{"paper_id":"2fe7f159-da58-4f4a-b209-bf2a47ab3fb5","body_text":"PREPRINT\nAuthor-formatted, not peer-reviewed document posted on 30/08/2024\nDOI: https://doi.org/10.3897/arphapreprints.e135900\nSuppression of an invasive pine by a native shrub\nfollowing a megafire\n Víctor Escobedo, Persy Gomez,  Marco A. Molina-Montenegro, Ian S. Acuña-Rodríguez\n\nSuppression of an invasive pine by a native shrub following a megafire 1 \nVíctor M. Escobedo 1,2, Persy Gómez 1, Marco A. Molina -Montenegro1,3, and Ian S. Acuña -2 \nRodríguez1,2 3 \n1Centro de Ecología Integrativa, Instituto de Ciencias Biológicas, Universidad de Talca, Talca, 4 \nChile 5 \n2Dirección de Investigación, Vicerrectoría Académica, Universidad de Talca, Talca, Chile 6 \n3Centro de Investigación en Estudios Avanzados del Maule (CIEAM), Universidad Católica del 7 \nMaule, Talca, Chile 8 \nFor correspondence: victor.escobedo@utalca.cl, ian.acuna@utalca.cl. 9 \n  10 \nAuthor-formatted, not peer-reviewed document posted on 30/08/2024. DOI:  https://doi.org/10.3897/arphapreprints.e135900\n\nAbstract 11 \nSeedling density of the Chilean wineberry Aristotelia chilensis negatively correlates with the 12 \nseedlings’ abundance of an invasive pine Pinus radiata, particularly in post-fire areas. This pattern 13 \nemerged following a megafire in Chile’s Coastal Maulino Forest, a biodiversity hotspot facing 14 \nincreasing fire threats. This pattern, coupled with a high proportion of plots lacking pine seedlings, 15 \nsuggests that A. chilensis may play a role in limiting P . radiata invasion. The negative relationship 16 \nwas strongest in areas with moderate fire severity, likely reflecting differences in shade tolerance. 17 \nA. chilensis, a light-demanding species with some degree of shade tolerance, can persist in partially 18 \nshaded environments. In contrast, P . radiata, a more strictly light-demanding species, struggles to 19 \nestablish under significant shade. In h igh-severity fires, however , we found no significant 20 \nrelationship between these species , likely due to detrimental effects on both species, including 21 \npotential microbiome dependence for A. chilensis. As A. chilensis shows successful establishment 22 \nat low fire severity, enhancing its post-fire recruitment, particularly in moderately burned areas, 23 \ncould be a valuable strategy for mitigating P . radiata invasion and restoring fire -affected 24 \nMediterranean ecosystems. 25 \nKeywords 26 \nInvasion resistance, Fire severity, Post-fire establishment, Soil microbiome 27 \n  28 \nAuthor-formatted, not peer-reviewed document posted on 30/08/2024. DOI:  https://doi.org/10.3897/arphapreprints.e135900\n\nIntroduction 29 \nWildfires pose a significant threat to biodiversity, disrupting ecosystem functions and threatening 30 \nsensitive habitats  worldwide. Their increased frequency and intensity are attributed to various 31 \nfactors, including climate change and land -use modifications (McLauchlan et al. 2020) . The 32 \nCoastal Maulino Forest, a biodiversity hotspot in central Chile (Myers et al. 2000), is facing more 33 \nfrequent and intense  wildfires in last decades, driven by factors such as rising temperatures , a 34 \nmegadrought and the forestry plantations of non-native species (González et al. 2018, 2023), some 35 \nof which could become invasive after fire disturbances . Primarily, the invasive species is Pinus 36 \nradiata (Pinaceae) which covers approximately 60% of the country’s 2.5 million hectares of forest 37 \nplantations (Bustamante and Simonetti 2005, Gonzá lez et al. 2018) . The devastating 2017 “Las 38 \nMáquinas” megafire burned over 200,000 ha of the Coastal Maulino Forest , a stark reminder of 39 \nthe vulnerability of this ecosystem (Valencia et al. 2018) . Despite ongoing active and passive 40 \nrestoration efforts  in south -central Chile  (Morales et al. 2021, Souza -Alonso et al. 2022) , 41 \nchallenges persist, including the rapid arrival of post-fire pine regeneration that hinders restoration 42 \nsuccess (Gómez et al. 2019, González et al. 2020, 2023). This highlights the need for conservation 43 \nand restoration practices tailored to this unique ecosystem. 44 \nInvasive species often display rapid resource utili sation, potentially outcompeting native 45 \nspecies and promoting more frequent fire events. This can create a positive invasion-fire feedback 46 \nloop (Contreras et al. 2011, Taylor et al. 2017). P . radiata is a light-demanding and shade-intolerant 47 \nspecies known for its aggressive post-fire regeneration through serotinous cones, which release 48 \nlarge amounts of viable wind -dispersed seeds after fire events (Franzese and Raffaele 2017) . 49 \nStudies have shown a higher probability of fire ignition in areas dominated  by P . radiata 50 \nplantations compared to native forests in south -central Chile (Contreras et al. 2011, Gómez -51 \nGonzález et al. 2019). 52 \nPrevious research suggests limited success in controlling P . radiata invasion through 53 \noverall native species diversity (Gómez et al. 2019, González et al. 2020) . However, recent field 54 \nstudies provide evidence that the native wineberry species Aristotelia chilensis (Elaeocarpaceae) 55 \nefficiently recolonises burnt areas even where P . radiata is present (Promis et al. 2019, Becerra et 56 \nal. 2022) . A. chilensis  is a fast -growing, light -demanding, fleshy -fruiting bird -dispersed tree 57 \nspecies with a semi -dioecious leaf habit. Th ese traits allow it to not only colonise clearings but 58 \nAuthor-formatted, not peer-reviewed document posted on 30/08/2024. DOI:  https://doi.org/10.3897/arphapreprints.e135900\n\nalso persist after plantations replace native forest s because it can exhibit some shade tolerance 59 \n(Guerra et al. 2010, Salgado -Luarte and Gianoli 2012). This rapid establishment and fast growth 60 \nof A. chilensis would align with the concept of the “ pre-emptive resource effect” – a mechanism 61 \nwhere early colonising native species can out compete invasive plants by monopoli sing essential 62 \nresources (Byun et al. 2013, Byun and Lee 2017, Delavaux et al. 2023) . Additionally, studies 63 \nsuggest that P . radiata, being a shade-intolerant species, might struggle to establish into a darker 64 \nunderstory dominated by A. chilensis and other na tive species (Gómez et al. 2019, Becerra and  65 \nSimonetti 2020). The efficient colonisation and fast growth of A. chilensis suggest that it has the 66 \npotential to act as a native plant competitor against P . radiata invasion in fire-affected ecosystems. 67 \nBuilding upon competition-based biotic resistance (Elton 1958) and the theory of limiting 68 \nsimilarity, where native species can limit invasive plant establishment due to niche overlap,  we 69 \nhypothesised that P . radiata abundance would negatively correlate with increasing A. chilensis 70 \nabundance. Specifically, we tested the relationship between the abundance of A. chilensis and P . 71 \nradiata in plots affected by varying fire severity levels caused by the Las Máquinas mega -fire in 72 \nthe Maulino Coastal Forest. Additionally, we explored whether fire severity modulates this 73 \nrelationship. Moderate - or low -severity fires  that increase light penetration while retaining 74 \nunderstory vegetation could favour A. chilensis  establishment, potentially strengthening its 75 \ncompetitive effect on P . radiata (i.e., a negative relationship). In contrast, high-severity fires that 76 \ncreate harsher conditions and potential soil disruption (i.e., depleting the soil microbiome) could 77 \nhinder the establishment of both A. chilensis and P . radiata, obscuring any competitive effects. By 78 \nelucidating these dynamics, we aim to provide valuable data to guide  and enhance conservation 79 \nand restoration efforts in fire-affected areas across the central Mediterranean region of Chile. 80 \nMaterials and methods 81 \nStudy Site 82 \nThe study was conducted at El Porvenir (35°42’ S, 72°22’ W), located at the northern edge of the 83 \nCoastal Maulino Forest in central -south Chile (Gómez et al. 2022) . El Porvenir is a fragment of 84 \nnative mesic forest type, surrounded by large stands of planted P . radiata and Eucalyptus globulus 85 \n(Myrtaceae). The dominant tree species include Nothofagus glauca  (Nothofagaceae), N. 86 \nalessandrii, N. obliqua, Cryptocarya alba (Lauraceae), Aextoxicon punctatum (Aextoxicaceae), 87 \nGevuina avellana (Proteaceae), and A. chilensis. Study area ha s a Mediterranean climate with a 88 \nAuthor-formatted, not peer-reviewed document posted on 30/08/2024. DOI:  https://doi.org/10.3897/arphapreprints.e135900\n\nmean annual precipitation of 918 mm and a mean annual temperature of 12.7 °C (Becerra and 89 \nSimonetti 2020). 90 \nIn January 2017, the Las Máquinas megafire affected El Porvenir, which experienced fire 91 \nseverity ranging from low to high (see Valencia et al. 2018, Gómez et al. 2022). Following the fire, 92 \nseveral species exhibited regeneration at different levels, with high seedling recruitment for  the 93 \ninvasive P . radiata and the native A. chilensis (Gómez et al. 2022). 94 \nPlot establishment and seedling survey 95 \nTo assess the potential role of A. chilensis in limiting P . radiata invasion, we established twenty-96 \nthree 625 m 2 plots across El Porvenir. These plots were randomly distributed within areas 97 \nexperiencing low (n = 8), moderate (n = 10), and high (n = 5) fire severity (see Gómez et al. 2022). 98 \nSeedling surveys were conducted at 8 and 24 months following the Las Máquinas mega -fire 99 \n(hereafter 2017 and 2019). To estimate the density of A. chilensis and P . radiata seedlings, three 1 100 \nm2 sub-plots were randomly located within each plot to search for regenerating A.chilensis and P . 101 \nradiata seedlings under 60 cm in height. The average number of seedlings per 3 m2 sampled area 102 \nwas 21.98 ± 20.34 for A. chilensis and 7.46 ± 12.78 for P . radiata. To confirm that the seedlings 103 \noriginated from seeds and not resprouts , we collected at least three random plant samples per 104 \nspecies from each sub-plot for root system examination. 105 \nData analysis 106 \nWe performed a negative binomial Generali sed Linear Mixed-effects Model ( NB GLMM) to 107 \nanalyse the relationship between the abundance of A. chilensis  and P . radiata seedlings. This 108 \nstatistical method is suitable for counting data with overdispersion, a common characteristic of 109 \necological data. Here, we accoun t for the potential influence of sampling time at each plot by 110 \nincluding time since the fire as a random factor nested within fire severity. This nested structure 111 \nconsiders the variation in fire severity across the landscape while acknowledging the potenti al 112 \ninfluence of sampling time within each fire severity category (see above).  Additionally, we 113 \nconducted separate NB GLMM analyses for each fire severity level, including time sampling as a 114 \nrandom factor. 115 \nResults and Discussion 116 \nAuthor-formatted, not peer-reviewed document posted on 30/08/2024. DOI:  https://doi.org/10.3897/arphapreprints.e135900\n\nOur analysis revealed a negative relationship between A. chilensis and P . radiata abundance across 117 \nthe study site (ꭕ2(1,46) = 8.0707, p < 0.01; Fig. 1). Thus, areas with higher numbers of A. chilensis 118 \nseedlings have fewer P . radiata seedlings, potentially indicating a suppressive effect of native 119 \nspecies on invasive tree establishment. 120 \nFurthermore, our results suggest that the strength of this negative relationship varied 121 \ndepending on fire severity. A significantly negative relationship between A. chilensis and P . radiata 122 \nwas found in areas with moderate fire severity (ꭕ2(1,20) = 16.385, p < 0.01; Fig. 2). In contrast, these 123 \ntwo species had no significant relationship  in plots with high or low fire severity (Fig. 2). This 124 \npattern hints that fire severity might play a role in mediating the interaction between A. chilensis 125 \nand P . radiata. Wildfire severity plays a crucial role in shaping post-fire succession and ecosystem 126 \ndynamics. Understanding the severity-specific effects of fires is essential for developing effective 127 \nforest restoration and conservation management strategies. 128 \nThe observed negative correlation between the abundance of A. chilensis and P . radiata 129 \nsuggests that the former’s presence, as a component of the pre-fire native flora, can be considered 130 \na predictor variable influencing P . radiata establishment in post-fire areas. Given that A. chilensis 131 \nwas already present in these ecosystems before the fires, its abundance at the time of the fire event 132 \nlikely influenced the available resources and habitat conditions for P . radiata establishment. 133 \nSeveral mechanisms could explain this phenomenon, including the priority effect by pre-empting 134 \nresources and habitat filtering (Byun et al. 2013, Byun and Lee 2017). First, A. chilensis is a fast-135 \ngrowing, light-demanding species. In areas with a higher abundance of A. chilensis, competition 136 \nfor light, water, or nutrients could be hindering the successful establishment of P . radiata seedlings. 137 \nFuture studies that quantify resource availability and seedling performance concerning A. chilensis 138 \ndensity could provide stronger evidence for this hypothesis.  Second, fire can have profound  and 139 \ndifferent effects on plant community assembly depending on its severity (McLauchlan et al. 2020). 140 \nThe environmental conditions created by moderate fire severity may be  more favourable for the 141 \nestablishment of native compared to invasive species. Specifically, these fires create a more open 142 \ncanopy with increased light availability in the understory, typically forming a patchy mosaic of 143 \nburned and unburned areas rather than eliminating the entire canopy. While A. chilensis is a light-144 \ndemanding species with some degree of shade tolerance  (Guerra et al. 2010, Salgado -Luarte and 145 \nGianoli 2012) , P . radiata is a strictly shade -intolerant species (Gómez et al. 2019) . Thus, this 146 \nAuthor-formatted, not peer-reviewed document posted on 30/08/2024. DOI:  https://doi.org/10.3897/arphapreprints.e135900\n\nvariation in light availability could favour A. chilensis over P . radiata establishment in suitable 147 \nmicrosites within the burned landscape. In low-severity fires with more remaining canopy cover, 148 \nP . radiata pine showed very low establishment (only two plots with 9 and 15 seedlings), likely due 149 \nto limited light availability for germination and seedling growth. In contrast, A. chilensis, which 150 \ncan exhibit some shade tolerance, could persist and thrive, with an average of 16 seedlings per plot 151 \nand up to 67 in one case (Fig. 2). High-severity fires present a vastly different scenario, where 152 \nmost or all vegetation is fire-consumed and heat sterilises the soil, eliminating vital microbes and 153 \ndisrupting biogeochemical processes. These harsh conditions are detrimental to the establishment 154 \nof both species, resulting in the lack of relationship observed in Fig. 2. In this line, A. chilensis’s 155 \nlower establishment suggests a dependence on healthy soil microbes (Escobedo et al. 156 \nunpublished), which are eliminated by high -severity fires. P . radiata, meanwhile,  sometimes 157 \nshowed higher abundance in these areas, potentially benefiting from the open and A. chilensis-free 158 \nconditions since its establishment and survival are less reliant upon microbe communities 159 \n(Escobedo et al. unpublished). This resilience disparity highlights the threat of high-severity fires 160 \nto the Coastal Maulino forest. 161 \nOur findings suggest that native A. chilensis  might play a role in limiting P. radiata 162 \ninvasion, potentially through competition  and/or habitat filtering. However, this beneficial effect 163 \nmight be compromised in high -severity fire areas, where A. chilensis establishment is hampered 164 \ndue to its dependence on a healthy soil microbiome. High-severity fires that disrupt soil microbial 165 \ncommunities pose a significan t threat to native plant communities and their ability to resist 166 \ninvasion. Promoting the establishment of native species like A. chilensis, particularly in areas with 167 \nmoderate fire severity, could be a valuable strategy for mitigating tree invasion and fostering the 168 \nrecovery of fire-affected Mediterranean ecosystems like the Maulino forest in central Chile. 169 \nAcknowledgements 170 \nPG was supported by the Global Botanic Garden Fund number 2022/022 (Botanic Gardens 171 \nConservation International, BGCI). ISAR was supported by ANID-FONDECYT grant 172 \n11240628. 173 \nData Availability 174 \nAuthor-formatted, not peer-reviewed document posted on 30/08/2024. 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Forest Ecology and Management 509: 120083. 286 \nhttps://doi.org/10.1016/j.foreco.2022.120083  287 \nTaylor K, Maxwell B, McWethy D, Pauchard A, Nunez M, Whitlock C (2017) Pinus contorta 288 \ninvasions increase wildfire fuel loads and may create a positive feedback with fire. 289 \nEcology 98: 678–687. https://doi.org/10.1002/ecy.1673  290 \nAuthor-formatted, not peer-reviewed document posted on 30/08/2024. DOI:  https://doi.org/10.3897/arphapreprints.e135900\n\nValencia D, Saavedra J, Brull J, Santelices R (2018) Severidad del daño causado por los 291 \nincendios forestales en los bosques remanentes de Nothofagus alessandrii Espinosa en la 292 \nRegión del Maule de Chile. Gayana. Botánica 75: 531–534. 293 \nhttps://doi.org/10.4067/S0717-66432018000100531 294 \n 295 \nFigures 296 \nFigure 1. Model-predicted relationship between Aristotelia chilensis and Pinus radiata seedlings 297 \nabundance for two sampling times (2017 and 2019). Line indicates a statistically significant 298 \nnegative relationship (p < 0.05) based on a negative binomial GLMM. 299 \n 300 \n  301 \nAuthor-formatted, not peer-reviewed document posted on 30/08/2024. DOI:  https://doi.org/10.3897/arphapreprints.e135900\n\nFigure 2. Model-predicted relationships between Aristotelia chilensis and Pinus radiata seedling 302 \nabundance across fire severity levels (a, low; b, medium; c, high) for two sampling times (2017 303 \nand 2019). The solid line in the middle panel (b, moderate-severity fire area) indicates a 304 \nstatistically significant negative relationship (p < 0.05) based on a negative binomial GLMM. 305 \nRelationships were not statistically significant in high- or low-fired-severity areas. 306 \n 307 \nAuthor-formatted, not peer-reviewed document posted on 30/08/2024. DOI:  https://doi.org/10.3897/arphapreprints.e135900","source_license":"CC-BY-4.0","license_restricted":false}