Preferences of rodent scatter-hoard and predators determine dissemination oak species.

Preferences of rodent scatter-hoard and predators determine dissemination oak species. | Research Square window.SnipcartSettings = { analytics: { enabled: false } }; (function() { var accessVector = localStorage.getItem('access_vector') || ''; window.dataLayer = window.dataLayer || []; if (accessVector) { window.dataLayer.push({ user: { profile: { profileInfo: { snid: accessVector } } } }); } })(); (function(w,d,s,l,i){w[l]=w[l]||[];w[l].push({'gtm.start':new Date().getTime(),event:'gtm.js'});var f=d.getElementsByTagName(s)[0],j=d.createElement(s),dl=l!='dataLayer'?'&l='+l:'';j.async=true;j.src='https://www.googletagmanager.com/gtm.js?id='+i+dl;f.parentNode.insertBefore(j,f);})(window,document,'script','dataLayer','GTM-K279D39R'); Browse Preprints In Review Journals COVID-19 Preprints AJE Video Bytes Research Tools Research Promotion AJE Professional Editing AJE Rubriq About Preprint Platform In Review Editorial Policies Our Team Advisory Board Help Center Sign In Submit a Preprint Cite Share Download PDF Research Article Preferences of rodent scatter-hoard and predators determine dissemination oak species. Sergio Del Arco, Jose María Del Arco This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-3410260/v1 This work is licensed under a CC BY 4.0 License Status: Under Review Version 1 posted 4 You are reading this latest preprint version Abstract The relationship between scatter-hoarding rodents and oak species has been considered on a scale from antagonism to mutualism. Depending on the costs and benefits, the outcome of the relationship can be found at one extreme or the other. Costs have included destruction of acorns that occurs during predation, but not all acorns attacked lose their embryos. As representatives of the mutualistic end, we present two species that preserve embryo ( Mus spretus and Apodemus sylvaticus ). Representing the antagonistic extreme, we present a predatory species that destroys the embryo ( Microtus arvalis ). The objective of this study is to test the preferences of both rodent groups for acorns. The results showed that there is one acorn species ( Quercus ilex ) that is preferred by the three rodent species. This acorn species has high concentrations of nutrients, low concentrations of tannins and thin shell. These characteristics attract the attention of rodents that could carry these acorns. There are two other acorn species ( Quercus suber and Quercus rubra ) that are consumed with little intensity for having low concentrations of nutrients, high concentration of tannins and thick shell. These characteristics escaping predators although transportation is not guaranteed. These acorns would germinate and emerge under the trees that have produced them and increase intraspecific competition. Coincidence preferences shown by the three rodent species poses a risk for the oak species, since the recent arrival of the predator (antagonist) species in the study area could paralyze the dispersal process carried out by the other two mutualistic species through predation. Acorn preferences Rodents Preserved embryo Mutualistic Antagonism acorn composition Figures Figure 1 Figure 2 Figure 3 Significance Statement The relationship between rodents and oak species has been termed antagonistic or mutualistic, depending on the outcome. In the antagonistic relationship, one of the species is harmed because it loses its seeds. In the mutualistic relationship, both species obtain benefits. We used rodent species representative of both relationships. We have provided these rodents with several different acorn species to observe if their different behavior during acorn handling causes differences in the choice of these acorn species. The selected acorn species exhibit different characteristics. Some seem intended to avoid predation, but others seem intended to attract the attention of rodents. We have observed that these differences lead to different behaviors in the choice of acorns. Introduction The relationship between scatter-hoarding rodents and oak species varies in their outcomes (Muñoz and Bonal 2011 ; Yi and Wang 2015 ; Gu et al. 2017 ; Lichti et al. 2017 ; Bogdziewicz et al. 2020 ; Mittelman et al. 2021 ). At one end of the ranging is antagonism, a relationship called predation which the species rodent takes resources from species plant that they suffer costs (Galetti et al. 2015 ). In this predation relationship, rodents obtain resources as benefits and plants pay costs. These costs involve the loss of seeds through their destruction by rodent ingestion. As a representative of this group, we present Microtus arvalis . This rodent species devours acorns by opening them at the apical end where the embryo is inserted. This organ is the first element of the acorn to be destroyed. At the other end of the ranging is mutualism. In this relationship, the two participating species obtain benefits, but both must contribute resources for the relationship to remain in equilibrium (Zhang et al. 2015 ; Zhang et al. 2016a ; Moore and Dittel 2020 ). In mutualism, rodents obtain resources by ingesting whole acorns or part of the cotyledons of the acorns they attack (Cao et al. 2018a ; Wróbel and Zwolak 2017 ; Moore and Dittel 2020 ). The costs they bear is the energy they must expend to transport the acorns to places safe for themselves. Normally, these rodents do not consume acorns in situ because their predators are waiting for them at the acorn concentration sites under the acorn-producing trees. They transport the acorns to protected places where they make numerous caches (Lichti et al. 2015 ; Wang et al. 2018 ; Mittelman et al. 2021 ). Some of these caches are forgotten (Lichti et al. 2020 ; Martínez-Baroja et al. 2021 ). Another cost is that they must expend energy to open acorns and remove their shells. Plants in this mutualistic relationship have the cost of losing acorns to rodent ingestion, to make acorns from high nutrient concentrations, synthesis of costly substances such as tannins and thick coatings. However, the benefits they obtain are that part of their acorns are transported by rodents to protected places under bushes where they are buried. There they gain humidity which favors their germination ((Perea et al. 2011 ; Yang and Yi 2012 ). Both species obtain benefits but pay costs. As representatives of mutualism, we present two species: Mus spretus and Apodemus sylvaticus . These two rodent species present a peculiar behavior during the acorn consumption process. This behavior was observed independently by two research groups (Perea et al. 2011 and Del Arco et al. 2018 ). These two species start most of the acorns they process at the basal end away from the embryo and although they consume a variable part of the cotyledons (Yi and Wang 2016 ) they leave for last the apical end with the embryo intact. This behavior allows some acorns (or at least embryos) to escape predation. Perea et al. 2011 and Yang and Yi 2012 , found that acorns with ¾ of the cotyledons consumed if they retain the embryo germinate. For this reason, these authors consider that these two species show a behavior that tends to conserve the embryo. This behavior is innate and not learned (Del Arco et al. 2018 ; Del Arco and Del Arco 2022 ). It is also an active behavior (Muñoz and Bonal 2008a ) as these mutualistic species use the basal end to open the acorns, consuming more energy even though it is easier to open them from the narrower distal end. The origin of the partial consumption of acorns may be in the size of the acorns in relation to the body size of the rodent (Muñoz and Bonal 2008b ). The origin of the conservative behavior of the embryo starting the acorns at the basal end may be that the plant species place high concentrations of tannins at the distal end where the embryo is to deter rodents from ingesting them (Steele et al. 1993 ; Hou et al. 2010 ; Wang et al. 2020 ). Zwolak and Crone ( 2012 ), Bogdziewicz et al. ( 2020 ) and Zwolak et al. ( 2020 ) proposed a mathematical model to assess whether the outcome of scatter-hoarding relationship is antagonism or mutualism (Chang and Zhang 2014 ; Gleditsch et al. 2017 ). When the benefits of transportation and burial are greater than the costs caused by predation during this process of scatter-hoarding, we can call the relationship mutualistic. To evaluate the cost of dispersal, acorn loss due to predation is typically included, however, not every acorn that is attacked loses its ability to germinate. The usual calculations carried out do not include the beneficial effect of partial acorn consumption, which significantly reduces these costs (Del Arco et al. 2018 ; Del Arco and Del Arco 2022 ). We have chosen six species of acorns to feed to the three rodent species. There are species of acorns that possess structures and characteristics associated with defense against predation, such as high concentrations of tannins and thick coatings that lengthen handling times. But there are other characteristics that are more associated with attracting the attention of rodents (Sivy et al. 2011 ; Zhang et al. 2016b ; Xiao et al. 2021 ), such as high nutrient concentrations or thin coatings. The main goal of this study is to test the food preferences of the two groups of rodents that show different behavior during the handling of acorns (preservatives ( A. sylvaticus and M. spretus ) and embryo destroyers ( M. arvalis )) (Pons and Pausas 2007 ; Rosalino et al. 2013 ). We want to know if there are differences between the two groups in the choice of acorn species for their diet (Gómez et al. 2019 ). The different physical and chemical characteristics exhibited by acorns may condition choice in each group (Gong et al. 2015 ; Perea et al. 2016 ). We want to check if these differences are due to the different characteristics of the acorns. Methods Study system For this study, we captured rodents of three different rodent species, Apodemus sylvaticus (Linnaeus, 1758) (wood mouse), Mus spretus (Lataste, 1883) (Algerian mouse), and Microtus arvalis (Pallas, 1778) (common vole). The captured specimens of the three rodent species were placed in terrariums for reproduction. Nine descendants of each rodent species were selected for the experiment to eliminate any previous acorn consumption experience. They were placed in a fenced enclosure of 100 m 2 . Inside we built 9 plots of 9 m 2 (3*3 m side). Each of them was isolated by placing on each side of the square a 3 m wide sheet of metal buried in the ground at a depth of 50 cm and a height from the ground of 1 m to prevent rodents from escaping by jumping. Under these conditions, the mice behaved as if they were in semi-wild conditions: building burrows, creating shallow storages, galleries, pantries, and caches to hide the acorns, which they moved around the plot. The 9 specimens of each species were placed in these plots successively, starting with M. spretus and ending with M. arvalis . The wood mouse has inhabited the Iberian Peninsula since ancient times (Michaux et al. 2003). Acorns of various Quercus species are among its food sources and are abundant in the area it inhabits (Pulido and Diaz 2005; Muñoz et al . 2009). The Algerian mouse is of North African origin but has been present in parts of the Iberian Peninsula for a long time (Palomo et al . 2008). It lives practically in the same habitat as the wood mouse except in the northern mountains. It also consumes acorns from the Quercus species (Muñoz and Bonal 2007). These two species partially consume acorns, opening them at the basal end and preserving the embryo. For this reason, we include them in the mutualism group. A third rodent species, Microtus arvalis , has recently arrived in the region (Luque-Larena et al . 2013; Paz et al . 2012) and does not use acorns for its diet, but consumes them (Del Arco et al . 2018). Common vole is abundant in central Europe. Until recently, its distribution area in the Iberian Peninsula has been confined to the northern mountains, where it can access fresh food such as soft green herbaceous plants. Due to the increase in the distribution area of irrigated crops, the common vole has expanded to the center of the Iberian Peninsula, where it experiences phases of overpopulation, and it is considered a recurrent crop pest (Luque-Larena et al . 2013; Paz et al . 2012). There are no references to this species feeding on acorns despite that the mountain pastures where it originally lives are surrounded by Q. pyrenaica and Q. petraea forest formations, but the humidity of the area means that fresh vegetables are continuously available as food. This species is a risk for the oaks in the center of the Iberian Peninsula because due to phases of overpopulation or by the increasingly pronounced drought can exhaust their traditional food and seek new sources of resources in the acorns and this species is clearly a predator of these seeds (Del Arco et al . 2018). To feed the rodents and check their food preferences, we have selected six species of acorns from the environment where the rodents live because they may constitute their source of resources and maintain some kind of relationship with them. Acorns from the following Quercus species were collected: Quercus ilex subsp. ballota (Desf.) Samp. (holm oak), Q. faginea Lam. (Portuguese oak) at the site where the rodents were caught, Q. pyrenaica Willd. (Pyreneen oak), Q. petraea (Matt.) Liebl. (sessile oak), in the northern mountains of the Iberian Peninsula where the common vole originally lived, Q. suber L. (cork oak) in Salamanca (Spain), and Q. rubra L (American oak) (Bieberich et al . 2016; Merceron et al . 2017) in experimental reforestations in Galicia (Spain). The only non-native species is Quercus. Rubra . We have chosen this species to test its integration in this system of relationships between rodents and oaks if it can adapt to the existing dispersal mechanisms in the area (Bieberich et al . 2016; Merceron et al . 2017). Experimental Procedures and Design Nine specimens of the descendants of each rodent species were placed in isolated plots. Every day, for five days each specimen had six acorns available to them from each of the Quercus species used. During the experiment, each specimen was offered 180 acorns, 30 from each of the six Quercus species. Before being given to the rodents, each acorn was weighed and labeled to estimate the individual mass consumed of each acorn. The remains of the acorns from the previous day were removed and weighed to estimate their mass after consumption. To estimate the daily mass consumed of each acorn species, we calculated the sum of the quantities consumed of each acorn by each rodent species specimen for one day. We also estimated the mean mass consumed per acorn of each plant species by each rodent species. Some acorns from each species were analyzed for their nutrient content (Table 1). We analyzed the chemical composition of the acorns to see whether it influenced selection during consumption (Table 1). A high lignin content is related to a higher tannin content (De Blas et al . 2003), which, in turn, could have deterred rodents from eating acorns (Zhang et al . 2013). On the contrary, the higher fat, protein, and sugar contents are usually the characteristics that most attract rodents to ingest acorns (Table 1). To study the similarity in acorn composition, we grouped the species with the most similar composition together (Figure 1) according to the Czekanowski C = (2W/A+B)*100 index. Where W = the sum of the minimum values of each compound in the side-by-side comparison of the two species A = the sum of the values of each component in the first species in the side-by-side comparison B = the sum of the values of each component in the second species in the side-by-side comparison. Another group of acorns was set aside to study their shell thickness. A Mitutoyo caliper was used to measure thickness at three different points: the circular scar where the acorn is attached to the cupule, the periphery of this circle located at the base of the acorn, and the apical end of the acorn, where the shell covers the embryo. Data Analyses The possible effects of rodent species (wood mouse, Algerian mouse, and common vole), day (5 levels), Quercus species ( Q. petraea, Q. ilex, Q. pyrenaica, Q. faginea, Q. suber, Q. rubra ), and their interactions on the number and mass of acorns eaten per specimen, were analyzed using Linear Mixed Models (LMM) with the Restricted Maximum Likelihood method (REML). The specimens were treated as a random factor and the day as a repeated factor. Finally, working on the model matrix, contrasts were carried out to test differences between fixed factor levels (Pinheiro and Bates 2000). Consequently, the Bonferroni correction was used to adjust for the significance level for each t-test (Sokal and Rohlf 1995). The statistical calculations were implemented in the R software environment (version 2.15.3; Core Team R 2013), using the nlme package for LMM (Pinheiro et al . 2013). Results The LMM analysis showed a highly significant interaction between ‘species of acorns’, ‘species of rodent’ and ‘number of acorns’, ‘daily acorn mass’, and ‘mean mass per acorn’ consumed (Table 2), suggesting that rodent species show a preference for some species of acorns. The number of acorns attacked by each rodent species, the daily acorn mass, and the mean mass per acorn did not vary significantly in time. There were no changes in rodent feeding behavior during acorn consumption. Preference for acorn species can be seen in Figure 2. We show the mean number of acorns attacked daily by each mouse of each species (Fig 2a). The two species with the highest number of acorns consumed per day are Q. ilex and Q. pyrenaica . In figure 2b, we show the daily mass consumed by mice of each rodent species. The highest mass of acorns consumed daily in the three rodent species corresponds to Q. ilex acorns. In Figure 2c we present the mean mass per acorn consumed daily by each mouse of the three rodent species. Also in this variable, the highest mass consumed per acorn in the three rodent species belongs to Q. ilex . The species with the lowest number of acorns consumed daily are Q. suber and Q. rubra in the rodent species common vole and Algerian mouse. Wood mouse attacks a daily mean acorns of these species like rest of the acorn species. However, the daily mass consumed of Q. suber and Q. rubra acorns is very low in all three rodent species. The mean daily mass consumed per acorn is also very low in all three rodent species for Q. rubra. Wood mouse shows more preference for Q. suber than Q. rubra as shown by the mass consumed in both species. We analyzed the chemical composition of the acorns to see whether it influenced selection during consumption. There were significant differences in acorn composition (Table 1). Figure 1, we show a cluster analysis for the six acorn species based on seed composition. Q. ilex is the species that differs most in terms of composition from other species. We also verified whether shell thickness influenced selection. Figure 3 shows the mean shell thickness in the six plant species. The two most rejected species ( Q. suber and Q. rubra ) have thicker shells. Discussion Looking at the results obtained, we have verified that the two groups of rodents' species of this study (preservers embryo (wood mouse and Algerian mouse) and embryo destructors Common vole) have the same preferences for acorns (Pons and Pausas 2007 ; Rosalino et al. 2013 ; Gómez et al. 2019 ). Both groups of rodents showed a preference for the Q. ilex and Q. pyrenaica acorns and consumed Q. suber and Q. rubra with less intensity (Fig. 2 ). These two groups of acorn species differ in their physical and chemical characteristics (Gong et al. 2015 ; Perea et al. 2016 ). (Fig. 1 ). The most appreciated ones have a higher concentration of nutrients, lower concentration of tannins (Table 1 ) (Perea et al. 2012 ) and thin shells (Huang et al. 2011 ) (Fig. 3 ). The least appreciated, on the contrary, high tannin concentrations, low in nutrients (Table 1 ) and thick shells (Fig. 3 ). Q. ilex , which is the most appreciated, differs significantly from the rest according to the cluster analysis (Fig. 1 ). Investing high proportions of protein, sugars and fats gives the acorns of this specie characteristics associated with intense consumption. It is a good candidate for obtaining resources during adverse periods (Lichti et al. 2017 ). Why invest high concentration of resources and what benefits do they get in return? The answer to this question is probably to be found in the mutualistic relationship that this plant species has long maintained with Wood mouse and Algerian mouse. These two rodent species practices partial consumption of acorns, preserving the embryo, and transport their acorns to caches because they do not consume them in situ. It is possible that the relationship was initiated by the attraction of rodents to the appetizing characteristics of the acorns of this species (Xiao and Krebs 2015 ; Bogdziewicz et al. 2020 ; Moore and Dittel 2020 ; Zwolak et al. 2020 ). For this reason, this species of acorns is the most transported to caches (Lu and Zhang 2004 ; Lichti et al. 2020 ). This means that it is the most favored in the dispersal process (Correa and Uriarte 2013 ; Wang et al. 2014b ; Kang et al. 2020 ). But this acorn specie has a problem, which also attracts the attention of the predatory specie in this study (Common vole) more intensely than other acorn species with lower nutrient concentrations (Lichti et al. 2020 ) (Fig. 2 ). Therefore, most of their acorns are destroyed more intensely than in other less preferred species by this rodent specie (Common vole). The risk for this acorn specie is that the natural food of Common vole disappears due to continuous drought in the study area or recurrent high densities of these rodent, that they look for new alternative sources of resources and end up incorporating acorns to their usual diet (Luque-Larena et al. 2013 ; Paz et al. 2012 ). We have already seen that the acorns it destroys are a very appetizing food for this rodent. The dispersal process of these Q. ilex acorns could be paralyzed by the presence of this rodent specie (Pesendorfer et al. 2016 ; Del Arco et al. 2018 ; Del Arco and Del Arco 2022 ). There are other acorns species ( Q. suber and Q. rubra) that are not appreciated for consumption by the two groups of rodents (mutualists and antagonists) (Fig. 2 ). These species present defensive characteristics to avoid consumption. High concentrations of tannins (Table 1 ) and thick coatings to prevent access (Fig. 3 ) (Zhang et al. 2013 ; Lai et al. 2014 ; Wang et al. 2014a b ; Lichti et al. 2017 ; Zhang et al. 2018 ). If they escape predation they can germinate and emerge, but they have the problem that there is no transport for these acorns. If they emerge, it is under the adult tree that has originated them, and this obviously increases intraspecific competition (Moore and Dittel 2020 ). The process of colonization of new sites will be paralyzed. They are not buried because this mission is carried out by dispersing rodents (wood mouse and Algerian mouse) that do not select them as a priority for food. These two less preferred species ( Q. suber and Q. rubra) have high concentrations of tannins. This may corroborate the statements of Steele et al. ( 1993 ); the partial consumption of acorns and the preservation of the embryo is a behavior that may have been generated by the inclusion of high concentrations of tannins around the embryo during made of acorns. These bad-tasting tannins discourage rodents from ingesting the embryo end by forcing them to open the acorns at the basal end. If acorns with higher concentrations of tannins are rejected, so will be the end with high amounts of these substances. However, since three rodent species reject the same species of acorns possibly because of the presence of tannins, they would also avoid opening the acorns at the apical end where these substances are most concentrated, however, the predatory specie (Common vole) initiates the acorns at the apical end destroying the embryo, not preserving it (Del Arco et al. 2018 ; Del Arco and Del Arco 2022 ). Why do they show different behavior in this open acorn process? The answer may lie in Del Arco et al. ( 2018 ) and Del Arco and Del Arco ( 2022 ). By these, the embryonic conservative behavior is embedded in the genetic code of the rodent species that practice it ( Apodemus sylvaticus and Mus spretus ). Captive-bred specimens behave like wild specimens, does not change over time and is therefore free of learning. The predatory species as it is confronted for the first time with acorns as a food source could not foresee the higher concentration of tannins at that end and opened the acorns at the apical part because it had easier access. The benefits of mutualism for acorn species have already been shown. But what benefit do embryo-preserving rodents get from using more energy to open acorns at the basal end rather than the more easily accessible apical end? Maybe just resources? (Kellner et al. 2016 ; Pesendorfer et al. 2016 ; Cao et al. 2018b ; Moore and Dittel 2020 ), but they would obtain them more easily by opening the acorns apically, despite the presence of tannins. Perhaps in the future there will be acorn-producing trees to feed their descendants? But this is a subject for future research. The acorn species most appreciated by all rodents ( Q. ilex ) is also the most abundant in the study region. Does it obtain this success because it is the species most dispersed by mutualistic rodents in this region? This will also be the subject of future studies. Conclusions The two groups of mutualistic and antagonistic rodents show the same preferences for the different acorn species offered. The acorn species with higher nutrient concentration are more appreciated for consumption and are also more transported to appropriate places to germinate by the two mutualistic species ( A. sylvaticus and M. spretus ) but are destroyed by the predator species ( M. arvalis ). Species with lower nutrient concentrations, characteristic to avoid predation, are not consumed, but neither are they transported. Declarations Acknowledgements We thank Junta de Castilla y León for permission granted to carry out this research, in their mission to safeguard ethics in animal welfare during their handling. We thank Ángel José Álvarez Barcia, the director of S.I.B.A. (Servicio de Investigación y Bienestar Animal) at the University of Valladolid, for his advice on correct treating and handling of rodents. Funding This study was partially supported by the VA002A07 and VA035G18 projects granted by the Junta de Castilla y León to J.M. Del Arco. Compliance with ethical standards Conflicts of Interest: The authors declare that they have no conflicts of interest. Ethical approval: All applicable international, national, and/or institutional guidelines for the care and use of animals were followed. All procedures performed in this study involving animals were in accordance with the ethical standards of the institution at which the studies were conducted (CEEBA University of Valladolid, Spain). The experimental procedures were designed in accordance with the requirements of replacement, reduction and refinement. Data Availability statement The datasets generated during the current study are available in the [https://uvadoc.uva.es/handle/10324/61865] repository. All data generated during this study are included in this published article and its supplementary information files. References Bieberich J, Lauerer M, Aas G (2016) Acorns of introduced Quercus rubra are neglected by European Jay but spread by mice. Ann For Res 59:249–258 Bogdziewicz M, Crone EE, Zwolak R (2020) Do benefits of seed dispersal and caching by scatterhoarders outweigh the costs of predation? An example with oaks and yellow-necked mice. J Ecol 108:10091018. 10.1111/1365-2745.13307 Cao L, Yan C, Wang B (2018a) Differential seed mass selection on hoarding decisions among three sympatric rodents. Behav Ecol Sociobiol 72:1–9. doi.org/10.1007/s00265-018-2576-5 Cao L, Yan C, Wang B, Zhengyu W, Zhang H, Geng Y, Chen J, Zhang Z (2018b) Risk of cache pilferage determines hoarding behavior of rodents and seed fate. Behav Ecol 29:984–991. 10.1093/beheco/ary040 Chang G, Zhang Z (2014) Functional traits determine formation of mutualism and predation interactions in seed-rodent dispersal system of a subtropical forest. Act Oecol 55:43–50 Correa M, Uriarte M (2013) Integrating frugivory and animal movement: a review of the evidence and implications for scaling seed dispersal. Biol Rev 88:255–272. 10.1111/j.1469-185X.2012.00250.x De Blas C, Mateos GG, Rebollar PG (2003) Spanish Foundation for the development of animal nutrition. Madrid, Spain Del Arco JM, Beltrán D, Martínez-Ruiz C (2018) Risk for the natural regeneration of Quercus species due to the expansion of rodent species (Microtus arvalis). Behav Ecol Sociobiol 72:1–8. doi.org/10.1007/s00265-018-2575-6 Del Arco JM, Del Arco S (2022) Partial Consumption of Acorns by Some Rodents Leads Their Relationship with Oaks Species Towards Mutualism. Ecol Evolutionary Biology 7(1):1–6. 10.11648/j. eeb.20220701.11 Galetti M, Guevara R, Galbiati LA, Neves CL, Rodarte RR, Mendes CP (2015) Seed Predation by Rodents and Implications for Plant Recruitment in Defaunated Atlantic Forests. Biotropica 47:521–525 Gleditsch JM, Hruska AM, Foster JT (2017) Connecting Resource Tracking by Frugivores to Temporal Variation in Seed Dispersal Networks. Ecol Evol 5:98. https://doi.org/10.3389/fevo.2017.00098 Gómez JM, Schupp EW, Jordano P (2019) Synzoochory: the ecological and evolutionary relevance of a dual interaction. Biol rev 94:874–902. https://doi.org/10.1111/brv.12481 Gong H, Tang C, Wang B (2015) Post-dispersal seed predation and its relations with seed traits: a thirty-species-comparative study. Plant Species Biol 30:193–201. 10.1111/1442-1984.12051 Gu H, Zhao Q, Zhang Z (2017) Does scatter-hoarding of seeds benefit cache owners or pilferers? Integrat Zool 12:477–488 Hou XG, Yi XF, Yang YQ, Liu WJ (2010) Acorn germination and seedling survival of Q. variabilis : effects of cotyledon excision. Ann For Sci 67:711. 10.1051/forest/2010036 Huang Z, Wang Y, Zhang H, Wu F, Zhang Z (2011) Behavioural responses of sympatric rodents to complete pilferage. Anim Behav 81:831–836 Kang H, Chang M, Liu S, Chao Z, Zhang X, Wang D (2020) Rodent-mediated plant community competition: what happens to the seeds after entering the adjacent stands? For Ecosist 7:1–14. https://doi.org/10.1186/s40663-020-00270-z Kellner KF, Lichti NI, Swihart RK (2016) Midstory removal reduces effectiveness of oak (Quercus) acorn dispersal by small mammals in the Central Hardwood Forest region. For Ecol Manag 375:182–190 Lai X, Guo C, Xiao Z (2014) Trait-mediated seed predation, dispersal and survival among frugivore-dispersed plants in a fragmented subtropical forest, Southwest China. Integrat Zool 9:246–254 Lichti NI, Steele MA, Swihart RK (2015) Seed fate and decision-making processes in scatter-hoarding rodents Biol Rev 92: 474–504. doi: 101111/brv12240 Lichti NI, Steele MA, Swihar RK (2017) Seed fate and decision-making processes in scatter-hoarding rodents. Biol Rev 92:474–504. 10.1111/brv.12240 Lichti NI, Dalgleish HJ, Steele MA (2020) Interactions among Shade, Caching Behavior, and Predation Risk May Drive Seed Trait Evolution in Scatter-Hoarded Plants. Diversity 12 :416. 10.3390/d12110416 Lu J, Zhang Z (2004) Effects of habitat and season on removal and hoarding of seeds of wild apricot (Prunus armeniaca) by small rodents. Acta Oecol 26:247–254 Luque-Larena JJ, Mougeot F, Viñuela J, Jareño D, Arroyo L, Lambin X, Arroyo B (2013) Recent large-scale range expansion and outbreaks of the common vole (Microtus arvalis) in NW Spain. Basic Appl Ecol 14:432–441. https://doi.org/10.1016/j.baae.2013.04.006 Martínez-Baroja L, Pérez-Camacho L, Villar-Salvador P, Rebollo S, Leverkus AB, Pesendorfer MB, Molina-Morale M, Castro J, Rey-Benayas JM (2021) Caching territoriality and site preferences by a scatter-hoarder drive the spatial pattern of seed dispersal and affect seedling emergence. J Ecol 109:2342–2353. 10.1111/1365-2745.13642 Merceron NR, Langhe A, Dubois H, Garin O, Gerarts F, Jacquemin F, Balligand B, Otjacques M, Sabbe T, Servranckx M, Wautelet S, Kremer A, Porté AJ, Monty A (2017) Removal of acorns of the alien oak Quercus rubra on the ground by scatter-hoarding animals in Belgian forests. Biotechnol Agron Soc Environ 21:127–130 Michaux JR, Magnanou E, Paradis E, Nieberding C, Libois R (2003) Mitochondrial phylogeography of the Woodmouse ( Apodemus sylvaticus ) in the Western Paleoartic region. Molec Ecol 12:685–697 Mittelman P, Pires AS, Fernandez FAS (2021) The intermediate dispersal hypothesis: seed dispersal is maximized in areas with intermediate usage by hoarders. Plant Ecol 222:221–231. https://doi.org/10.1007/s11258-020-01100-6 Moore CM, Dittel JW (2020) On mutualism, models, and masting: the effects of seed-dispersing animals on the plants they disperse. J Ecol 108:1775–1783 Muñoz A, Bonal R (2007) Rodents change acorn dispersal behaviour in response to ungulate presence. Oikos 116:1631–1638 Muñoz A, Bonal R (2008a) Seed choice by rodents: learning or inheritance? Behav Ecol Sociobiol 62:913–922. 10.1007/s00265-007-0515-y Muñoz A, Bonal R (2008b) Are you strong enough to carry that seed? Seed size/body size ratios influence seed choices by rodents. Anim Behav 76:709–715 Muñoz A, Bonal R, Díaz M (2009) Ungulates, rodents, shrubs: interactions in a diverse Mediterranean ecosystem. Basic Appl Ecol 10:151–160 Muñoz A, Bonal R (2011) Linking seed dispersal to cache protection strategies. J Ecol 99:1016–1025 Palomo LJ, Justo ER, Vargas JM (2008) Mus spretus ( Rodentia: Muridae ). Mann Sp 840:1–10 Paz A, Jareño D, Arroyo L, Viñuela J, Arroyo B, Mougeot F, Luque-Larena JJ, Fargallo JA (2012) Avian predators as a biological control system of common vole (Microtus arvalis) populations in North-Western Spain: experimental set-up and preliminary results. Pest Manag Sci 69:444–450. https://doi.org/10.1002/ps.3289 Perea R, San Miguel A, Gil L (2011) Leftovers in seed dispersal: ecological implications of partial seed consumption for oak regeneration. J Ecol 99:94–201 Perea R, San Miguel A, Martínez-Jauregui M, Valbuena-Carabaña M, Gil L (2012) Effects of seed quality and seed location on the removal of acorns and beechnuts. Eur J Forest Res 131:623–631. 10.1007/s10342-011-0536-y Perea R, Dirzo R, San Miguel A, Gil L (2016) Post-dispersal seed recovery by animals: is it a plant- or an animal-driven process? Oikos 125:1203–1210. doi:101111/oik02556 Pesendorfer MB, Sillett TS, Koenig WD, Morrison SA (2016) Scatter-hoarding corvids as seed dispersers for oaks and pines: A review of a widely distributed mutualism and its utility to habitat restoration. Am Ornithol Soc 118:215–237. 10.1650/CONDOR-15-125.1 Pinheiro J, Bates D (2000) Mixed-effects models in S and S-Plus. Springer, New York Pinheiro J, Bates D, DebRoy S, Sarkar D (2013) The R Development Core Team. nlme: linear and nonlinear mixed effects models. R Packag Version 3:1–108 Pons J, Pausas JG (2007) Rodent acorn selection in a Mediterranean oak landscape. Ecol Res 22:535–541. 10.1007/s11284-006-0053-5 Pulido FJ, Díaz M (2005) Regeneration of a Mediterranean oak: a whole-cycle approach. Ecoscience 12:92–102 Rosalino LM, Nóbrega F, Santos-Reis M, Teixeira G, Rebelo R (2013) Acorn selection by the wood mouse Apodemus sylvaticus: a semi-controlled experiment in a Mediterranean environment. Zoolog Sci 30(9):724–730. 10.2108/zsj.30.724 Sivy KJ, Ostoja SM, Schupp EW, Durham S (2011) Effects of rodent species, seed species, and predator cues on seed fate. Acta Oecol 37:321–328 Sokal RR, Rohlf FJ (1995) Biometry, 3rd edn. WH Freeman and Co, New York Steele MA, Knowles T, Bridle K, Simms EL (1993) Tannins and Partial Consumption of Acorns: Implications for Dispersal of Oaks by Seed Predators. Am Midl Nat 130:229–238 Wang B, Chen J, Corlett RT (2014a) Factors influencing repeated seed movements by scatter-hoarding rodents in an alpine forest. Scien Repor 4:1–6. 10.1038/srep04786 Wang Z, Cao L, Zhang Z (2014b) Seed traits and taxonomic relationships determine the occurrence of mutualisms versus seed predation in a tropical forest rodent and seed dispersal system. Integ Zool 9:309–319 Wang Z, Wang B, Yi X, Yan C, Cao L, Zhang Z (2018) Scatter-hoarding rodents are better pilferers than larder-hoarders. Anim Behav 141:151–159 Wang Z, Wang B, Yan C, Yuan S, Cao L (2020) Neighborhood effects on the tannin-related foraging decisions of two rodent species under semi-natural conditions. Integr Zool 15:569–577. 10.1111/1749-4877.12473 Wróbel A, Zwolak R (2017) Deciphering the effects of disperser assemblages and seed mass on patterns of seed dispersal in a rodent community. Integrat Zool 12:457–467. 10.1111/1749-4877.12265 Xiao Z, Krebs CJ (2015) Modeling the costs and benefits of seed scatterhoarding to plants. Ecosphere 6:1–8. https://doi.org/10.1890/ES14-00438.1 Xiao Z, Holyoak M, Krebs CJ, Huang X (2021) Palatability and profitability of co-occurring seeds alter indirect interactions among rodent-dispersed trees. Integrat Zool 17:206–216. 10.1111/1749-4877.12543 Yang Y, Yi X (2012) Partial acorn consumption by small rodents: implication for regeneration of white oak, Quercus mongolica. Plant Ecol 213:197–205. 10.1007/s11258-011-0016-y Yi X, Wang Z (2015) Context-dependent seed dispersal determines acorn survival of sympatric oak species. Plant Ecol 216:123–132. 10.1007/s11258-014-0421-0 Yi X, Wang Z (2016) The Importance of Cotyledons for Early-Stage Oak Seedlings Under Different Nutrient Levels: A Multi-species Study. J Plant Growth Regul 35:183–189. 10.1007/s00344-015-9516-7 Zhang M, Steele MA, Yi X (2013) Reconsidering the effects of tannin on seed dispersal by rodents: Evidence from enclosure and field experiments with artificial seeds. Behav Proc 100:200–207 Zhang H, Wang Z, Zeng Q, Chang G (2015) Mutualistic and predatory interactions are driven by rodent body size and seed traits in a rodent–seed system in warm-temperate forest in northern China. Wild Res 42:149–157 Zhang H, Yan C, Chang G, Zhang Z (2016a) Seed trait–mediated selection by rodents affects mutualistic interactions and seedling recruitment of co–occurring tree species. Oecol 180:475–484. 10.1007/s00442-015-3490-4 Zhang Z, Wang Z, Chang G, Yi X, Lu J, Xiao Z, Zhang H, Cao L, Wang F, Li H, Yan C (2016b) Trade-off between seed defensive traits and impacts on interaction patterns between seeds and rodents in forest ecosystems. Plant Ecol 217:253–265. 10.1007/s11258-016-0566-0 Zhang Y, Bartlow AW, Wang Z, Yi X (2018) Effects of tannins on population dynamics of sympatric seed-eating rodents: the potential role of gut tannin-degrading bacteria. Oecologia 187:667–678. doi.org/10.1007/s00442-018-4151-1 Zwolak R, Crone EE (2012) Quantifying the outcome of plant–granivore interactions. Oikos 121:20–27. 10.1111/j.1600-0706.2011.019849.x Zwolak R, Bogdziewicz M, Crone EE (2020) On the need to evaluate costs and benefits of synzoochory for plant populations. J Ecol 108:1784–1788. 10.1111/1365-2745.13463 Tables Table 1 Chemical composition. n Dry matter (%) Lignin Protein Fat Sugar Q. petraea 3 0.58 ± 0.02ac 0.47 ± 0.05a 3.87 ± 0.41a 11.51 ± 0.30a Q. ilex subsp. ba llot a 5 0.25 ± 0.02b 6.34 ± 0.22bc 8.54 ± 0.66b 13.18 ± 0.30a Q. pyrenaica 3 0.16 ± 0.02b 4.91 ± 0.11b 5.45 ± 0.11a 11.15 ± 0.57a Q. faginea 3 0.31 ± 0.07bc 4.94 ± 0.19b 4.76 ± 0.22a 16.58 ± 0.80b Q. suber Q. rubra 3 3 0.81 ± 0.11c 0.60 ± 0.02ac 25 ± 0.72c 6.66 ± 0.18bc 4.35 ± 0.27a 3.38 ± 0.18a 12.78 ± 1.64a 8.92 ± 0.22a Composition of acorns from different oaks species (mean ± SE). Different letters indicate significance differences between oaks species (Bonferroni test, p < 0.05) after significant one-way ANOVA tests (lignin: F(4, 12) = 21.28, p < 0.001; protein: F(4, 12) = 57.31, p< 0.001; fat: F(4, 12) = 16.29, p < 0.001; sugar: F(4, 12) = 6.88, p = 0.004) Table 2 Summary of the results from the linear mixed models testing the effects of oaks species, rodent species, and day, and their interaction on the number of acorns consumed, total mass of acorns (g) consumed per day, and mean mass of acorns (g) consumed per individual, in the laboratory experiment. The F values of the fixed factors and their significance ( p ) are presented. df F p Number of acorns Intercept 2325 1846.0430 <0.0001 Quercus species 2325 54.5766 <0.0001 Rodent species 2325 30.8616 <0.0001 Time 2325 0.2883 0.8857 Quercus*Rodent 2325 9.4229 <0.0001 Quercus*Time 2325 0.2626 0.9996 Rodent*Time 2325 0.2099 0.9893 Quercus*Rodent*Time 2325 0.0680 1.0000 df F p Daily acorn mass Intercept 712 2469.6575 <0.0001 Quercus species 712 549.6110 <0.0001 Rodent species 712 360.8723 <0.0001 Time 712 0.3293 0.8584 Quercus*Rodent 712 62.5560 <0.0001 Quercus*Time 712 0.6025 0.9126 Rodent*Time 712 1.1698 0.3147 Quercus*Rodent*Time 712 0.7617 0.8572 Supplementary Files DATAavailability.pdf Cite Share Download PDF Status: Under Review Version 1 posted Reviewers agreed at journal 14 Oct, 2023 Reviewers invited by journal 10 Oct, 2023 Editor assigned by journal 08 Oct, 2023 First submitted to journal 04 Oct, 2023 You are reading this latest preprint version Research Square lets you share your work early, gain feedback from the community, and start making changes to your manuscript prior to peer review in a journal. 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Also discoverable on Platform About Our Team In Review Editorial Policies Advisory Board Help Center Resources Author Services Accessibility API Access RSS feed Manage Cookie Preferences © Research Square 2026 | ISSN 2693-5015 (online) Privacy Policy Terms of Service Do Not Sell My Personal Information {"props":{"pageProps":{"initialData":{"identity":"rs-3410260","acceptedTermsAndConditions":true,"allowDirectSubmit":false,"archivedVersions":[],"articleType":"Research Article","associatedPublications":[],"authors":[{"id":239070221,"identity":"df63ee73-e967-4316-bec2-4d3dafa31638","order_by":0,"name":"Sergio Del Arco","email":"","orcid":"","institution":"University of Salamanca: Universidad de Salamanca","correspondingAuthor":false,"submittingAuthor":false,"prefix":"","firstName":"Sergio","middleName":"Del","lastName":"Arco","suffix":""},{"id":239070222,"identity":"c0b4f89e-2056-4621-83f6-3895eb4756c0","order_by":1,"name":"Jose María Del Arco","email":"data:image/png;base64,iVBORw0KGgoAAAANSUhEUgAAAZAAAAAyAQMAAABI0h/eAAAABlBMVEX///8AAABVwtN+AAAACXBIWXMAAA7EAAAOxAGVKw4bAAAA6UlEQVRIiWNgGAWjYBACxgYIzQPl25CuJY10Ww8TVsLc3v7w0Y2aezL8DczPPvyoOJ+44fgBxoc/8Dms54yxcc6xYh6JA2zGM3vO3E7ccCaB2ZgHn5YZOWzSOWwJPAwHGIyZGdtuJ85sSGCTxucwxhnpz3/n/EvgkT/A/hmo5VzizP4H7D/xOmxGghlzblsCj8EBHpAtBxL7JRLYGPA6DOgX6dy+BB7DwzzFQE6ycb/Ew2ZpfFoMgSH2Oedbgr3c8fbNDD8q7GTb+JMPfsTnMMMGGIsZYXMDFoUIII9XdhSMglEwCkYBCAAAjvZKQj4O1NQAAAAASUVORK5CYII=","orcid":"https://orcid.org/0000-0002-6439-2184","institution":"University of Valladolid: Universidad de Valladolid","correspondingAuthor":true,"submittingAuthor":false,"prefix":"","firstName":"Jose","middleName":"María Del","lastName":"Arco","suffix":""}],"badges":[],"createdAt":"2023-10-04 11:12:53","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-3410260/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-3410260/v1","draftVersion":[],"editorialEvents":[],"editorialNote":"","failedWorkflow":false,"files":[{"id":44598281,"identity":"71127370-a0ca-48b5-a243-fce3235462ce","added_by":"auto","created_at":"2023-10-13 20:36:22","extension":"png","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":44053,"visible":true,"origin":"","legend":"\u003cp\u003eChemical affinity. Cluster analysis for six acorn species based on seed composition. Qi = \u003cem\u003eQ. ilex\u003c/em\u003e; Qr = \u003cem\u003eQ. rubra\u003c/em\u003e; Qp = \u003cem\u003eQ.\u003c/em\u003e \u003cem\u003epetraea\u003c/em\u003e; Qpy = \u003cem\u003eQ. pyrenaica\u003c/em\u003e; Qs = \u003cem\u003eQ. suber\u003c/em\u003e; Qf = \u003cem\u003eQ. faginea\u003c/em\u003e.\u003c/p\u003e","description":"","filename":"1.png","url":"https://assets-eu.researchsquare.com/files/rs-3410260/v1/41db572bd7196282b3ea1665.png"},{"id":44598283,"identity":"16c34d2b-d9bc-42aa-8d80-989e2b408272","added_by":"auto","created_at":"2023-10-13 20:36:22","extension":"png","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":84602,"visible":true,"origin":"","legend":"\u003cp\u003eConsumption of acorns.\u003cstrong\u003e \u003c/strong\u003eMean number of acorns attacked daily (a), mean mass of acorns consumed per day(b)and mean mass per acorn consumed daily (c) by each rodent species in each acorn class.\u003c/p\u003e\n\u003cp\u003eQi = \u003cem\u003eQ. ilex\u003c/em\u003e; Qr = \u003cem\u003eQ. rubra\u003c/em\u003e; Qp = \u003cem\u003eQ. petraea\u003c/em\u003e; Qpy = \u003cem\u003eQ. pyrenaica\u003c/em\u003e; Qs = \u003cem\u003eQ. suber\u003c/em\u003e; Qf = \u003cem\u003eQ. faginea\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003eMs = \u003cem\u003eM. spretus\u003c/em\u003e; As = \u003cem\u003eA. sylvaticus\u003c/em\u003e; Ma = \u003cem\u003eM. arvalis\u003c/em\u003e.\u003c/p\u003e","description":"","filename":"2.png","url":"https://assets-eu.researchsquare.com/files/rs-3410260/v1/a85b20bb05b7d8be28ad894c.png"},{"id":44598282,"identity":"e4c847b1-4b39-4e13-98f0-021eca1513dd","added_by":"auto","created_at":"2023-10-13 20:36:22","extension":"png","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":21989,"visible":true,"origin":"","legend":"\u003cp\u003eCover thickness.\u003cstrong\u003e \u003c/strong\u003eShell thickness of the acorns from the six oaks species (mm).\u003c/p\u003e","description":"","filename":"3.png","url":"https://assets-eu.researchsquare.com/files/rs-3410260/v1/a633f2c08c2e64fa623d6cb3.png"},{"id":44599699,"identity":"675260ff-aa29-43ed-ba9f-cc050c021211","added_by":"auto","created_at":"2023-10-13 20:44:27","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":395938,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-3410260/v1/55b38c4d-159b-4003-ac6e-d04e2f0324f6.pdf"},{"id":44598284,"identity":"0477ccf5-8072-4ad1-b473-4db2a95e9f73","added_by":"auto","created_at":"2023-10-13 20:36:22","extension":"pdf","order_by":1,"title":"","display":"","copyAsset":false,"role":"supplement","size":3307369,"visible":true,"origin":"","legend":"","description":"","filename":"DATAavailability.pdf","url":"https://assets-eu.researchsquare.com/files/rs-3410260/v1/3098c16d8020e7b6447687af.pdf"}],"financialInterests":"","formattedTitle":"Preferences of rodent scatter-hoard and predators determine dissemination oak species.","fulltext":[{"header":"Significance Statement","content":"\u003cp\u003eThe relationship between rodents and oak species has been termed antagonistic or mutualistic, depending on the outcome. In the antagonistic relationship, one of the species is harmed because it loses its seeds. In the mutualistic relationship, both species obtain benefits. We used rodent species representative of both relationships. We have provided these rodents with several different acorn species to observe if their different behavior during acorn handling causes differences in the choice of these acorn species.\u003c/p\u003e\n\u003cp\u003eThe selected acorn species exhibit different characteristics. Some seem intended to avoid predation, but others seem intended to attract the attention of rodents. We have observed that these differences lead to different behaviors in the choice of acorns.\u003c/p\u003e"},{"header":"Introduction","content":"\u003cp\u003eThe relationship between scatter-hoarding rodents and oak species varies in their outcomes (Mu\u0026ntilde;oz and Bonal \u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e2011\u003c/span\u003e; Yi and Wang \u003cspan citationid=\"CR57\" class=\"CitationRef\"\u003e2015\u003c/span\u003e; Gu et al. \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e2017\u003c/span\u003e; Lichti et al. \u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e2017\u003c/span\u003e; Bogdziewicz et al. \u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Mittelman et al. \u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). At one end of the ranging is antagonism, a relationship called predation which the species rodent takes resources from species plant that they suffer costs (Galetti et al. \u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e2015\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eIn this predation relationship, rodents obtain resources as benefits and plants pay costs. These costs involve the loss of seeds through their destruction by rodent ingestion. As a representative of this group, we present \u003cem\u003eMicrotus arvalis\u003c/em\u003e. This rodent species devours acorns by opening them at the apical end where the embryo is inserted. This organ is the first element of the acorn to be destroyed.\u003c/p\u003e \u003cp\u003eAt the other end of the ranging is mutualism. In this relationship, the two participating species obtain benefits, but both must contribute resources for the relationship to remain in equilibrium (Zhang et al. \u003cspan citationid=\"CR60\" class=\"CitationRef\"\u003e2015\u003c/span\u003e; Zhang et al. \u003cspan citationid=\"CR61\" class=\"CitationRef\"\u003e2016a\u003c/span\u003e; Moore and Dittel \u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). In mutualism, rodents obtain resources by ingesting whole acorns or part of the cotyledons of the acorns they attack (Cao et al. \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2018a\u003c/span\u003e; Wr\u0026oacute;bel and Zwolak \u003cspan citationid=\"CR53\" class=\"CitationRef\"\u003e2017\u003c/span\u003e; Moore and Dittel \u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). The costs they bear is the energy they must expend to transport the acorns to places safe for themselves. Normally, these rodents do not consume acorns in situ because their predators are waiting for them at the acorn concentration sites under the acorn-producing trees. They transport the acorns to protected places where they make numerous caches (Lichti et al. \u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e2015\u003c/span\u003e; Wang et al. \u003cspan citationid=\"CR51\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; Mittelman et al. \u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). Some of these caches are forgotten (Lichti et al. \u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Mart\u0026iacute;nez-Baroja et al. \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). Another cost is that they must expend energy to open acorns and remove their shells. Plants in this mutualistic relationship have the cost of losing acorns to rodent ingestion, to make acorns from high nutrient concentrations, synthesis of costly substances such as tannins and thick coatings. However, the benefits they obtain are that part of their acorns are transported by rodents to protected places under bushes where they are buried. There they gain humidity which favors their germination ((Perea et al. \u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e2011\u003c/span\u003e; Yang and Yi \u003cspan citationid=\"CR56\" class=\"CitationRef\"\u003e2012\u003c/span\u003e). Both species obtain benefits but pay costs. As representatives of mutualism, we present two species: \u003cem\u003eMus spretus\u003c/em\u003e and \u003cem\u003eApodemus sylvaticus\u003c/em\u003e. These two rodent species present a peculiar behavior during the acorn consumption process. This behavior was observed independently by two research groups (Perea et al. \u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e2011\u003c/span\u003e and Del Arco et al. \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). These two species start most of the acorns they process at the basal end away from the embryo and although they consume a variable part of the cotyledons (Yi and Wang \u003cspan citationid=\"CR58\" class=\"CitationRef\"\u003e2016\u003c/span\u003e) they leave for last the apical end with the embryo intact. This behavior allows some acorns (or at least embryos) to escape predation. Perea et al. \u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e2011\u003c/span\u003e and Yang and Yi \u003cspan citationid=\"CR56\" class=\"CitationRef\"\u003e2012\u003c/span\u003e, found that acorns with \u0026frac34; of the cotyledons consumed if they retain the embryo germinate. For this reason, these authors consider that these two species show a behavior that tends to conserve the embryo. This behavior is innate and not learned (Del Arco et al. \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; Del Arco and Del Arco \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). It is also an active behavior (Mu\u0026ntilde;oz and Bonal \u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e2008a\u003c/span\u003e) as these mutualistic species use the basal end to open the acorns, consuming more energy even though it is easier to open them from the narrower distal end. The origin of the partial consumption of acorns may be in the size of the acorns in relation to the body size of the rodent (Mu\u0026ntilde;oz and Bonal \u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e2008b\u003c/span\u003e). The origin of the conservative behavior of the embryo starting the acorns at the basal end may be that the plant species place high concentrations of tannins at the distal end where the embryo is to deter rodents from ingesting them (Steele et al. \u003cspan citationid=\"CR48\" class=\"CitationRef\"\u003e1993\u003c/span\u003e; Hou et al. \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e2010\u003c/span\u003e; Wang et al. \u003cspan citationid=\"CR52\" class=\"CitationRef\"\u003e2020\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eZwolak and Crone (\u003cspan citationid=\"CR64\" class=\"CitationRef\"\u003e2012\u003c/span\u003e), Bogdziewicz et al. (\u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2020\u003c/span\u003e) and Zwolak et al. (\u003cspan citationid=\"CR65\" class=\"CitationRef\"\u003e2020\u003c/span\u003e) proposed a mathematical model to assess whether the outcome of scatter-hoarding relationship is antagonism or mutualism (Chang and Zhang \u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e2014\u003c/span\u003e; Gleditsch et al. \u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e2017\u003c/span\u003e). When the benefits of transportation and burial are greater than the costs caused by predation during this process of scatter-hoarding, we can call the relationship mutualistic. To evaluate the cost of dispersal, acorn loss due to predation is typically included, however, not every acorn that is attacked loses its ability to germinate. The usual calculations carried out do not include the beneficial effect of partial acorn consumption, which significantly reduces these costs (Del Arco et al. \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; Del Arco and Del Arco \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e2022\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eWe have chosen six species of acorns to feed to the three rodent species. There are species of acorns that possess structures and characteristics associated with defense against predation, such as high concentrations of tannins and thick coatings that lengthen handling times. But there are other characteristics that are more associated with attracting the attention of rodents (Sivy et al. \u003cspan citationid=\"CR46\" class=\"CitationRef\"\u003e2011\u003c/span\u003e; Zhang et al. \u003cspan citationid=\"CR62\" class=\"CitationRef\"\u003e2016b\u003c/span\u003e; Xiao et al. \u003cspan citationid=\"CR55\" class=\"CitationRef\"\u003e2021\u003c/span\u003e), such as high nutrient concentrations or thin coatings.\u003c/p\u003e \u003cp\u003eThe main goal of this study is to test the food preferences of the two groups of rodents that show different behavior during the handling of acorns (preservatives (\u003cem\u003eA. sylvaticus\u003c/em\u003e and \u003cem\u003eM. spretus\u003c/em\u003e) and embryo destroyers (\u003cem\u003eM. arvalis\u003c/em\u003e)) (Pons and Pausas \u003cspan citationid=\"CR43\" class=\"CitationRef\"\u003e2007\u003c/span\u003e; Rosalino et al. \u003cspan citationid=\"CR45\" class=\"CitationRef\"\u003e2013\u003c/span\u003e). We want to know if there are differences between the two groups in the choice of acorn species for their diet (G\u0026oacute;mez et al. \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). The different physical and chemical characteristics exhibited by acorns may condition choice in each group (Gong et al. \u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e2015\u003c/span\u003e; Perea et al. \u003cspan citationid=\"CR39\" class=\"CitationRef\"\u003e2016\u003c/span\u003e). We want to check if these differences are due to the different characteristics of the acorns.\u003c/p\u003e"},{"header":"Methods","content":"\u003cp\u003e\u003cem\u003eStudy system\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003eFor this study, we captured rodents of\u0026nbsp;three different rodent species,\u0026nbsp;\u003cem\u003eApodemus sylvaticus\u003c/em\u003e (Linnaeus, 1758) (wood mouse), \u003cem\u003eMus spretus\u003c/em\u003e (Lataste, 1883) (Algerian mouse), and \u003cem\u003eMicrotus arvalis\u003c/em\u003e (Pallas, 1778) (common vole).\u0026nbsp;The captured specimens of the three rodent species were placed in terrariums for reproduction. Nine descendants of each rodent species were selected for the experiment to eliminate any previous acorn consumption experience.\u003c/p\u003e\n\u003cp\u003eThey were placed in a fenced enclosure of 100 m\u003csup\u003e2\u003c/sup\u003e.\u0026nbsp;Inside we built\u0026nbsp;9\u0026nbsp;plots of 9 m\u003csup\u003e2\u003c/sup\u003e (3*3 m side). Each of them was isolated by placing on each side of the square a 3 m wide sheet of metal buried in the ground at a depth of 50 cm and a height from the ground of 1 m to prevent rodents from escaping by jumping. Under these conditions, the mice behaved as if they were in semi-wild conditions: building burrows, creating shallow storages, galleries, pantries, and caches to hide the acorns, which they moved around the\u0026nbsp;plot.\u0026nbsp;The 9 specimens of each species were placed in these plots successively, starting with \u003cem\u003eM. spretus\u003c/em\u003e and ending with \u003cem\u003eM. arvalis\u003c/em\u003e.\u003c/p\u003e\n\u003cp\u003eThe wood mouse\u0026nbsp;has inhabited the Iberian Peninsula since ancient times (Michaux \u003cem\u003eet al.\u003c/em\u003e 2003). Acorns of various \u003cem\u003eQuercus\u003c/em\u003e species are among its food sources and are abundant in the area it inhabits (Pulido and Diaz 2005; Mu\u0026ntilde;oz \u003cem\u003eet al\u003c/em\u003e. 2009). The\u0026nbsp;Algerian mouse\u0026nbsp;is of North African origin but has been present in parts of the Iberian Peninsula for a long time (Palomo \u003cem\u003eet al\u003c/em\u003e. 2008). It lives practically in the same habitat as the wood mouse\u0026nbsp;except in the northern mountains. It also consumes acorns from the \u003cem\u003eQuercus\u003c/em\u003e species (Mu\u0026ntilde;oz and Bonal 2007). These two species partially consume acorns, opening them at the basal end and preserving the embryo. For this reason, we include them in the mutualism group. A third rodent species, \u003cem\u003eMicrotus arvalis\u003c/em\u003e, has recently arrived in the region (Luque-Larena \u003cem\u003eet al\u003c/em\u003e. 2013; Paz \u003cem\u003eet al\u003c/em\u003e. 2012) and does not use acorns for its diet, but consumes them (Del Arco \u003cem\u003eet al\u003c/em\u003e. 2018).\u0026nbsp;Common vole\u0026nbsp;is abundant in central Europe. Until recently, its distribution area in the Iberian Peninsula has been confined to the northern mountains, where it can access fresh food such as soft green herbaceous plants. Due to the increase in the distribution area of irrigated crops, the common vole\u0026nbsp;has expanded to the center of the Iberian Peninsula, where it experiences phases of overpopulation,\u0026nbsp;and it is considered a recurrent crop pest (Luque-Larena \u003cem\u003eet al\u003c/em\u003e. 2013; Paz \u003cem\u003eet al\u003c/em\u003e. 2012). There are no references to this species feeding on acorns despite that the mountain pastures where it originally lives are surrounded by \u003cem\u003eQ. pyrenaica\u003c/em\u003e and \u003cem\u003eQ. petraea\u0026nbsp;\u003c/em\u003eforest formations, but the humidity of the area means that fresh vegetables are continuously available as food. This species is a risk for the oaks in the center of the Iberian Peninsula because due to phases of overpopulation or by the increasingly pronounced drought can exhaust their traditional food and seek new sources of resources in the acorns and this species is clearly a predator of these seeds\u0026nbsp;(Del Arco \u003cem\u003eet al\u003c/em\u003e. 2018).\u003c/p\u003e\n\u003cp\u003eTo feed the rodents and check their food preferences, we have selected six species of acorns from the environment where the rodents live because they may constitute their source of resources and maintain some kind of relationship with them.\u003c/p\u003e\n\u003cp\u003eAcorns from the following \u003cem\u003eQuercus\u003c/em\u003e species were collected: \u003cem\u003eQuercus ilex\u003c/em\u003e subsp. ballota (Desf.) Samp. (holm oak), \u003cem\u003eQ. faginea\u003c/em\u003e Lam. (Portuguese oak) at the site where the rodents were caught, \u003cem\u003eQ. pyrenaica\u0026nbsp;\u003c/em\u003eWilld. (Pyreneen oak), \u003cem\u003eQ. petraea\u003c/em\u003e (Matt.) Liebl. (sessile oak), in the northern mountains of the Iberian Peninsula where the common vole originally lived, \u003cem\u003eQ. suber\u003c/em\u003e L. (cork oak) in Salamanca (Spain), and \u003cem\u003eQ. rubra\u0026nbsp;\u003c/em\u003eL (American oak) (Bieberich \u003cem\u003eet al\u003c/em\u003e. 2016; Merceron \u003cem\u003eet al\u003c/em\u003e. 2017) in experimental reforestations in Galicia (Spain). The only non-native species is \u003cem\u003eQuercus. Rubra\u003c/em\u003e. We have chosen this species to test its integration in this system of relationships between rodents and oaks if it can adapt to the existing dispersal mechanisms in the area (Bieberich \u003cem\u003eet al\u003c/em\u003e. 2016; Merceron \u003cem\u003eet al\u003c/em\u003e. 2017).\u003c/p\u003e\n\u003cp\u003e\u003cem\u003eExperimental Procedures and Design\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003eNine specimens of the descendants of each rodent species were placed in isolated plots.\u0026nbsp;Every day, for five days each specimen had six acorns available to them from each of the \u003cem\u003eQuercus\u003c/em\u003e species used. During the experiment, each specimen\u0026nbsp;was offered\u0026nbsp;180 acorns, 30 from each of the six \u003cem\u003eQuercus\u003c/em\u003e species.\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eBefore being given to the rodents, each acorn was weighed and labeled to estimate the individual mass consumed of each acorn. The remains of the acorns from the previous day were removed and weighed to estimate their mass after consumption. To estimate the daily mass consumed of each acorn species, we calculated the sum of the quantities consumed of each acorn by each rodent species specimen for one day. We also estimated the mean mass consumed per acorn of each plant species by each rodent species.\u003c/p\u003e\n\u003cp\u003eSome acorns from each species were analyzed for their nutrient content (Table 1).\u0026nbsp;We analyzed the chemical composition of the acorns to see whether it influenced selection during consumption (Table 1).\u0026nbsp;A high lignin content is related to a higher tannin content (De Blas \u003cem\u003eet al\u003c/em\u003e. 2003), which, in turn, could have deterred rodents from eating acorns (Zhang \u003cem\u003eet al\u003c/em\u003e. 2013). On the contrary, the higher fat, protein, and sugar contents\u0026nbsp;are usually the characteristics that most attract rodents to ingest acorns\u0026nbsp;(Table 1).\u003c/p\u003e\n\u003cp\u003eTo study the similarity in acorn composition, we grouped the species with the most similar composition together (Figure 1) according to the\u0026nbsp;Czekanowski C = (2W/A+B)*100 index.\u003c/p\u003e\n\u003cp\u003eWhere W = the sum of the minimum values of each compound in the side-by-side comparison of the two species\u003c/p\u003e\n\u003cp\u003eA = the sum of the values of each component\u0026nbsp;in the first species\u0026nbsp;in the side-by-side comparison\u003c/p\u003e\n\u003cp\u003eB = the sum of the values of each component\u0026nbsp;in the second species\u0026nbsp;in the side-by-side comparison.\u003c/p\u003e\n\u003cp\u003eAnother group of acorns was set aside to study their shell thickness. A Mitutoyo caliper was used to measure thickness at three different points: the circular scar where the acorn is attached to the cupule, the periphery of this circle located at the base of the acorn, and the apical end of the acorn, where the shell covers the embryo.\u003c/p\u003e\n\u003cp\u003e\u003cem\u003eData Analyses\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003eThe possible effects of rodent species (wood mouse,\u0026nbsp;Algerian mouse,\u0026nbsp;and common vole), day (5 levels), \u003cem\u003eQuercus\u003c/em\u003e species (\u003cem\u003eQ. petraea, Q. ilex, Q. pyrenaica, Q. faginea, Q. suber, Q. rubra\u003c/em\u003e), and their interactions on the number and mass of acorns eaten per specimen, were analyzed using Linear Mixed Models (LMM) with the Restricted Maximum Likelihood method (REML). The specimens were treated as a random factor and the day as a repeated factor. Finally, working on the model matrix, contrasts were carried out to test differences between fixed factor levels (Pinheiro and Bates 2000). Consequently, the Bonferroni correction was used to adjust for the significance level for each t-test (Sokal and Rohlf 1995). The statistical calculations were implemented in the R software environment (version 2.15.3; Core Team R 2013), using the nlme package for LMM (Pinheiro \u003cem\u003eet al\u003c/em\u003e. 2013).\u003c/p\u003e"},{"header":"Results","content":"\u003cp\u003eThe LMM analysis showed a highly significant interaction between \u0026lsquo;species of acorns\u0026rsquo;, \u0026lsquo;species of rodent\u0026rsquo; and \u0026lsquo;number of acorns\u0026rsquo;, \u0026lsquo;daily acorn mass\u0026rsquo;, and \u0026lsquo;mean mass per acorn\u0026rsquo; consumed (Table 2), suggesting that rodent species show a preference for some species of acorns. The number of acorns attacked by each rodent species, the daily acorn mass, and the mean mass per acorn did not vary significantly in time. There were no changes in rodent feeding behavior during acorn consumption.\u0026nbsp;\u003c/p\u003e\n\u003cp\u003ePreference for acorn species can be seen in Figure 2. We show the mean number of acorns attacked daily by each mouse of each species (Fig 2a).\u0026nbsp;The two species with the highest number of acorns consumed per day are \u003cem\u003eQ. ilex\u003c/em\u003e and \u003cem\u003eQ. pyrenaica\u003c/em\u003e. In figure 2b, we show the daily mass consumed by mice of each rodent species. The highest mass of acorns consumed daily in the three rodent species corresponds to \u003cem\u003eQ. ilex\u003c/em\u003e acorns. In Figure 2c we present the mean mass per acorn consumed daily by each mouse of the three rodent species. Also in this variable, the highest mass consumed per acorn in the three rodent species belongs to\u0026nbsp;\u003cem\u003eQ. ilex\u003c/em\u003e.\u003cbr\u003eThe species with the lowest number of acorns consumed daily are \u003cem\u003eQ. suber\u0026nbsp;\u003c/em\u003eand \u003cem\u003eQ. rubra\u003c/em\u003e in the rodent species common vole and Algerian mouse. Wood mouse attacks a daily mean acorns of these species like rest of the acorn species. However, the daily mass consumed of \u003cem\u003eQ. suber\u003c/em\u003e and \u003cem\u003eQ. rubra\u003c/em\u003e acorns is very low in all three rodent species. The mean daily mass consumed per acorn is also very low in all three rodent species for \u003cem\u003eQ. rubra.\u003c/em\u003e Wood mouse shows more preference for \u003cem\u003eQ. suber\u003c/em\u003e than\u0026nbsp;\u003cem\u003eQ. rubra\u003c/em\u003e as shown by the mass consumed in both species.\u003cbr\u003e\u0026nbsp;We analyzed the chemical composition of the acorns to see whether it influenced selection during consumption. There were significant differences in acorn composition (Table 1).\u003c/p\u003e\n\u003cp\u003eFigure 1, we show a cluster analysis for the six acorn species based on seed composition. \u003cem\u003eQ. ilex\u0026nbsp;\u003c/em\u003eis the species that differs most in terms of composition from other species.\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eWe also verified whether shell thickness influenced selection. Figure 3 shows the mean shell thickness in the six plant species. The two most rejected species (\u003cem\u003eQ. suber\u0026nbsp;\u003c/em\u003eand \u003cem\u003eQ. rubra\u003c/em\u003e) have thicker shells.\u003c/p\u003e"},{"header":"Discussion","content":"\u003cp\u003eLooking at the results obtained, we have verified that the two groups of rodents' species of this study (preservers embryo (wood mouse and Algerian mouse) and embryo destructors Common vole) have the same preferences for acorns (Pons and Pausas \u003cspan citationid=\"CR43\" class=\"CitationRef\"\u003e2007\u003c/span\u003e; Rosalino et al. \u003cspan citationid=\"CR45\" class=\"CitationRef\"\u003e2013\u003c/span\u003e; G\u0026oacute;mez et al. \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). Both groups of rodents showed a preference for the \u003cem\u003eQ. ilex\u003c/em\u003e and \u003cem\u003eQ. pyrenaica\u003c/em\u003e acorns and consumed \u003cem\u003eQ. suber\u003c/em\u003e and \u003cem\u003eQ. rubra\u003c/em\u003e with less intensity (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). These two groups of acorn species differ in their physical and chemical characteristics (Gong et al. \u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e2015\u003c/span\u003e; Perea et al. \u003cspan citationid=\"CR39\" class=\"CitationRef\"\u003e2016\u003c/span\u003e). (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). The most appreciated ones have a higher concentration of nutrients, lower concentration of tannins (Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e) (Perea et al. \u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e2012\u003c/span\u003e) and thin shells (Huang et al. \u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e2011\u003c/span\u003e) (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003e). The least appreciated, on the contrary, high tannin concentrations, low in nutrients (Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e) and thick shells (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003e). \u003cem\u003eQ. ilex\u003c/em\u003e, which is the most appreciated, differs significantly from the rest according to the cluster analysis (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). Investing high proportions of protein, sugars and fats gives the acorns of this specie characteristics associated with intense consumption. It is a good candidate for obtaining resources during adverse periods (Lichti et al. \u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e2017\u003c/span\u003e). Why invest high concentration of resources and what benefits do they get in return? The answer to this question is probably to be found in the mutualistic relationship that this plant species has long maintained with Wood mouse and Algerian mouse. These two rodent species practices partial consumption of acorns, preserving the embryo, and transport their acorns to caches because they do not consume them in situ. It is possible that the relationship was initiated by the attraction of rodents to the appetizing characteristics of the acorns of this species (Xiao and Krebs \u003cspan citationid=\"CR54\" class=\"CitationRef\"\u003e2015\u003c/span\u003e; Bogdziewicz et al. \u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Moore and Dittel \u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Zwolak et al. \u003cspan citationid=\"CR65\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). For this reason, this species of acorns is the most transported to caches (Lu and Zhang \u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e2004\u003c/span\u003e; Lichti et al. \u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). This means that it is the most favored in the dispersal process (Correa and Uriarte \u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e2013\u003c/span\u003e; Wang et al. \u003cspan citationid=\"CR50\" class=\"CitationRef\"\u003e2014b\u003c/span\u003e; Kang et al. \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). But this acorn specie has a problem, which also attracts the attention of the predatory specie in this study (Common vole) more intensely than other acorn species with lower nutrient concentrations (Lichti et al. \u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e2020\u003c/span\u003e) (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). Therefore, most of their acorns are destroyed more intensely than in other less preferred species by this rodent specie (Common vole). The risk for this acorn specie is that the natural food of Common vole disappears due to continuous drought in the study area or recurrent high densities of these rodent, that they look for new alternative sources of resources and end up incorporating acorns to their usual diet (Luque-Larena et al. \u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e2013\u003c/span\u003e; Paz et al. \u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e2012\u003c/span\u003e). We have already seen that the acorns it destroys are a very appetizing food for this rodent. The dispersal process of these \u003cem\u003eQ. ilex\u003c/em\u003e acorns could be paralyzed by the presence of this rodent specie (Pesendorfer et al. \u003cspan citationid=\"CR40\" class=\"CitationRef\"\u003e2016\u003c/span\u003e; Del Arco et al. \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; Del Arco and Del Arco \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e2022\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eThere are other acorns species (\u003cem\u003eQ. suber\u003c/em\u003e and \u003cem\u003eQ. rubra)\u003c/em\u003e that are not appreciated for consumption by the two groups of rodents (mutualists and antagonists) (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). These species present defensive characteristics to avoid consumption. High concentrations of tannins (Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e) and thick coatings to prevent access (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003e) (Zhang et al. \u003cspan citationid=\"CR59\" class=\"CitationRef\"\u003e2013\u003c/span\u003e; Lai et al. \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2014\u003c/span\u003e; Wang et al. \u003cspan citationid=\"CR49\" class=\"CitationRef\"\u003e2014a\u003c/span\u003e\u003cspan citationid=\"CR50\" class=\"CitationRef\"\u003eb\u003c/span\u003e; Lichti et al. \u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e2017\u003c/span\u003e; Zhang et al. \u003cspan citationid=\"CR63\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). If they escape predation they can germinate and emerge, but they have the problem that there is no transport for these acorns. If they emerge, it is under the adult tree that has originated them, and this obviously increases intraspecific competition (Moore and Dittel \u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). The process of colonization of new sites will be paralyzed. They are not buried because this mission is carried out by dispersing rodents (wood mouse and Algerian mouse) that do not select them as a priority for food. These two less preferred species (\u003cem\u003eQ. suber\u003c/em\u003e and \u003cem\u003eQ. rubra)\u003c/em\u003e have high concentrations of tannins. This may corroborate the statements of Steele et al. (\u003cspan citationid=\"CR48\" class=\"CitationRef\"\u003e1993\u003c/span\u003e); the partial consumption of acorns and the preservation of the embryo is a behavior that may have been generated by the inclusion of high concentrations of tannins around the embryo during made of acorns. These bad-tasting tannins discourage rodents from ingesting the embryo end by forcing them to open the acorns at the basal end. If acorns with higher concentrations of tannins are rejected, so will be the end with high amounts of these substances. However, since three rodent species reject the same species of acorns possibly because of the presence of tannins, they would also avoid opening the acorns at the apical end where these substances are most concentrated, however, the predatory specie (Common vole) initiates the acorns at the apical end destroying the embryo, not preserving it (Del Arco et al. \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; Del Arco and Del Arco \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). Why do they show different behavior in this open acorn process? The answer may lie in Del Arco et al. (\u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e2018\u003c/span\u003e) and Del Arco and Del Arco (\u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). By these, the embryonic conservative behavior is embedded in the genetic code of the rodent species that practice it (\u003cem\u003eApodemus sylvaticus\u003c/em\u003e and \u003cem\u003eMus spretus\u003c/em\u003e). Captive-bred specimens behave like wild specimens, does not change over time and is therefore free of learning. The predatory species as it is confronted for the first time with acorns as a food source could not foresee the higher concentration of tannins at that end and opened the acorns at the apical part because it had easier access.\u003c/p\u003e \u003cp\u003eThe benefits of mutualism for acorn species have already been shown. But what benefit do embryo-preserving rodents get from using more energy to open acorns at the basal end rather than the more easily accessible apical end? Maybe just resources? (Kellner et al. \u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e2016\u003c/span\u003e; Pesendorfer et al. \u003cspan citationid=\"CR40\" class=\"CitationRef\"\u003e2016\u003c/span\u003e; Cao et al. \u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e2018b\u003c/span\u003e; Moore and Dittel \u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e2020\u003c/span\u003e), but they would obtain them more easily by opening the acorns apically, despite the presence of tannins. Perhaps in the future there will be acorn-producing trees to feed their descendants? But this is a subject for future research.\u003c/p\u003e \u003cp\u003eThe acorn species most appreciated by all rodents (\u003cem\u003eQ. ilex\u003c/em\u003e) is also the most abundant in the study region. Does it obtain this success because it is the species most dispersed by mutualistic rodents in this region? This will also be the subject of future studies.\u003c/p\u003e"},{"header":"Conclusions","content":"\u003cp\u003eThe two groups of mutualistic and antagonistic rodents show the same preferences for the different acorn species offered. The acorn species with higher nutrient concentration are more appreciated for consumption and are also more transported to appropriate places to germinate by the two mutualistic species (\u003cem\u003eA. sylvaticus\u003c/em\u003e and \u003cem\u003eM. spretus\u003c/em\u003e) but are destroyed by the predator species (\u003cem\u003eM. arvalis\u003c/em\u003e).\u003c/p\u003e \u003cp\u003eSpecies with lower nutrient concentrations, characteristic to avoid predation, are not consumed, but neither are they transported.\u003c/p\u003e"},{"header":"Declarations","content":"\u003cp\u003e\u003cstrong\u003eAcknowledgements\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eWe thank Junta de Castilla y Le\u0026oacute;n for permission granted to carry out this research, in their mission to safeguard ethics in animal welfare during their handling. We thank \u0026Aacute;ngel Jos\u0026eacute; \u0026Aacute;lvarez Barcia, the director of S.I.B.A. (Servicio de Investigaci\u0026oacute;n y Bienestar Animal) at the University of Valladolid, for his advice on correct treating and handling of rodents.\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eFunding\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThis study was partially supported by the VA002A07 and VA035G18 projects granted by the Junta de Castilla y Le\u0026oacute;n to J.M. Del Arco.\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eCompliance with ethical standards\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eConflicts of Interest: The authors declare that they have no conflicts of interest.\u003c/p\u003e\n\u003cp\u003eEthical approval: All applicable international, national, and/or institutional guidelines for the care and use of animals were followed. All procedures performed in this study involving animals were in accordance with the ethical standards of the institution at which the studies were conducted (CEEBA University of Valladolid, Spain). The experimental procedures were designed in accordance with the requirements of replacement, reduction and refinement.\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eData Availability statement\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThe datasets generated during the current study are available in the [https://uvadoc.uva.es/handle/10324/61865] repository.\u003cbr\u003eAll data generated during this study are included in this published article and its supplementary information files.\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\u003cli\u003e\u003cspan\u003eBieberich J, Lauerer M, Aas G (2016) Acorns of introduced Quercus rubra are neglected by European Jay but spread by mice. Ann For Res 59:249\u0026ndash;258\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eBogdziewicz M, Crone EE, Zwolak R (2020) Do benefits of seed dispersal and caching by scatterhoarders outweigh the costs of predation? An example with oaks and yellow-necked mice. J Ecol 108:10091018. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003e10.1111/1365-2745.13307\u003c/span\u003e\u003cspan address=\"10.1111/1365-2745.13307\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eCao L, Yan C, Wang B (2018a) Differential seed mass selection on hoarding decisions among three sympatric rodents. Behav Ecol Sociobiol 72:1\u0026ndash;9. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003edoi.org/10.1007/s00265-018-2576-5\u003c/span\u003e\u003cspan address=\"10.1007/s00265-018-2576-5\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eCao L, Yan C, Wang B, Zhengyu W, Zhang H, Geng Y, Chen J, Zhang Z (2018b) Risk of cache pilferage determines hoarding behavior of rodents and seed fate. Behav Ecol 29:984\u0026ndash;991. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003e10.1093/beheco/ary040\u003c/span\u003e\u003cspan address=\"10.1093/beheco/ary040\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eChang G, Zhang Z (2014) Functional traits determine formation of mutualism and predation interactions in seed-rodent dispersal system of a subtropical forest. Act Oecol 55:43\u0026ndash;50\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eCorrea M, Uriarte M (2013) Integrating frugivory and animal movement: a review of the evidence and implications for scaling seed dispersal. Biol Rev 88:255\u0026ndash;272. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003e10.1111/j.1469-185X.2012.00250.x\u003c/span\u003e\u003cspan address=\"10.1111/j.1469-185X.2012.00250.x\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eDe Blas C, Mateos GG, Rebollar PG (2003) Spanish Foundation for the development of animal nutrition. Madrid, Spain\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eDel Arco JM, Beltr\u0026aacute;n D, Mart\u0026iacute;nez-Ruiz C (2018) Risk for the natural regeneration of Quercus species due to the expansion of rodent species (Microtus arvalis). Behav Ecol Sociobiol 72:1\u0026ndash;8. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003edoi.org/10.1007/s00265-018-2575-6\u003c/span\u003e\u003cspan address=\"10.1007/s00265-018-2575-6\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eDel Arco JM, Del Arco S (2022) Partial Consumption of Acorns by Some Rodents Leads Their Relationship with Oaks Species Towards Mutualism. Ecol Evolutionary Biology 7(1):1\u0026ndash;6. 10.11648/j. eeb.20220701.11\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eGaletti M, Guevara R, Galbiati LA, Neves CL, Rodarte RR, Mendes CP (2015) Seed Predation by Rodents and Implications for Plant Recruitment in Defaunated Atlantic Forests. Biotropica 47:521\u0026ndash;525\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eGleditsch JM, Hruska AM, Foster JT (2017) Connecting Resource Tracking by Frugivores to Temporal Variation in Seed Dispersal Networks. Ecol Evol 5:98. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.3389/fevo.2017.00098\u003c/span\u003e\u003cspan address=\"10.3389/fevo.2017.00098\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eG\u0026oacute;mez JM, Schupp EW, Jordano P (2019) Synzoochory: the ecological and evolutionary relevance of a dual interaction. Biol rev 94:874\u0026ndash;902. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1111/brv.12481\u003c/span\u003e\u003cspan address=\"10.1111/brv.12481\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eGong H, Tang C, Wang B (2015) Post-dispersal seed predation and its relations with seed traits: a thirty-species-comparative study. Plant Species Biol 30:193\u0026ndash;201. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003e10.1111/1442-1984.12051\u003c/span\u003e\u003cspan address=\"10.1111/1442-1984.12051\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eGu H, Zhao Q, Zhang Z (2017) Does scatter-hoarding of seeds benefit cache owners or pilferers? Integrat Zool 12:477\u0026ndash;488\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eHou XG, Yi XF, Yang YQ, Liu WJ (2010) Acorn germination and seedling survival of \u003cem\u003eQ. variabilis\u003c/em\u003e: effects of cotyledon excision. Ann For Sci 67:711. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003e10.1051/forest/2010036\u003c/span\u003e\u003cspan address=\"10.1051/forest/2010036\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eHuang Z, Wang Y, Zhang H, Wu F, Zhang Z (2011) Behavioural responses of sympatric rodents to complete pilferage. Anim Behav 81:831\u0026ndash;836\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eKang H, Chang M, Liu S, Chao Z, Zhang X, Wang D (2020) Rodent-mediated plant community competition: what happens to the seeds after entering the adjacent stands? For Ecosist 7:1\u0026ndash;14. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1186/s40663-020-00270-z\u003c/span\u003e\u003cspan address=\"10.1186/s40663-020-00270-z\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eKellner KF, Lichti NI, Swihart RK (2016) Midstory removal reduces effectiveness of oak (Quercus) acorn dispersal by small mammals in the Central Hardwood Forest region. For Ecol Manag 375:182\u0026ndash;190\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eLai X, Guo C, Xiao Z (2014) Trait-mediated seed predation, dispersal and survival among frugivore-dispersed plants in a fragmented subtropical forest, Southwest China. Integrat Zool 9:246\u0026ndash;254\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eLichti NI, Steele MA, Swihart RK (2015) Seed fate and decision-making processes in scatter-hoarding rodents Biol Rev 92: 474\u0026ndash;504. doi: 101111/brv12240\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eLichti NI, Steele MA, Swihar RK (2017) Seed fate and decision-making processes in scatter-hoarding rodents. Biol Rev 92:474\u0026ndash;504. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003e10.1111/brv.12240\u003c/span\u003e\u003cspan address=\"10.1111/brv.12240\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eLichti NI, Dalgleish HJ, Steele MA (2020) Interactions among Shade, Caching Behavior, and Predation Risk May Drive Seed Trait Evolution in Scatter-Hoarded Plants. \u003cem\u003eDiversity 12\u003c/em\u003e:416. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003e10.3390/d12110416\u003c/span\u003e\u003cspan address=\"10.3390/d12110416\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eLu J, Zhang Z (2004) Effects of habitat and season on removal and hoarding of seeds of wild apricot \u003cem\u003e(Prunus armeniaca)\u003c/em\u003e by small rodents. Acta Oecol 26:247\u0026ndash;254\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eLuque-Larena JJ, Mougeot F, Vi\u0026ntilde;uela J, Jare\u0026ntilde;o D, Arroyo L, Lambin X, Arroyo B (2013) Recent large-scale range expansion and outbreaks of the common vole (Microtus arvalis) in NW Spain. Basic Appl Ecol 14:432\u0026ndash;441. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1016/j.baae.2013.04.006\u003c/span\u003e\u003cspan address=\"10.1016/j.baae.2013.04.006\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eMart\u0026iacute;nez-Baroja L, P\u0026eacute;rez-Camacho L, Villar-Salvador P, Rebollo S, Leverkus AB, Pesendorfer MB, Molina-Morale M, Castro J, Rey-Benayas JM (2021) Caching territoriality and site preferences by a scatter-hoarder drive the spatial pattern of seed dispersal and affect seedling emergence. J Ecol 109:2342\u0026ndash;2353. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003e10.1111/1365-2745.13642\u003c/span\u003e\u003cspan address=\"10.1111/1365-2745.13642\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eMerceron NR, Langhe A, Dubois H, Garin O, Gerarts F, Jacquemin F, Balligand B, Otjacques M, Sabbe T, Servranckx M, Wautelet S, Kremer A, Port\u0026eacute; AJ, Monty A (2017) Removal of acorns of the alien oak \u003cem\u003eQuercus rubra\u003c/em\u003e on the ground by scatter-hoarding animals in Belgian forests. Biotechnol Agron Soc Environ 21:127\u0026ndash;130\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eMichaux JR, Magnanou E, Paradis E, Nieberding C, Libois R (2003) Mitochondrial phylogeography of the Woodmouse (\u003cem\u003eApodemus sylvaticus\u003c/em\u003e) in the Western Paleoartic region. Molec Ecol 12:685\u0026ndash;697\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eMittelman P, Pires AS, Fernandez FAS (2021) The intermediate dispersal hypothesis: seed dispersal is maximized in areas with intermediate usage by hoarders. Plant Ecol 222:221\u0026ndash;231. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1007/s11258-020-01100-6\u003c/span\u003e\u003cspan address=\"10.1007/s11258-020-01100-6\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eMoore CM, Dittel JW (2020) On mutualism, models, and masting: the effects of seed-dispersing animals on the plants they disperse. J Ecol 108:1775\u0026ndash;1783\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eMu\u0026ntilde;oz A, Bonal R (2007) Rodents change acorn dispersal behaviour in response to ungulate presence. Oikos 116:1631\u0026ndash;1638\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eMu\u0026ntilde;oz A, Bonal R (2008a) Seed choice by rodents: learning or inheritance? Behav Ecol Sociobiol 62:913\u0026ndash;922. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003e10.1007/s00265-007-0515-y\u003c/span\u003e\u003cspan address=\"10.1007/s00265-007-0515-y\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eMu\u0026ntilde;oz A, Bonal R (2008b) Are you strong enough to carry that seed? Seed size/body size ratios influence seed choices by rodents. Anim Behav 76:709\u0026ndash;715\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eMu\u0026ntilde;oz A, Bonal R, D\u0026iacute;az M (2009) Ungulates, rodents, shrubs: interactions in a diverse Mediterranean ecosystem. Basic Appl Ecol 10:151\u0026ndash;160\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eMu\u0026ntilde;oz A, Bonal R (2011) Linking seed dispersal to cache protection strategies. J Ecol 99:1016\u0026ndash;1025\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003ePalomo LJ, Justo ER, Vargas JM (2008) \u003cem\u003eMus spretus\u003c/em\u003e (\u003cem\u003eRodentia: Muridae\u003c/em\u003e). Mann Sp 840:1\u0026ndash;10\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003ePaz A, Jare\u0026ntilde;o D, Arroyo L, Vi\u0026ntilde;uela J, Arroyo B, Mougeot F, Luque-Larena JJ, Fargallo JA (2012) Avian predators as a biological control system of common vole (Microtus arvalis) populations in North-Western Spain: experimental set-up and preliminary results. Pest Manag Sci 69:444\u0026ndash;450. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1002/ps.3289\u003c/span\u003e\u003cspan address=\"10.1002/ps.3289\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003ePerea R, San Miguel A, Gil L (2011) Leftovers in seed dispersal: ecological implications of partial seed consumption for oak regeneration. J Ecol 99:94\u0026ndash;201\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003ePerea R, San Miguel A, Mart\u0026iacute;nez-Jauregui M, Valbuena-Caraba\u0026ntilde;a M, Gil L (2012) Effects of seed quality and seed location on the removal of acorns and beechnuts. Eur J Forest Res 131:623\u0026ndash;631. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003e10.1007/s10342-011-0536-y\u003c/span\u003e\u003cspan address=\"10.1007/s10342-011-0536-y\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003ePerea R, Dirzo R, San Miguel A, Gil L (2016) Post-dispersal seed recovery by animals: is it a plant- or an animal-driven process? \u003cem\u003eOikos\u003c/em\u003e 125:1203\u0026ndash;1210. doi:101111/oik02556\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003ePesendorfer MB, Sillett TS, Koenig WD, Morrison SA (2016) Scatter-hoarding corvids as seed dispersers for oaks and pines: A review of a widely distributed mutualism and its utility to habitat restoration. Am Ornithol Soc 118:215\u0026ndash;237. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003e10.1650/CONDOR-15-125.1\u003c/span\u003e\u003cspan address=\"10.1650/CONDOR-15-125.1\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003ePinheiro J, Bates D (2000) Mixed-effects models in S and S-Plus. Springer, New York\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003ePinheiro J, Bates D, DebRoy S, Sarkar D (2013) The R Development Core Team. nlme: linear and nonlinear mixed effects models. R Packag Version 3:1\u0026ndash;108\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003ePons J, Pausas JG (2007) Rodent acorn selection in a Mediterranean oak landscape. Ecol Res 22:535\u0026ndash;541. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003e10.1007/s11284-006-0053-5\u003c/span\u003e\u003cspan address=\"10.1007/s11284-006-0053-5\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003ePulido FJ, D\u0026iacute;az M (2005) Regeneration of a Mediterranean oak: a whole-cycle approach. Ecoscience 12:92\u0026ndash;102\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eRosalino LM, N\u0026oacute;brega F, Santos-Reis M, Teixeira G, Rebelo R (2013) Acorn selection by the wood mouse Apodemus sylvaticus: a semi-controlled experiment in a Mediterranean environment. Zoolog Sci 30(9):724\u0026ndash;730. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003e10.2108/zsj.30.724\u003c/span\u003e\u003cspan address=\"10.2108/zsj.30.724\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eSivy KJ, Ostoja SM, Schupp EW, Durham S (2011) Effects of rodent species, seed species, and predator cues on seed fate. Acta Oecol 37:321\u0026ndash;328\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eSokal RR, Rohlf FJ (1995) Biometry, 3rd edn. WH Freeman and Co, New York\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eSteele MA, Knowles T, Bridle K, Simms EL (1993) Tannins and Partial Consumption of Acorns: Implications for Dispersal of Oaks by Seed Predators. Am Midl Nat 130:229\u0026ndash;238\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eWang B, Chen J, Corlett RT (2014a) Factors influencing repeated seed movements by scatter-hoarding rodents in an alpine forest. Scien Repor 4:1\u0026ndash;6. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003e10.1038/srep04786\u003c/span\u003e\u003cspan address=\"10.1038/srep04786\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eWang Z, Cao L, Zhang Z (2014b) Seed traits and taxonomic relationships determine the occurrence of mutualisms \u003cem\u003eversus\u003c/em\u003e seed predation in a tropical forest rodent and seed dispersal system. Integ Zool 9:309\u0026ndash;319\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eWang Z, Wang B, Yi X, Yan C, Cao L, Zhang Z (2018) Scatter-hoarding rodents are better pilferers than larder-hoarders. Anim Behav 141:151\u0026ndash;159\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eWang Z, Wang B, Yan C, Yuan S, Cao L (2020) Neighborhood effects on the tannin-related foraging decisions of two rodent species under semi-natural conditions. Integr Zool 15:569\u0026ndash;577. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003e10.1111/1749-4877.12473\u003c/span\u003e\u003cspan address=\"10.1111/1749-4877.12473\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eWr\u0026oacute;bel A, Zwolak R (2017) Deciphering the effects of disperser assemblages and seed mass on patterns of seed dispersal in a rodent community. Integrat Zool 12:457\u0026ndash;467. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003e10.1111/1749-4877.12265\u003c/span\u003e\u003cspan address=\"10.1111/1749-4877.12265\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eXiao Z, Krebs CJ (2015) Modeling the costs and benefits of seed scatterhoarding to plants. Ecosphere 6:1\u0026ndash;8. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1890/ES14-00438.1\u003c/span\u003e\u003cspan address=\"10.1890/ES14-00438.1\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eXiao Z, Holyoak M, Krebs CJ, Huang X (2021) Palatability and profitability of co-occurring seeds alter indirect interactions among rodent-dispersed trees. Integrat Zool 17:206\u0026ndash;216. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003e10.1111/1749-4877.12543\u003c/span\u003e\u003cspan address=\"10.1111/1749-4877.12543\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eYang Y, Yi X (2012) Partial acorn consumption by small rodents: implication for regeneration of white oak, Quercus mongolica. Plant Ecol 213:197\u0026ndash;205. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003e10.1007/s11258-011-0016-y\u003c/span\u003e\u003cspan address=\"10.1007/s11258-011-0016-y\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eYi X, Wang Z (2015) Context-dependent seed dispersal determines acorn survival of sympatric oak species. Plant Ecol 216:123\u0026ndash;132. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003e10.1007/s11258-014-0421-0\u003c/span\u003e\u003cspan address=\"10.1007/s11258-014-0421-0\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eYi X, Wang Z (2016) The Importance of Cotyledons for Early-Stage Oak Seedlings Under Different Nutrient Levels: A Multi-species Study. J Plant Growth Regul 35:183\u0026ndash;189. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003e10.1007/s00344-015-9516-7\u003c/span\u003e\u003cspan address=\"10.1007/s00344-015-9516-7\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eZhang M, Steele MA, Yi X (2013) Reconsidering the effects of tannin on seed dispersal by rodents: Evidence from enclosure and field experiments with artificial seeds. Behav Proc 100:200\u0026ndash;207\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eZhang H, Wang Z, Zeng Q, Chang G (2015) Mutualistic and predatory interactions are driven by rodent body size and seed traits in a rodent\u0026ndash;seed system in warm-temperate forest in northern China. Wild Res 42:149\u0026ndash;157\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eZhang H, Yan C, Chang G, Zhang Z (2016a) Seed trait\u0026ndash;mediated selection by rodents affects mutualistic interactions and seedling recruitment of co\u0026ndash;occurring tree species. Oecol 180:475\u0026ndash;484. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003e10.1007/s00442-015-3490-4\u003c/span\u003e\u003cspan address=\"10.1007/s00442-015-3490-4\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eZhang Z, Wang Z, Chang G, Yi X, Lu J, Xiao Z, Zhang H, Cao L, Wang F, Li H, Yan C (2016b) Trade-off between seed defensive traits and impacts on interaction patterns between seeds and rodents in forest ecosystems. Plant Ecol 217:253\u0026ndash;265. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003e10.1007/s11258-016-0566-0\u003c/span\u003e\u003cspan address=\"10.1007/s11258-016-0566-0\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eZhang Y, Bartlow AW, Wang Z, Yi X (2018) Effects of tannins on population dynamics of sympatric seed-eating rodents: the potential role of gut tannin-degrading bacteria. Oecologia 187:667\u0026ndash;678. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003edoi.org/10.1007/s00442-018-4151-1\u003c/span\u003e\u003cspan address=\"10.1007/s00442-018-4151-1\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eZwolak R, Crone EE (2012) Quantifying the outcome of plant\u0026ndash;granivore interactions. Oikos 121:20\u0026ndash;27. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003e10.1111/j.1600-0706.2011.019849.x\u003c/span\u003e\u003cspan address=\"10.1111/j.1600-0706.2011.019849.x\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eZwolak R, Bogdziewicz M, Crone EE (2020) On the need to evaluate costs and benefits of synzoochory for plant populations. J Ecol 108:1784\u0026ndash;1788. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003e10.1111/1365-2745.13463\u003c/span\u003e\u003cspan address=\"10.1111/1365-2745.13463\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003c/ol\u003e"},{"header":"Tables","content":"\u003cp\u003e\u003cstrong\u003eTable 1\u003c/strong\u003e Chemical composition.\u003c/p\u003e\n\u003ctable border=\"0\" cellspacing=\"0\" cellpadding=\"0\" width=\"567\"\u003e\n \u003ctbody\u003e\n \u003ctr\u003e\n \u003ctd width=\"19.54225352112676%\" rowspan=\"2\" valign=\"top\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"15.140845070422536%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003en\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"14.788732394366198%\" valign=\"top\"\u003e\n \u003cp\u003eDry\u0026nbsp;matter\u0026nbsp;(%)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"50.528169014084504%\" colspan=\"3\" valign=\"top\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"18.818380743982495%\" valign=\"top\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"18.38074398249453%\" valign=\"top\"\u003e\n \u003cp\u003eLignin\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"22.538293216630198%\" valign=\"top\"\u003e\n \u003cp\u003e\u0026nbsp; Protein\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"22.319474835886215%\" valign=\"top\"\u003e\n \u003cp\u003e\u0026nbsp; \u0026nbsp; \u0026nbsp;Fat\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"17.943107221006564%\" valign=\"top\"\u003e\n \u003cp\u003e\u0026nbsp; \u0026nbsp;Sugar\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"19.54225352112676%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003eQ.\u0026nbsp;petraea\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"15.140845070422536%\" valign=\"top\"\u003e\n \u003cp\u003e3\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"14.788732394366198%\" valign=\"top\"\u003e\n \u003cp\u003e0.58\u0026nbsp;\u0026plusmn;\u0026nbsp;0.02ac\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"18.133802816901408%\" valign=\"top\"\u003e\n \u003cp\u003e0.47\u0026nbsp;\u0026plusmn;\u0026nbsp;0.05a\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"17.95774647887324%\" valign=\"top\"\u003e\n \u003cp\u003e3.87\u0026nbsp;\u0026plusmn;\u0026nbsp;0.41a\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"14.43661971830986%\" valign=\"top\"\u003e\n \u003cp\u003e11.51\u0026nbsp;\u0026plusmn;\u0026nbsp;0.30a\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"19.54225352112676%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003eQ. ilex\u0026nbsp;\u003c/em\u003esubsp. \u003cem\u003eba\u003c/em\u003e\u003cem\u003ellot\u003c/em\u003e\u003cem\u003ea\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"15.140845070422536%\" valign=\"top\"\u003e\n \u003cp\u003e5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"14.788732394366198%\" valign=\"top\"\u003e\n \u003cp\u003e0.25\u0026nbsp;\u0026plusmn;\u0026nbsp;0.02b\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"18.133802816901408%\" valign=\"top\"\u003e\n \u003cp\u003e6.34\u0026nbsp;\u0026plusmn;\u0026nbsp;0.22bc\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"17.95774647887324%\" valign=\"top\"\u003e\n \u003cp\u003e8.54\u0026nbsp;\u0026plusmn;\u0026nbsp;0.66b\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"14.43661971830986%\" valign=\"top\"\u003e\n \u003cp\u003e13.18\u0026nbsp;\u0026plusmn;\u0026nbsp;0.30a\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"19.54225352112676%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003eQ.\u0026nbsp;pyrenaica\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"15.140845070422536%\" valign=\"top\"\u003e\n \u003cp\u003e3\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"14.788732394366198%\" valign=\"top\"\u003e\n \u003cp\u003e0.16\u0026nbsp;\u0026plusmn;\u0026nbsp;0.02b\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"18.133802816901408%\" valign=\"top\"\u003e\n \u003cp\u003e4.91\u0026nbsp;\u0026plusmn;\u0026nbsp;0.11b\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"17.95774647887324%\" valign=\"top\"\u003e\n \u003cp\u003e5.45\u0026nbsp;\u0026plusmn;\u0026nbsp;0.11a\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"14.43661971830986%\" valign=\"top\"\u003e\n \u003cp\u003e11.15\u0026nbsp;\u0026plusmn;\u0026nbsp;0.57a\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"19.54225352112676%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003eQ.\u0026nbsp;faginea\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"15.140845070422536%\" valign=\"top\"\u003e\n \u003cp\u003e3\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"14.788732394366198%\" valign=\"top\"\u003e\n \u003cp\u003e0.31\u0026nbsp;\u0026plusmn;\u0026nbsp;0.07bc\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"18.133802816901408%\" valign=\"top\"\u003e\n \u003cp\u003e4.94\u0026nbsp;\u0026plusmn;\u0026nbsp;0.19b\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"17.95774647887324%\" valign=\"top\"\u003e\n \u003cp\u003e4.76\u0026nbsp;\u0026plusmn;\u0026nbsp;0.22a\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"14.43661971830986%\" valign=\"top\"\u003e\n \u003cp\u003e16.58\u0026nbsp;\u0026plusmn;\u0026nbsp;0.80b\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"19.54225352112676%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003eQ.\u0026nbsp;suber\u003c/em\u003e\u003c/p\u003e\n \u003cp\u003e\u003cem\u003eQ. rubra\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"15.140845070422536%\" valign=\"top\"\u003e\n \u003cp\u003e3\u003c/p\u003e\n \u003cp\u003e3\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"14.788732394366198%\" valign=\"top\"\u003e\n \u003cp\u003e0.81\u0026nbsp;\u0026plusmn;\u0026nbsp;0.11c\u003c/p\u003e\n \u003cp\u003e0.60\u0026nbsp;\u0026plusmn;\u0026nbsp;0.02ac\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"18.133802816901408%\" valign=\"top\"\u003e\n \u003cp\u003e25\u0026nbsp;\u0026plusmn;\u0026nbsp;0.72c\u003c/p\u003e\n \u003cp\u003e6.66\u0026nbsp;\u0026plusmn;\u0026nbsp;0.18bc\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"17.95774647887324%\" valign=\"top\"\u003e\n \u003cp\u003e4.35\u0026nbsp;\u0026plusmn;\u0026nbsp;0.27a\u003c/p\u003e\n \u003cp\u003e3.38\u0026nbsp;\u0026plusmn;\u0026nbsp;0.18a\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"14.43661971830986%\" valign=\"top\"\u003e\n \u003cp\u003e12.78\u0026nbsp;\u0026plusmn;\u0026nbsp;1.64a\u003c/p\u003e\n \u003cp\u003e8.92\u0026nbsp;\u0026plusmn;\u0026nbsp;0.22a\u003c/p\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003c/tbody\u003e\n\u003c/table\u003e\n\u003cp\u003e\u003cspan style=\"text-align: inherit;\"\u003eComposition of acorns from different oaks species (mean \u0026plusmn; SE). Different letters indicate significance differences between oaks species (Bonferroni test, p \u0026lt; 0.05) after significant one-way ANOVA tests (lignin: F(4, 12) = 21.28, p \u0026lt; 0.001; protein: F(4, 12) = 57.31, p\u0026lt; 0.001; fat: F(4, 12) = 16.29, p \u0026lt; 0.001; sugar: F(4, 12) = 6.88, p = 0.004)\u003c/span\u003e\u003c/p\u003e\n\u003cp\u003e\u003cspan style=\"text-align: inherit;\"\u003e\u003cstrong\u003eTable 2\u0026nbsp;\u003c/strong\u003eSummary of the results from the linear mixed models testing the effects of oaks species, rodent species, and day, and their interaction on the number of acorns consumed, total mass of acorns (g) consumed per day, and mean mass of acorns (g) consumed per individual, in the laboratory experiment. The \u003cem\u003eF\u0026nbsp;\u003c/em\u003evalues of the fixed factors and their significance (\u003cem\u003ep\u003c/em\u003e) are presented.\u003c/span\u003e\u003c/p\u003e\n\u003ctable border=\"0\" cellspacing=\"0\" cellpadding=\"0\" width=\"415\"\u003e\n \u003ctbody\u003e\n \u003ctr\u003e\n \u003ctd width=\"36.47342995169082%\" valign=\"bottom\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"13.285024154589372%\" colspan=\"3\" valign=\"bottom\"\u003e\n \u003cp\u003e\u003cstrong\u003edf\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"15.942028985507246%\" colspan=\"2\" valign=\"bottom\"\u003e\n \u003cp\u003e\u003cstrong\u003e\u003cem\u003eF\u003c/em\u003e\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"34.29951690821256%\" colspan=\"2\"\u003e\n \u003cp\u003e\u003cstrong\u003e\u003cem\u003ep\u003c/em\u003e\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"36.47342995169082%\" valign=\"bottom\"\u003e\n \u003cp\u003e\u003cstrong\u003eNumber of acorns\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"13.285024154589372%\" colspan=\"3\" valign=\"bottom\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"15.942028985507246%\" colspan=\"2\" valign=\"bottom\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"34.29951690821256%\" colspan=\"2\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"36.47342995169082%\" valign=\"bottom\"\u003e\n \u003cp\u003e\u0026nbsp; \u0026nbsp; \u0026nbsp; Intercept\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"13.285024154589372%\" colspan=\"3\" valign=\"bottom\"\u003e\n \u003cp\u003e2325\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"15.942028985507246%\" colspan=\"2\" valign=\"bottom\"\u003e\n \u003cp\u003e1846.0430\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"34.29951690821256%\" colspan=\"2\"\u003e\n \u003cp\u003e\u0026lt;0.0001\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"36.47342995169082%\" valign=\"bottom\"\u003e\n \u003cp\u003e\u003cem\u003eQuercus\u003c/em\u003e species\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"13.285024154589372%\" colspan=\"3\" valign=\"bottom\"\u003e\n \u003cp\u003e2325\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"15.942028985507246%\" colspan=\"2\" valign=\"bottom\"\u003e\n \u003cp\u003e54.5766\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"34.29951690821256%\" colspan=\"2\"\u003e\n \u003cp\u003e\u0026lt;0.0001\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"36.47342995169082%\" valign=\"bottom\"\u003e\n \u003cp\u003e\u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp;Rodent species\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"13.285024154589372%\" colspan=\"3\" valign=\"bottom\"\u003e\n \u003cp\u003e2325\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"15.942028985507246%\" colspan=\"2\" valign=\"bottom\"\u003e\n \u003cp\u003e30.8616\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"34.29951690821256%\" colspan=\"2\"\u003e\n \u003cp\u003e\u0026lt;0.0001\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"36.47342995169082%\" valign=\"bottom\"\u003e\n \u003cp\u003e\u0026nbsp; \u0026nbsp; Time\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"13.285024154589372%\" colspan=\"3\" valign=\"bottom\"\u003e\n \u003cp\u003e2325\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"15.942028985507246%\" colspan=\"2\" valign=\"bottom\"\u003e\n \u003cp\u003e0.2883\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"34.29951690821256%\" colspan=\"2\"\u003e\n \u003cp\u003e0.8857\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"36.47342995169082%\" valign=\"bottom\"\u003e\n \u003cp\u003e\u0026nbsp; \u0026nbsp; \u0026nbsp; Quercus*Rodent\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"13.285024154589372%\" colspan=\"3\" valign=\"bottom\"\u003e\n \u003cp\u003e2325\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"15.942028985507246%\" colspan=\"2\" valign=\"bottom\"\u003e\n \u003cp\u003e9.4229\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"34.29951690821256%\" colspan=\"2\"\u003e\n \u003cp\u003e\u0026lt;0.0001\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"36.47342995169082%\" valign=\"bottom\"\u003e\n \u003cp\u003e\u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp;Quercus*Time\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"13.285024154589372%\" colspan=\"3\" valign=\"bottom\"\u003e\n \u003cp\u003e2325\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"15.942028985507246%\" colspan=\"2\" valign=\"bottom\"\u003e\n \u003cp\u003e0.2626\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"34.29951690821256%\" colspan=\"2\"\u003e\n \u003cp\u003e0.9996\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"36.47342995169082%\" valign=\"bottom\"\u003e\n \u003cp\u003e\u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp;Rodent*Time\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"13.285024154589372%\" colspan=\"3\" valign=\"bottom\"\u003e\n \u003cp\u003e2325\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"15.942028985507246%\" colspan=\"2\" valign=\"bottom\"\u003e\n \u003cp\u003e0.2099\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"34.29951690821256%\" colspan=\"2\"\u003e\n \u003cp\u003e0.9893\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"36.47342995169082%\" valign=\"bottom\"\u003e\n \u003cp\u003e\u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp;Quercus*Rodent*Time\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"13.285024154589372%\" colspan=\"3\" valign=\"bottom\"\u003e\n \u003cp\u003e2325\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"15.942028985507246%\" colspan=\"2\" valign=\"bottom\"\u003e\n \u003cp\u003e0.0680\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"34.29951690821256%\" colspan=\"2\"\u003e\n \u003cp\u003e1.0000\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"38.795180722891565%\" colspan=\"2\" valign=\"bottom\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"9.156626506024097%\" valign=\"bottom\"\u003e\n \u003cp\u003e\u003cstrong\u003edf\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"16.3855421686747%\" colspan=\"2\" valign=\"bottom\"\u003e\n \u003cp\u003e\u003cstrong\u003e\u003cem\u003eF\u003c/em\u003e\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"13.012048192771084%\" colspan=\"2\"\u003e\n \u003cp\u003e\u003cstrong\u003e\u003cem\u003ep\u003c/em\u003e\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"22.650602409638555%\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"38.795180722891565%\" colspan=\"2\" valign=\"bottom\"\u003e\n \u003cp\u003e\u003cstrong\u003eDaily acorn mass\u0026nbsp;\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"9.156626506024097%\" valign=\"bottom\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"16.3855421686747%\" colspan=\"2\" valign=\"bottom\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"13.012048192771084%\" colspan=\"2\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"22.650602409638555%\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"38.795180722891565%\" colspan=\"2\" valign=\"bottom\"\u003e\n \u003cp\u003e\u0026nbsp; \u0026nbsp; \u0026nbsp; Intercept\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"9.156626506024097%\" valign=\"bottom\"\u003e\n \u003cp\u003e712\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"16.3855421686747%\" colspan=\"2\" valign=\"bottom\"\u003e\n \u003cp\u003e2469.6575\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"13.012048192771084%\" colspan=\"2\"\u003e\n \u003cp\u003e\u0026lt;0.0001\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"22.650602409638555%\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"38.795180722891565%\" colspan=\"2\" valign=\"bottom\"\u003e\n \u003cp\u003e\u003cem\u003eQuercus\u003c/em\u003e species\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"9.156626506024097%\" valign=\"bottom\"\u003e\n \u003cp\u003e712\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"16.3855421686747%\" colspan=\"2\" valign=\"bottom\"\u003e\n \u003cp\u003e549.6110\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"13.012048192771084%\" colspan=\"2\"\u003e\n \u003cp\u003e\u0026lt;0.0001\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"22.650602409638555%\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"38.795180722891565%\" colspan=\"2\" valign=\"bottom\"\u003e\n \u003cp\u003e\u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp;Rodent species\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"9.156626506024097%\" valign=\"bottom\"\u003e\n \u003cp\u003e712\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"16.3855421686747%\" colspan=\"2\" valign=\"bottom\"\u003e\n \u003cp\u003e360.8723\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"13.012048192771084%\" colspan=\"2\"\u003e\n \u003cp\u003e\u0026lt;0.0001\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"22.650602409638555%\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"38.795180722891565%\" colspan=\"2\" valign=\"bottom\"\u003e\n \u003cp\u003e\u0026nbsp; \u0026nbsp; Time\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"9.156626506024097%\" valign=\"bottom\"\u003e\n \u003cp\u003e712\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"16.3855421686747%\" colspan=\"2\" valign=\"bottom\"\u003e\n \u003cp\u003e0.3293\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"13.012048192771084%\" colspan=\"2\"\u003e\n \u003cp\u003e0.8584\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"22.650602409638555%\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"38.795180722891565%\" colspan=\"2\" valign=\"bottom\"\u003e\n \u003cp\u003e\u0026nbsp; \u0026nbsp; \u0026nbsp; Quercus*Rodent\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"9.156626506024097%\" valign=\"bottom\"\u003e\n \u003cp\u003e712\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"16.3855421686747%\" colspan=\"2\" valign=\"bottom\"\u003e\n \u003cp\u003e62.5560\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"13.012048192771084%\" colspan=\"2\"\u003e\n \u003cp\u003e\u0026lt;0.0001\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"22.650602409638555%\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"38.795180722891565%\" colspan=\"2\" valign=\"bottom\"\u003e\n \u003cp\u003e\u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp;Quercus*Time\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"9.156626506024097%\" valign=\"bottom\"\u003e\n \u003cp\u003e712\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"16.3855421686747%\" colspan=\"2\" valign=\"bottom\"\u003e\n \u003cp\u003e0.6025\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"13.012048192771084%\" colspan=\"2\"\u003e\n \u003cp\u003e0.9126\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"22.650602409638555%\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"38.795180722891565%\" colspan=\"2\" valign=\"bottom\"\u003e\n \u003cp\u003e\u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp;Rodent*Time\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"9.156626506024097%\" valign=\"bottom\"\u003e\n \u003cp\u003e712\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"16.3855421686747%\" colspan=\"2\" valign=\"bottom\"\u003e\n \u003cp\u003e1.1698\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"13.012048192771084%\" colspan=\"2\"\u003e\n \u003cp\u003e0.3147\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"22.650602409638555%\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"38.795180722891565%\" colspan=\"2\" valign=\"bottom\"\u003e\n \u003cp\u003e\u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp;Quercus*Rodent*Time\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"9.156626506024097%\" valign=\"bottom\"\u003e\n \u003cp\u003e712\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"16.3855421686747%\" colspan=\"2\" valign=\"bottom\"\u003e\n \u003cp\u003e0.7617\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"13.012048192771084%\" colspan=\"2\"\u003e\n \u003cp\u003e0.8572\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"22.650602409638555%\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003c/tbody\u003e\n\u003c/table\u003e\n\u003cp\u003e\u003cbr\u003e\u003c/p\u003e\n\u003cp\u003e\u003cbr\u003e\u003c/p\u003e"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":false,"hideJournal":false,"highlight":"","institution":"","isAcceptedByJournal":false,"isAuthorSuppliedPdf":false,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":false,"isPdf":false,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","identity":"behavioral-ecology-and-sociobiology","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"beas","sideBox":"Learn more about [Behavioral Ecology and Sociobiology](http://link.springer.com/journal/265)","snPcode":"265","submissionUrl":"https://www.editorialmanager.com/beas/default.aspx","title":"Behavioral Ecology and Sociobiology","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false},"keywords":"Acorn preferences, Rodents, Preserved embryo, Mutualistic, Antagonism, acorn composition","lastPublishedDoi":"10.21203/rs.3.rs-3410260/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-3410260/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003eThe relationship between scatter-hoarding rodents and oak species has been considered on a scale from antagonism to mutualism. Depending on the costs and benefits, the outcome of the relationship can be found at one extreme or the other. Costs have included destruction of acorns that occurs during predation, but not all acorns attacked lose their embryos.\u003c/p\u003e\n\u003cp\u003eAs representatives of the mutualistic end, we present two species that preserve embryo (\u003cem\u003eMus spretus\u003c/em\u003e and \u003cem\u003eApodemus sylvaticus\u003c/em\u003e). Representing the antagonistic extreme, we present a predatory species that destroys the embryo (\u003cem\u003eMicrotus arvalis\u003c/em\u003e).\u003c/p\u003e\n\u003cp\u003eThe objective of this study is to test the preferences of both rodent groups for acorns.\u003c/p\u003e\n\u003cp\u003eThe results showed that there is one acorn species (\u003cem\u003eQuercus ilex\u003c/em\u003e) that is preferred by the three rodent species. This acorn species has high concentrations of nutrients, low concentrations of tannins and thin shell. These characteristics attract the attention of rodents that could carry these acorns. There are two other acorn species (\u003cem\u003eQuercus suber\u003c/em\u003eand \u003cem\u003eQuercus rubra\u003c/em\u003e) that are consumed with little intensity for having low concentrations of nutrients, high concentration of tannins and thick shell. These characteristics escaping predators although transportation is not guaranteed. These acorns would germinate and emerge under the trees that have produced them and increase intraspecific competition.\u003c/p\u003e\n\u003cp\u003eCoincidence preferences shown by the three rodent species poses a risk for the oak species, since the recent arrival of the predator (antagonist) species in the study area could paralyze the dispersal process carried out by the other two mutualistic species through predation.\u003c/p\u003e","manuscriptTitle":"Preferences of rodent scatter-hoard and predators determine dissemination oak species.","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2023-10-13 20:36:17","doi":"10.21203/rs.3.rs-3410260/v1","editorialEvents":[{"type":"communityComments","content":0},{"type":"reviewerAgreed","content":"","date":"2023-10-14T05:36:38+00:00","index":0,"fulltext":""},{"type":"reviewersInvited","content":"","date":"2023-10-10T11:05:12+00:00","index":"","fulltext":""},{"type":"editorAssigned","content":"","date":"2023-10-08T20:53:34+00:00","index":"","fulltext":""},{"type":"submitted","content":"Behavioral Ecology and Sociobiology","date":"2023-10-04T07:12:48+00:00","index":"","fulltext":""}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"behavioral-ecology-and-sociobiology","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"beas","sideBox":"Learn more about [Behavioral Ecology and Sociobiology](http://link.springer.com/journal/265)","snPcode":"265","submissionUrl":"https://www.editorialmanager.com/beas/default.aspx","title":"Behavioral Ecology and Sociobiology","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false}}],"origin":"","ownerIdentity":"6d0beb05-9a26-458d-97ce-5defa3f7e816","owner":[],"postedDate":"October 13th, 2023","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"under-review","subjectAreas":[],"tags":[],"updatedAt":"2023-10-13T20:36:17+00:00","versionOfRecord":[],"versionCreatedAt":"2023-10-13 20:36:17","video":"","vorDoi":"","vorDoiUrl":"","workflowStages":[]},"version":"v1","identity":"rs-3410260","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-3410260","identity":"rs-3410260","version":["v1"]},"buildId":"-HB7Z8yhvgn0wM9Nzuekk","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}