenotypic rebuilding after fire: Understanding within‐individual variability in a Mediterranean shrub, Journal of Ecology, 113 , 10, (2921-2934), (2025). https://doi.org/10.1111/1365-2745.70137 Crossref Loading... View Options View options PDF View PDF Figures Tables Media Share Share Share article link Copy Link Copied! Copying failed. 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