Escape from the soil surface: how burial by scatter-hoarding rodents may help tree seeds survive forest fire

Escape from the soil surface: how burial by scatter-hoarding rodents may help tree seeds survive forest fire | 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 Search Search ADVANCED SEARCH SCROLL This is a preprint and has not been peer reviewed. Data may be preliminary. 10 December 2025 V1 Latest version Share on Escape from the soil surface: how burial by scatter-hoarding rodents may help tree seeds survive forest fire Authors : Ryunosuke Okawa 0009-0007-1038-3024 [email protected] , Kaoru Kitajima , and Hiroki Sato 0000-0003-2290-2286 Authors Info & Affiliations https://doi.org/10.22541/au.176536813.37007469/v1 296 views 210 downloads Contents Abstract ABSTRACT Information & Authors Metrics & Citations View Options References Figures Tables Media Share Abstract Scatter-hoarding rodents not only contribute to secondary seed dispersal, but also, through burial, to protect seeds from various biotic and abiotic hazards on the soil surface. One such hazard is the heat from wildfires, which are increasing globally in forests that historically did not burn. In this study, we examined whether seed burial by native scatter-hoarding rodent species, Macrotarsomys ingens , would enhance the fire survival of seeds of a non-fire-adapted plant species, Ochna ciliata (seed width ± SD: 5.2 ± 0.3 mm), in a seasonally dry tropical forest of Madagascar acutely threatened by wildfire. We first quantified the removal and subsequent burial of tagged seeds by M. ingens . Using the observed depth (mean depth of 1.5 cm) as a guide, we then designed a mesocosm experiment in which seeds were buried at two soil depths (on the soil surface vs. 1.5 cm below the surface) and exposed to three levels of fire intensity (low, medium, and high) created by fuel load manipulation. The peak temperature was significantly lower at 1.5 cm below than at the soil surface. Buried seeds exhibited significantly higher viability following fire than those left at the soil surface under low and medium fire intensities. Our results suggest that seed burial by scatter-hoarding rodents helps seeds escape from fire, even in non-fire-adapted plant species. Escape from the soil surface: how burial by scatter-hoarding rodents may help tree seeds survive forest fire ABSTRACT Scatter-hoarding rodents not only contribute to secondary seed dispersal, but also, through burial, to protect seeds from various biotic and abiotic hazards on the soil surface. One such hazard is the heat from wildfires, which are increasing globally in forests that historically did not burn. In this study, we examined whether seed burial by native scatter-hoarding rodent species, Macrotarsomys ingens , would enhance the fire survival of seeds of a non-fire-adapted plant species, Ochna ciliata (seed width ± SD: 5.2 ± 0.3 mm), in a seasonally dry tropical forest of Madagascar acutely threatened by wildfire. We first quantified the removal and subsequent burial of tagged seeds by M. ingens. Using the observed depth (mean depth of 1.5 cm) as a guide, we then designed a mesocosm experiment in which seeds were buried at two soil depths (on the soil surface vs. 1.5 cm below the surface) and exposed to three levels of fire intensity (low, medium, and high) created by fuel load manipulation. The peak temperature was significantly lower at 1.5 cm below than at the soil surface. Buried seeds exhibited significantly higher viability following fire than those left at the soil surface under low and medium fire intensities. Our results suggest that seed burial by scatter-hoarding rodents helps seeds escape from fire, even in non-fire-adapted plant species.Keywords: Forest fires, non-fire-adapted plant species, seed dispersal, scatter-hoarding, seed-caching behavior, rodents Introduction Seeds of many plant species are dispersed by scatter-hoarding animals, many of which are rodents that store food for later use in multiple caches, each containing one or a few items (Morris 1962; Gómez et al . 2019). Unlike other dispersal agents (e.g., water, wind, gravity, and frugivorous animals), scatter-hoarders bury seeds below litter and soil surface. The burial depths by scatter-hoarding rodents are shallow at many concealed locations, yet enough to reduce the risk of seed death due to desiccation, freezing, and predation (as reviewed in Lichti et al . 2017). Hence, cached seeds that are not retrieved may have a greater chance of germination (Vander Wall 1990) and subsequent seedling establishment (Giannoni et al. 2001).Scatter-hoarding may also place seeds in safe sites protected from fire, an important ecological process that unpredictably affects plant survival and reproduction over time and space (Keeley et al. 2011; Pausas & Keeley 2014; Lamont et al. 2019). While the temperature at the soil surface rises above 400 °C during a forest fire, it remains below 100 °C just a few centimetres below the soil surface (DeBano 1988; Ando et al. 2014; Daibes et al . 2018). From ecosystems where fires occur frequently, it is reported that shallow belowground caches of scatter-hoarding rodents enhance post-fire seed germination (Rusch et al. 2014; Moore & Vander Wall 2015). Seeds of many plant species in such fire-frequent ecosystems have fire-adaptive traits, such as serotiny, physical dormancy, and smoke-induced germination (Keeley et al. 2011; Enright et al. 2014; Keeley & Pausas 2022; Simpson et al. 2025). In contrast, plants that inhabit environments that do not have periodic fires may not have heat-tolerant seeds, and thus it remains unknown whether shallow burial may enhance the survival of such non-fire-adapted seeds.Globally, incidents of wildfires are increasing in ecosystems where fire was historically rare or absent (Chisholm et al. 2016; Barlow et al . 2020; Hu et al . 2015). Various anthropogenic drivers, including climate change, land-use management, and invasive exotic plants, alter fire regimes (Kelly et al. 2020), causing significant loss of forest cover and biodiversity (Whelan 1995; Bond & Keeley 2005; Bowman et al . 2009). Such is the case in Madagascar, where fires caused by human activities, including burning forests to expand agricultural and pastoral lands, are causing rapid and widespread forest losses (Ralimanana et al . 2022). Despite the urgency of this situation, few studies have examined the potential roles of seed dispersers for post-fire vegetation recovery in these ecosystems. In particular, it remains unclear whether seed burial by scatter-hoarding rodents contributes to the survival of even seeds from non-fire-adapted plant species following fire.We designed the current study in order to evaluate the potential benefit of seed burial by scatter-hoarding rodents through escape of the acute heat at the soil surface during the fire. Using a non-fire adapted tree species in a seasonally dry forest in Madagascar, we quantified: 1) the proportion and burial depth of seeds by scatter-hoarding rodents in a natural forest; 2) the temperature at the surface and 1.5 cm below the soil surface during experimental burning of dry leaf litter with three levels of fuel load; and 3) the proportion of survival and subsequent germination of unburied and buried seeds under three levels of experimental burn intensity. The dense natural forest at our study site lacks historical fire regimes, but fire incidents have increased sharply since 2014, resulting in severe forest losses (Percival et al. 2024, Rasolozaka et al. 2025). At our study site, we reported the first case of a native rodent species engaging in scatter-hoarding in Madagascar, removing seeds of various tree species (Okawa et al . 2025). Thus, it is an ideal site to examine whether seed burial by scatter-hoarding rodents may contribute to fire survival of non-fire-adapted plant species. Materials and Methods Study site & species The study was conducted in Ankarafantsika National Park (136,513 ha) in the Boeny Region of northwestern Madagascar, that protects the largest semi-deciduous dry forest in the country under a strongly seasonal climate of rainy (November–April: with a total precipitation of 1371.9 ± 158.3 mm) and dry (May–October: with a total precipitation of 5.2 ± 4.5 mm) seasons (Sato 2022). In particular, within the primary forest of Jardin Botanique A (JBA) designated for scientific research within the park, many ecological and biodiversity studies have been conducted, including studies on plant species diversity and forest community structure (Fujimoto et al. 2024; Komada et al. 2025) and seed dispersal by animals (primates by Sato 2012, 2022 and Valenta et al. 2013; reptiles by Fukuyama et al. 2025; rodents by Okawa et al. 2025).We examined seed removal and caching by an endemic rodent, Macrotarsomys ingens (Nesomyidae) (mean weight: 56.5 g; Figure 1A) (Razafindratsima et al. 2018). Our earlier study has shown that M. ingens , but not other rodent species, remove seeds of various plant species in this forest (Okawa et al . 2025). For the current experiment, we chose a medium-sized tree species, Ochna ciliata (Ochnaceae), with a maximum diameter at breast height of 15.4 cm. The mean fruit and seed widths ± SD are 6.4 ± 0.2 mm and 5.2 ± 0.3 mm, respectively (Figure. 1B). O. ciliata is not dominant at the study site (Fujimoto et al . 2024), but it bears fruits during the late dry season when most forest fires occur (Percival et al. 2024), and the lead author has observed that M. ingens frequently remove its seeds (Lead author unpublished data). Seed tagging & tracking To investigate the burial depth of O. ciliata seeds cached by M. ingens , we conducted seed tracking experiments during October–November in 2024. We selected four mature trees of Ochna ciliata , which were separated by more than 50 m from each other. Ripe fruits were collected from underneath the four focal trees or non-focal trees in October 2024, and seeds were extracted from fruit pulp. We excluded damaged seeds that floated in water. Experimental seeds were marked individually by attaching a coded plastic tag (3.5 × 2.5 cm) with a 12 cm thin steel thread following the method of Xiao et al. (2006) (Figure S1A). To locate removed seed at night, each plastic tag was painted with non-toxic, UV fluorescent paint (MLC-20, Ohira Seisakusho, Aichi, Japan).We cleared a small area (50 × 35 cm) of any existing fruits and seeds from the ground under each focal tree to create a gridded arena. Within each gridded arena, we placed 30 tagged seeds for a total of 120 tagged seeds across all arenas and identified the animals that removed seeds using infrared cameras with motion-triggered shutters (Strike Force Pro X 1080, Browning). Cameras were mounted 50–70-cm aboveground on focal or adjacent tree trunks and set to record 20-s videos with 1-s intervals between recordings. We searched for removed seeds within a 20 m radius around focal trees using a UV flashlight (UV-SU385-01W, Kontec Inc., Osaka, Japan) at dawn to minimize disturbance (Figure S1B; C). After a thorough search, we categorized seed fates as eaten, intact (on the litter surface), cached, or missing. When we found an intact or cached seed, we recorded the dispersal distance from its initial position (i.e., the arena). For cached seeds, the depth from the top of the seed to soil surface was measured. Litter collection & measurement To measure the abundance of potential fuel, we chose 30 random locations, separated with intervals of more than 20 m, within the study site. To estimate the variation of fuel loads in the study site, we collected litter of various levels of decomposition (e.g., fallen leaves, twigs, bark, cones, and small branches) from 30 cm x 30 cm areas in October 2024. Because they were naturally air dried after ca. 6 months of no rain, we considered their weight as the dry fuel weight. After confirming the normality (Shapiro-Wilk test: W = 0.97, p = 0.68), we determined the minimum, mean, and maximum densities of dry fuel load per unit area of ground as 371.1, 877.2, and 1389.4 g/m 2 , respectively (Figure S2). Mesocosm litter burning experiment We constructed an open-top brick enclosure (45 cm tall × 120 cm wide × 70 cm long) divided into four compartments (Figure 1C) to conduct an experimental burning with three levels of fire intensity (low, medium, and high). We filled the bottom with sandy soil containing roots collected from JBA. The study had two levels of seed placement (at the surface and 1.5 cm below the soil surface) and three levels of fuel load (low, medium, and high, corresponding to 400, 900, 1400 g/m 2 of litter). We collected fruits of O. ciliata and natural litter as fuel from JBA. Litter was re-dried to increase flammability in a metal box heated by a gas burner for one hour prior to pacing in each experimental burn compartment (Figure 1C). Seeds were extracted from fruits and verified that they don’t float in water. Within each compartment, we placed 25 seeds at the soil surface in one half of the area, and 25 seeds buried at 1.5 cm below the surface in the other half. We chose this burial depth to simulate the mean burial depth of scatter-hoarded seeds by M. ingens found in the experiment described in Section 2.2. We then spread the re-dried litter over of the set quantity in each burn compartment. In total, 300 seeds were used at each soil depth (surface and belowground) across the three fuel loads (low: 100, medium: 100, high: 100).We ignited litter of one compartment at a time (Figure 1C). We used two stick thermometers (KT-01U, Custom Corp., Tokyo, Japan; TP510, ThermoPro), the sensor bar of one placed at the soil surface and that of the other placed at 1.5 cm below the soil surface. After ignition of the fuel, temperature was recorded every 15 seconds during the first five minutes, and then every one minute until 30 minutes had elapsed. The fire intensity was quantified as the area under the curve of fire temperature over the 30-min experimental period, using the initial temperature (0-min) of each experimental burn as the baseline. After the experimental burns, seeds were collected from the enclosures, cleaned from soil particles, and then subjected to germination experiments. Germination experiment To examine the viability and germination capacity of seeds after experimental burns, we placed seeds in round plastic tubs (40 cm diameter × 15 cm height) filled with soil (Figure 1D). We prepared 100 intact seeds of O. ciliata not used in the burn experiment to determine the reference level of survival and germination. Each tub was divided to four compartments to receive 25 seeds each from four randomly selected treatment combinations (two seed placement depth × three fuel load levels, plus reference). We planted seeds approximately 5 cm apart, 1.5 cm below the soil surface, and the tubs were placed in a seedling nursery covered with 30% grade shade cloth to reduce stress from direct sunlight and heavy rainfall. Tubs received water daily to keep the soil moist, and seeds were retrieved 12 weeks after planting, approximately 2 weeks after the last seed had germinated. Each seed was classified into three categories: (1) germinated (Figure 1D), (2) hard (ungerminated but firm to pinching with fingers), and (3) dead (ungerminated and soft or charred) following Soares et al. (2021). A small number of missing seeds were excluded from the further analysis. We considered germinated and hard seeds as viable seeds. Statistical analysis We used a generalized linear mixed model (GLMM) with a binomial distribution, in which the dependent variable was individual seed viability (viable or dead). Seed placement (soil surface vs. belowground), fuel load level (low, medium, and high), and their interactions were included as fixed effects, and burn experimental date, enclosure compartment ID, and planting tub ID were included as random effects. When the GLMM result was significant ( p < 0.05), we used Tukey’s multiple comparison test. Viability of the reference seeds was analysed separately since they were not part of the experimental design. We used one-way ANOVA and post hoc pairwise comparisons to examine differences between reference seeds and those exposed to six treatment combinations in the burn experiment. The Statistical Software R 4.4.0 (R Development Core Team 2024) was used for all analysis. Results From camera trapping, we found that only M. ingens removed seeds of O. ciliate dropped or placed under fruiting trees. In total, 70 (58.3%) of 120 tagged seeds were removed from their initial positions, and the rest were eaten in situ. Of the removed seeds, 17 (14.2%) were eaten, 37 (30.8%) were intact, 11 (9.2%) were cached, and 5 (4.2%) were not found. Intact and cached seeds were found at a mean distance of 2.4 m (range: 0.3–9.3 m) from their initial positions. The burial depths were 1.5 cm on average, ranging from 5.0 to 30.0 mm (Table S1). Of the 11 cached seeds, five (4.2% of 120 seeds, or 36% of the 11 cached seeds) were buried at or deeper than 1.5 cm below the soil surface.Fire intensity was significantly higher at the soil surface than at 1.5 cm belowground (Wilcoxon test: p < 0.001; Figure 2). Fire intensity at the soil surface varied among the three fuel load levels (Kruskal-Wallis test: p = 0.02; Figure 2A), with mean maximum temperature (± SD) of 163.7 ± 56.6 °C, 219.5 ± 44.7 °C, and 258.4 ± 69.2 °C for low, medium, and high fuel loads, respectively (N = 4 for each load condition). In contrast, fire intensity belowground did not differ among fuel load levels ( p = 0.06; Figure 2B), with mean maximum temperature (± SD) of 60.9 ± 6.1 °C, 69.8 ± 11.7 °C, and 99.6 ± 22.9 °C under low, medium, and high fuel loads, respectively (N = 4 for each condition).Results of germination experiments showed that approximately half of the seeds died regardless of seed placement and fuel load (Figure 3), while the proportion of viable seeds was affected by seed placement depths and fuel load (Figure 3; Table 1). The mean viability of seeds placed at the soil surface was approximately half of the viability of buried seeds (mean ± SD: 19.7 ± 9.6 % for soil surface; 39.0 ± 16.4 % for belowground). After experiencing heat from burning low and medium fuel loads, the viability of buried seeds was significantly higher compared to those placed at the soil surface, whereas no significant difference was found at high fuel loads (Figure 3; Table S2). Overall seed viability of buried seeds was significantly higher at low and medium fuel loads than at high fuel load, but no significant differences were observed across fuel load levels at the soil surface (Figure 3; Table S2). Seeds in the reference treatment showed significantly higher viability than seeds at the soil surface under all fuel load levels (Figure 3; Table S3). In contrast, only the high fuel load significantly reduced the viability of seeds belowground compared to the reference treatment, but under low and medium fuel load conditions, seeds belowground showed similar viability to the reference. The proportion of germinated seeds was also influenced by both seed placement and fuel load levels (Table S4). At least under low fuel load treatment, germination rates were significantly higher belowground than at the soil surface, and no seeds at the soil surface germinated under high fuel load (Figure 3; Tables S5). Discussion Although numerous studies have demonstrated the contributions of scatter-hoarding rodents to seed dispersal across various forest ecosystems (Gómez et al . 2019), few have examined how seed burial enhances seed survival by protecting seeds from physical hazards, including lethal heat from fire. Our study indicated that seeds buried just 1.5 cm below the soil surface could survive the heat generated by burning low to intermediate amounts of dry litter. As seeds surviving fire can be a significant contributor to vegetation recovery, our results suggest that scatter-hoarding rodents can potentially contribute not only to seed dispersal but also to post-fire recovery of non-fire-adapted plant species when their adults are killed by fire.We found that M. ingens cached 11 of 120 seeds and buried four of them at depths of 1.5 cm or greater, a depth at which seeds could survive the heat generated by medium fire intensity. In general, scatter-hoarding rodents cache seeds shallowly in the soil (less than 5 cm deep), where temperature and humidity are suitable for seedling establishment (Hollander & Vander Wall 2004). A previous study reported that even seeds smaller than those of O. ciliata can establish seedlings from a depth of 5 cm in the soil (Li et al. 2006). Thus, a depth of 3.0 cm, the maximum depth at which seeds of O. ciliata were buried by M. ingens in the current study, would not be too deep for seedling emergence. The observed probability of seed burial by M. ingens was low. Yet, given the large number of seeds produced at the population level, seed burial by scatter-hoarding rodents can make a critical contribution to the species’ persistence after wildfire.After experimental burns, buried seeds of O. ciliata showed higher viability than seeds at the soil surface. Fire-adapted plant species often have seeds that can be released or break dormancy in response to heat shocks and smoke to germinate in post-fire environments (Keeley & Fotheringham 2000). Our study species, O. ciliata , does not have dormant seeds that germinate in response to fire-related cues, as the germination level was similar between reference and those buried and experienced burn-generated heat. Hence, we can conclude that the soil’s thermal-insulating effect explains why burial would improve seed survival of this non-fire-adapted plant species. Furthermore, forest fires in the study region typically occur during the late dry season (Percival et al. 2024), and germination of O. ciliata required at least one month in our experiment (Figure S3). Thus, non-fire-adapted plant species dispersing seeds in late dry season, such as O. ciliata , would fail to regenerate after fire unless their seeds become buried by scatter-hoarding rodents.In natural forests, seed survival would be patchy because fire intensity and lethal temperatures vary spatially due to small-scale heterogeneity in fuel loads (Thaxton & Platt 2006; Brando et al. 2016). While the fire intensity belowground did not differ significantly among fuel load levels in our experiment, seed survival was higher for buried seeds experiencing heat generated from burning low to intermediate amounts of litter. This was probably because seed viability is influenced not only by the total amount of heat reaching the seeds, but also by the duration of temperature above the critical threshold (Daniell et al. 1969; Dayamba et al. 2010). The lethal temperatures for seeds are 70°C and 90°C in wet and dry soil, respectively (Martin et al. 1975). In our experiment, the temperature at 1.5 cm below the soil surface exceeded 90°C only under the high fuel load (Figure S4), which could explain the lower seed viability in this treatment. As field burning experiments conducted in fire-prone vegetation worldwide (Mucunguzi & Oryem-Origa 1995; Greenberg et al. 2012), direct contact with the flames was lethal for seeds regardless of fire intensity. In addition, the patchiness of dead and live wood, which generates longer-lasting high heat, would add to the spatial and temporal heterogeneity of lethal temperatures during a forest fire. Indeed, the maximum fire temperature at the soil surface near burning dead trees reached 235°C, even after the litter had burned out during an actual forest fire near our study site (N = 1; Lead author, personal observation).Rodents and other small animals tolerant of hunting and habitat fragmentation can play a particularly important role in post-fire vegetation recovery. Large frugivorous vertebrates that act as primary seed dispersal agents are disappearing due to anthropogenic disturbances (Barnosky et al. 2011; Dirzo et al. 2014; Harrison et al. 2013). In contrast, rodents are more tolerant of disturbances and less prone to extinction than large vertebrates (Cardillo et al. 2005; Ripple et al. 2015). Some dispersal agents other than scatter-hoarders also contribute to the burial of seeds (Vander Wall & Longland 2004). For example, dung beetles ubiquitous in tropical regions play an important role in burying animal dung containing seeds at depths favorable for seedling establishment (Shepherd & Chapman 1998; Andresen 2001; 2002). However, these secondary seed dispersal mechanisms depend on primary dispersal, such as seeds being defecated by animals. Thus, even as primary seed dispersers decline under increasing fragmentation, degradation, and human-driven wildfire, secondary dispersers that function independently of them—such as scatter-hoarding rodents—remain crucial for mitigating the decline in the resilience of tropical forest seed-dispersal systems. Table 1. Results of a generalized linear mixed model ANOVA testing the proportion of viable seeds of Ochna ciliate at the soil surface and at 1.5 cm below the soil surface in response to simulated fire under three levels of fire intensity. Seed placement (soil surface vs. belowground) 1 23.57 < 0.001 Fuel load level (low, medium, and high) 2 22.92 < 0.001 Interaction (Seed placement × Fuel load level) 2 5.67 0.06 df, degree of freedom Figure 1. Photographs of study objects: (A) Macrotatsomys ingens , a native scatter-hoarding rodent, (B) fruits (left and upper) and seeds (bottom) of Ochna ciliata , (C) brick enclosure with compartment containing natural forest soil (upper left), leaf and fine branch litter over seeds of O. ciliata placed on the soil surface and belowground (lower left), 30 minutes after fuel ignition (lower right), burned seeds collected after the burn experiment before germination trials (upper right), (D) plastic tub containing 25 seeds from four different treatment combinations (upper), and observation of their fates (germinated, hard, or dead) (lower). Figure 2. Fire temperature during experimental burns (A) at the soil surface and (B) at 1.5 cm below the soil surface with low (blue), medium (orange), high (red) fuel loads. Each fitted line with shaded areas indicating 95% CIs were generated from four replications. Figure 3. Proportion of germinated (black), hard (grey), and dead seeds (white) of Ochna ciliata at the soil surface and 1.5 cm below the soil surface under the reference treatment, and under low (blue), medium (orange), and high (red) fuel loads. For statistical parameters, see Table S2 and S3. References 1. Ando, K., Shinjo, H., Noro, Y., Takenaka, S., Miura, R., Sokotela, S. B., & Funakawa, S. (2014). Short-term effects of fire intensity on soil organic matter and nutrient release after slash-and-burn in Eastern Province, Zambia. Soil Science and Plant Nutrition , 60 (2), 173-182. 2. Andresen, E. (2001). 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Keywords forest fires non-fire-adapted plants rodents scatter-hoarding seed dispersal seed-caching behavior Authors Affiliations Ryunosuke Okawa 0009-0007-1038-3024 [email protected] Kyoto University View all articles by this author Kaoru Kitajima Kyoto University View all articles by this author Hiroki Sato 0000-0003-2290-2286 Kyoto University View all articles by this author Metrics & Citations Metrics Article Usage 296 views 210 downloads .FvxKWukQNSOunydq8rnd { width: 100px; } Citations Download citation Ryunosuke Okawa, Kaoru Kitajima, Hiroki Sato. Escape from the soil surface: how burial by scatter-hoarding rodents may help tree seeds survive forest fire. Authorea . 10 December 2025. DOI: https://doi.org/10.22541/au.176536813.37007469/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. 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