Scatter-hoarding rodents prefer slightly astringent food

The mutualistic interaction between scatter-hoarding rodents and their seed plants is highly complex yet poorly understood. Plants may benefit from the seed dispersal behavior of rodents, as long as seed consumption is minimized. In parallel, rodents may maximize foraging efficiency and cache high-quality resources for future consumption. Defensive compounds, such as tannins, are thought to be a major mechanism for plant control over rodent behavior. However, previous studies, using naturally occurring seeds, have not provided conclusive evidence supporting this hypothesis. Here, we test the importance of tannin concentrations on the scatter-hoarding behavior of rodents by using an artificial seed system. We combined feeding trials and field observations to examine the overall impact of seed tannin concentrations on rodent behavior and health. We found that rodents favored seeds with an intermediate amount of tannin (~5%) in the field. Meanwhile, in rodents that were fed a diet with different tannin content, only diets with high tannin content (25%, 15%, and 10%) caused a significant negative influence on rodent survival and health. Significant differences were not found among treatments with tannin levels of 0-5%. In contrast to many existing studies, our results clearly demonstrate that scatter-hoarding rodents prefer slightly 'astringent' food. In the co-evolutionary arms race between plants and animals, our results suggest that while tannins may play a significant role in reducing general predation levels by the faunal community, they have no precise control over the behavior of their mutualistic partner. Instead, the two partners appear to have reached an evolutionary point where both parties receive adequate benefits, with the year-to-year outcome being dependent on a wide range of factors beyond the control of either partner.     

Tannin concentration enhances seed caching by scatter-hoarding rodents: An experiment using artificial ‘seeds’

Tannins are very common among plant seeds but their effects on the fate of seeds, for example, via mediation of the feeding preferences of scatter-hoarding rodents, are poorly understood. In this study, we created a series of artificial ‘seeds’ that only differed in tannin concentration and the type of tannin, and placed them in a pine forest in the Shangri-La Alpine Botanical Garden, Yunnan Province of China. Two rodent species (Apodemus latronum and A. chevrieri) showed significant preferences for ‘seeds’ with different tannin concentrations. A significantly higher proportion of seeds with low tannin concentration were consumed in situ compared with seeds with a higher tannin concentration. Meanwhile, the tannin concentration was significantly positively correlated with the proportion of seeds cached. The different types of tannin (hydrolysable tannin vs condensed tannin) did not differ significantly in their effect on the proportion of seeds eaten in situ vs seeds cached. Tannin concentrations had no significant effect on the distance that cached seeds were carried, which suggests that rodents may respond to different seed traits in deciding whether or not to cache seeds and how far they will transport seeds.     

Teasing Apart the Effects of Seed Size and Energy Content on Rodent Scatter-Hoarding Behavior

Scatter-hoarding rodents are known to play a crucial role in the seed dispersal of many plant species. Numerous studies have indicated that both seed size and the energy content of seeds can affect rodent foraging behavior. However, seed size is usually associated with energy content per seed, making it difficult to isolate how seed size and energy affect rodent foraging preferences. This study used 99 treatments of artificial seeds (11 seed sizes×9 levels of energy content) to tease apart the effect of seed size and energy content on rodent seed-caching behavior. Both seed traits showed significant effects, but their details depended on the stage of the rodent foraging process. Seeds with higher energy content were harvested more rapidly while seed size only had a modest effect on harvest rate. However, after harvesting, seed size showed a much stronger effect on rodent foraging behavior. Rodents’ choice of which seeds to remove and cache, as well as seed dispersal distance, seemed to reflect an optimal seed size. Our findings could be adapted in future studies to gain a better understanding of scatter-hoarding rodent foraging behavior, and the co-evolutionary dynamics between plant seed production and seed dispersers.     

围栏条件下同域分布三种鼠对两种种子的贮藏行为差异

种子贮藏是啮齿动物利用食物资源的一种适应性行为。同域分布的啮齿动物可能进化出不同的贮藏方式和策略。为了解同域分布的啮齿动物的贮藏策略及种子特征在动物贮藏行为决策中的作用,于2014年10-12月,通过半自然围栏实验,在秦岭南坡的佛坪国家级自然保护区内调查了同域分布社鼠、中华姬鼠和甘肃仓鼠对板栗和锐齿槲栎种子的贮藏策略差异,并探讨了2种种子特征在鼠类贮藏行为决策中的作用。结果显示：1）社鼠主要集中贮藏板栗种子,未分散贮藏任何种子;中华姬鼠未贮藏板栗种子,对锐齿槲栎种子同时表现出集中和分散贮藏行为,以集中贮藏为主;甘肃仓鼠对2种种子均表现出集中贮藏行为,未表现分散贮藏行为。2）社鼠和甘肃仓鼠均倾向于集中贮藏有较高营养价值（高蛋白和脂肪）且单宁含量较低的板栗种子;中华姬鼠倾向于分散贮藏单宁含量高的锐齿槲栎种子。3）3种鼠均喜好取食有较高营养价值且单宁含量低的板栗种子,很少取食单宁含量高的锐齿槲栎种子。结果表明同域分布鼠类对不同种子的贮藏方式有所不同,种子特征影响鼠类的取食和贮藏策略。     

Post‐dispersal seed predation and its relations with seed traits: a thirty‐species‐comparative study

Post‐dispersal seed predation is a key process determining the variability in seed survival in forests, where most seeds are handled by rodents. Seed predation is thought to affect seedling regeneration, colonization ability and spatial distribution of plants. Basic seed traits are the essential factors affecting rodent foraging preferences and thus seed survival and seedling recruitment. Many studies have discussed several seed traits and their effects upon seed predation by rodents. However, the results of those previous studies are usually equivocal, likely because few seed traits and/or plant species tend to be incorporated into these studies. In order to elucidate the relationships between seed predation and seed traits, we surveyed the predation of 48 600 seeds in a natural pine forest, belonging to 30 species, for three consecutive years. The results demonstrated that: (i) seed size and seed coat hardness did not significantly affect seed predation; (ii) total phenolics had a negative effect upon seed predation; (iii) positive effects of nitrogen content upon seed predation were found. From our study, it seems that the better strategy to prevent heavy predation is for plants to produce seeds with higher total phenolics content rather than physical defenses (i.e. hard seed coat) or larger seeds. Additionally, rodent foraging preference may depend more on Nitrogen content than other nutrient content of seeds.     

Dissecting the decision making process of scatter‐hoarding rodents

Many animals scatter‐hoard seeds to ensure an even supply of food throughout the year and this behavior requires similar foraging decisions. Seed‐traits have been shown to affect the final foraging decision but little is known about the decision process itself. Here, we first defined four sequential steps comprising the decision process of scatter‐hoarding rodents: 1) upon encountering a seed, should it be ignored or manipulated; 2) if manipulated, should it be eaten in situ or removed elsewhere; 3) upon removal, how far away should it be carried; and finally 4) whether to eat or cache the removed seed. Using experimental seeds with controlled differences in size, tannin and nutrient content, we evaluated how different traits influence each step in this decision process. We found that different traits had distinct effects on each step. Seed size affected all four steps, while nutrient and tannin content primarily affected the first and third steps. By dissecting foraging behavior in relation to experimentally controlled seed‐traits, we have created an effective framework within which to understand the unique relationship between scatter‐hoarding rodents that both predate and disperse plant seeds.     

Fragmentation modifies seed trait effects on scatter-hoarders’ foraging decisions

Scatter-hoarding animals are crucial in seed dispersal of nut-bearing plants. We used the holm oak Quercus ilex—wood mouse Apodemus sylvaticus mutualism as a model system to evaluate the relative importance of seed size and fat content on scatter-hoarders’ foraging decisions influencing oak dispersal and potential recruitment. We performed a field experiment in which we offered holm oak acorns with contrasting seed size (2 vs 5 g) and fat content (3 vs 11%). Moreover, to test if the strength of these seed trait effects was context-dependent, experimental acorns were placed in small fragments, where natural regeneration is scarce or absent, and forest habitats. In small fragments, rodents had to face increased intraspecific competition for acorns and reduced anti-predator cover during transportation. As a result, they became more selective to ensure rapid acquisition of most valuable food items but, in turn, transported seeds closer to avoid unaffordable predation risks. During harvesting and caching, larger acorns were prioritized and preferentially cached. Fat content only had a minor effect in harvesting preferences. In contrast, in forest sites, where rodent abundance was four times lower and understory cover was well-developed, rodents were not selective but provided enhanced dispersal services to oaks (caching rates were 75% higher). From the plants’ perspective, our results imply that the benefits of producing costly seeds are context-dependent. Seed traits modified harvesting and caching rates only when rodents were forced to forage more efficiently in response to increased intraspecific competition. However, when landscape traits limited cache protection strategies, a more selective foraging behavior by scatter-hoarders did not result in enhanced dispersal services. Overall, our result shows that successful dispersal of acorns depends on how specific traits modulate their value and how landscape properties affect rodents’ ability to safeguard them for later consumption.     

Scatter-hoarding rodents use different foraging strategies for seeds from different plant species

Seed predation and dispersal by scatter-hoarding rodents are key processes that determine seed survival, and thus, plant regeneration within forests. For decades, there has been much debate on the important effects of seed size (one of the most important seed traits) on rodent foraging preference. Furthermore, the possible selective forces in the evolution of seed size may be influenced by primary selectivity and how rodents treat seeds after harvesting. In this study, different-sized seeds from four species (Pinus armandii, Pinus densata, Abies sp., and Viburnum sp.) harvested by scatter-hoarding rodents were studied in an alpine forest in Southwestern China for two consecutive years. Our results showed that seed size influenced rodent foraging preferences, with bigger seeds being preferred over smaller seeds, within and across species. Rodents only removed and cached the larger seeds of P. armandii, and ate the seeds of the other three species in situ. Rodents are purely seed predators for these three species. For the cached seeds of P. armandii, significantly positive correlations were observed between seed size and dispersal distance among both primary and secondary cached seeds in 2006, but not in 2005. Our results indicate that among many coexisting species with widely different-sized seeds, scatter-hoarding rodents played important roles in the seed dispersal of the big-seeded species alone. This caching behavior could offset the limited seed dispersal of large-seeded and wingless species (P. armandii), in comparison with that of small winged seed species (P. densata and Abies sp.) and frugivore-dispersed species (Viburnum sp.).     

The foraging behavior of granivorous rodents and short-term apparent competition among seeds

The foraging behavior of a predator species is thought to bethe cause of short-term apparent competition among those preyspecies that share the predator. Short-term apparent competitionis the negative indirect effect that one prey species has onanother prey species via its effects on predator foraging behavior.In theory, the density-dependent foraging behavior of granivorousrodents and their preference for certain seeds are capable of inducing short-term apparent competition among seed species.In this study, I examined the foraging behavior of two heteromyidrodent species (family Heteromyidae), Merriam's kangaroo rats(Dipodomys merriami) and little pocket mice (Perognathus longimembris).In one experiment I tested the preferences of both rodent speciesfor the seeds of eight plant species. Both rodent species exhibiteddistinct but variable preferences for some seeds and avoidanceof others. However, the differences in preference appearedto have only an occasional effect on the strength of the short-term apparent competition detected in a field experiment. In anotherexperiment, I found that captive individuals of both rodentspecies had approximately equal foraging effort (i.e., timespent foraging) in patches that contained a highly preferredseed type (Oryzopsis hymenoides) regardless of seed density and the presence of a less preferred seed type (Astragalus cicer)in the patches. The rodents also harvested a large proportionof O. hymenoides seeds regardless of initial seed density;this precluded a negative indirect effect of A. cicer on O.hymenoides. But there was a negative indirect effect of O.hymenoides on A. cicer caused by rodents having a lower foragingeffort in patches that only contained A. cicer seeds than inpatches that contained A. cicer and O. hymenoides seeds. Theindirect interaction between O. hymenoides and A. cicer thusrepresented a case of short-term apparent competition thatwas non-reciprocal. Most importantly, it was caused by theforaging behavior of the rodents.     

Endocarp thickness affects seed removal speed by small rodents in a warm-temperate broad-leafed deciduous forest, China

Seed traits are important factors affecting seed predation by rodents and thereby the success of recruitment. Seeds of many tree species have hard hulls. These are thought to confer mechanical protection, but the effect of endocarp thickness on seed predation by rodents has not been well investigated. Wild apricot (Prunus armeniaca), wild peach (Amygdalus davidiana), cultivated walnut (Juglans regia), wild walnut (Juglans mandshurica Maxim) and Liaodong oak (Quercus liaotungensis) are very common tree species in northwestern Beijing city, China. Their seeds vary greatly in size, endocarp thickness, caloric value and tannin content. This paper aims to study the effects of seed traits on seed removal speed of these five tree species by small rodents in a temperate deciduous forest, with emphasis on the effect of endocarp thickness. The results indicated that speed of removal of seeds released at stations in the field decreased significantly with increasing endocarp thickness. We found no significant correlations between seed removal speed and other seed traits such as seed size, caloric value and tannin content. In seed selection experiments in small cages, Père David's rock squirrel (Sciurotamias davidianus), a large-bodied, strong-jawed rodent, selected all of the five seed species, and the selection order among the five seed species was determined by endocarp thickness and the ratio of endocarp mass/seed mass. In contrast, the Korean field mouse (Apodemus peninsulae) and Chinese white-bellied rat (Niviventer confucianus), with relatively small bodies and weak jaws, preferred to select small seeds like acorns of Q. liaotungensis and seeds of P. armeniaca, indicating that rodent body size is also an important factor affecting food selection based on seed size. These results suggest endocarp thickness significantly reduces seed removal speed by rodents and then negatively affects dispersal fitness of seeds before seed removal of tree species in the study region. However, effect of endocarp thickness on final dispersal fitness needs further investigation because it may increase seed caching and survival after seed removal.     

种子特征和结实量对啮齿动物取食和扩散种子行为的影响

为了深入了解啮齿动物在不同种子丰富度条件下对不同大小和单宁含量种子的觅食行为策略及其与植物种群更新的关系,在宁夏六盘山区的华北落叶松人工林,研究了不同大小和单宁含量[0%Tannin(T)、2%T、8%T和15%T]的人工种子在模拟结实小年和结实大年对啮齿动物取食和扩散行为的影响.结果表明: 啮齿动物消耗种子速度在结实小年更快,结实大年的种子消耗速度相对缓慢. 种子就地取食率(ISPR)在不同结实年份间无显著差异,扩散后取食率(PRAD)在结实小年显著高于结实大年,但前者的扩散后贮藏率(HRAD)显著低于后者;种子扩散后的取食距离(PDAD)和贮藏距离(HDAD)在结实小年均显著大于结实大年.在结实小年,大种子的PDAD和HDAD均大于小种子,前者在不同大小种子间均差异显著,而后者仅在2%T和15%T的不同大小种子间差异显著;在结实大年,除0%T外的其他单宁含量种子的PDAD和HDAD在不同大小种子间均差异显著.ISPR在中等单宁含量种子最大,高单宁含量种子最小;PRAD分别在结实小年的高单宁含量种子和结实大年的无单宁种子最大;不论在结实大年还是结实小年,HRAD均在高单宁含量种子最大,中等单宁含量种子最小.这说明结实大年可延缓啮齿动物对种子的消耗速率,提高种子的HRAD,但种子扩散距离减小;啮齿动物在结实大年和小年均表现出对大种子的扩散偏好,且大种子被扩散的距离更远;啮齿动物在不同结实年份均偏好于就地取食中等单宁含量种子,而扩散高单宁含量种子.     

Segregating the Effects of Seed Traits and Common Ancestry of Hardwood Trees on Eastern Gray Squirrel Foraging Decisions

The evolution of specific seed traits in scatter-hoarded tree species often has been attributed to granivore foraging behavior. However, the degree to which foraging investments and seed traits correlate with phylogenetic relationships among trees remains unexplored. We presented seeds of 23 different hardwood tree species (families Betulaceae, Fagaceae, Juglandaceae) to eastern gray squirrels (Sciurus carolinensis), and measured the time and distance travelled by squirrels that consumed or cached each seed. We estimated 11 physical and chemical seed traits for each species, and the phylogenetic relationships between the 23 hardwood trees. Variance partitioning revealed that considerable variation in foraging investment was attributable to seed traits alone (27–73%), and combined effects of seed traits and phylogeny of hardwood trees (5–55%). A phylogenetic PCA (pPCA) on seed traits and tree phylogeny resulted in 2 “global” axes of traits that were phylogenetically autocorrelated at the family and genus level and a third “local” axis in which traits were not phylogenetically autocorrelated. Collectively, these axes explained 30–76% of the variation in squirrel foraging investments. The first global pPCA axis, which produced large scores for seed species with thin shells, low lipid and high carbohydrate content, was negatively related to time to consume and cache seeds and travel distance to cache. The second global pPCA axis, which produced large scores for seeds with high protein, low tannin and low dormancy levels, was an important predictor of consumption time only. The local pPCA axis primarily reflected kernel mass. Although it explained only 12% of the variation in trait space and was not autocorrelated among phylogenetic clades, the local axis was related to all four squirrel foraging investments. Squirrel foraging behaviors are influenced by a combination of phylogenetically conserved and more evolutionarily labile seed traits that is consistent with a weak or more diffuse coevolutionary relationship between rodents and hardwood trees rather than a direct coevolutionary relationship.     

Synchrony between fruit maturation and effective dispersers' foraging activity increases seed protection against seed predators

The evolution of pollination and seed dispersal mutualisms is conditioned by the spatial and temporal co-occurrence of animals and plants. In the present study we explore the timing of seed release of a myrmecochorous plant (Helleborus foetidus) and ant activity in two populations in southern Spain during 2 consecutive years. The results indicate that fruit dehiscence and seed shedding occur mostly in the morning and correspond to the period of maximum foraging activity of the most effective ant dispersers. By contrast, ant species that do not transport seeds and/or that do not abound near the plants are active either before or after H. foetidus diaspores are released. Experimental analysis of diet preference for three kinds of food shows that effective ant dispersers are mostly scavengers that readily feed on insect corpses and sugars. Artificial seed depots suggest that seeds deposited on the ground out of the natural daily time window of diaspore releasing are not removed by ants and suffer strong predation by nocturnal rodents Apodemus sylvaticus. Nevertheless, important inter-annual variations in rodent populations cast doubts on their real importance as selection agents. We argue that traits allowing synchrony between seed presentation and effective partners may constitute a crucial pre-adaptation for the evolution of plant-animal mutualisms involving numerous animal partners.     

Functional traits determine formation of mutualism and predation interactions in seed-rodent dispersal system of a subtropical forest

Network structure in plant-animal systems has been widely investigated but the roles of functional traits of plants and animals in formation of mutualism and predation interactions and community structure are still not fully understood. In this study, we quantitatively assessed interaction strength of mutualism and predation between 5 tree species and 7 rodent species by using semi-natural enclosures in a subtropical forest in southwest China. Seeds with high handling-time and nutrition traits (for both rat and mouse species) or high tannin trait (for mouse species) show high mutualism but low predation with rodents; while seeds with low handling-time and low nutrition traits show high predation but low mutualism with rodents. Large-sized rat species are more linked to seeds with high handling-time and high nutrition traits, while small-sized mouse species are more connected with seeds with low handling-time, low nutrition value and high tannin traits. Anti-predation seed traits tend to increase chance of mutualism instead of reducing predation by rodents, suggesting formation of mutualism may be connected with that of predation. Our study demonstrates that seed and animal traits play significant roles in the formation of mutualism and predation and network structure of the seed-rodent dispersal system.     

秦岭南北坡森林鼠类对板栗和锐齿栎种子扩散的影响

森林鼠类的种子贮藏行为对植物的扩散及更新会产生积极的影响。2012和2013年秋季,分别在秦岭北坡的周至国家级自然保护区和南坡的佛坪国家级自然保护区内,调查了森林鼠类对板栗(Castanea mollissima)和锐齿栎(Quercus aliena)种子的取食和扩散差异。结果显示:1)秦岭南北坡的环境因素,特别是植被因素,对鼠类扩散板栗和锐齿栎种子具有重要的影响。南坡较为丰富的壳斗科植被种类,导致2种种子在南坡存留时间均长于北坡,而北坡的扩散取食和丢失率均高于南坡。2)种子特征影响鼠类的取食或贮藏偏好。由于较高的蛋白、脂肪等营养含量,鼠类更喜好取食或搬运贮藏板栗种子。然而,低营养但高丹宁含量的锐齿栎种子仍然被鼠类大量贮藏。3)2种种子在南北坡的扩散历程在两个年份间有很大差异,在食物相对匮乏的年份(2012年),种子被扩散的速度更快且丢失的比率更高。这种差异反映了种子大小年现象对森林鼠类取食和贮藏策略的影响。4)无论在秦岭南坡还是北坡,营养价值含量(如蛋白和脂肪)较高的板栗种子的取食和贮藏距离都明显大于营养价值含量较低的锐齿栎种子,这与最优贮藏空间分布模型的预测一致。     

Pre- and postdispersal seed predation by rodents: balance of food and safety

Seed presentation and availability for seed predators changeduring every plant reproductive cycle. We know very little abouthow those changes impinge on both the ability of seed predatorsto impact plant populations and the foraging costs associatedwith seed consumption. Therefore, we conducted several fieldexperiments to evaluate whether wood mice Apodemus sylvaticusbalance food and safety while foraging on Helleborus foetidusseeds during both the pre- and early postdispersal phases ofthe plant reproductive cycle. Both food and safety were keydeterminants of mouse foraging on H. foetidus seeds, thoughtheir roles were not consistent along the plant reproductivecycle. Thus, augmenting ambient food reduced fruit removal bymice during the predispersal phase. During the postdispersalphase, seeds in sheltered microsites experienced higher removalrates than those located in nonsheltered microsites; however,no effect of food augmentation was detected. This apparent reversedrole of food and safety on decision making by mice seemed closelylinked to both the dramatic changes in accessibility and presentationof H. foetidus seeds and the coupled changing foraging costsfaced by mice at different phases of the plant reproductivecycle. For instance, because the cost of foraging for predispersalseeds was higher than for postdispersal seeds, the effect offood augmentation on foraging by wood mice was greater duringthe predispersal phase. Thus, our study illustrates the needof considering differences between pre- and postdispersal seedpredation in the study of granivore rodents and their impacton plant populations.     

Consistencies in post-dispersal seed predation of temperate fleshy-fruited species among seasons, years and sites

1. Seed predation of 12 fleshy-fruited species was recorded in experimental dishes under early successional forest in south-west Germany (four seasons 1992 and 1993) and in south England (summer 1995). On each occasion, 200 dishes were laid out, each containing five seeds of a given species. The mean time for three or more seeds to be removed was taken as a measure of granivore preferences. We tested correlations of these preferences with several physical and nutritional seed traits.
2. Live trapping and selectively accessible dishes indicated that rodents were the major granivores ( Apodemus sylvaticus, Apodemus flavicollis, Clethrionomys glareolus ); no predation by birds or insects was observed. The rank order of the rodents' seed preferences was consistent among sites, seasons and years, but mean predation differed between species, sites and seasons. Seed predation was highest in summer and lowest in winter. Possible implications of the seasonal pattern in predation risk are discussed.
3. The preferences of rodents were significantly correlated with the species-specific viability of seeds (i.e. percentage of sound seed) in five of eight experiments and with the percentage of water in the embryo-plus-endosperm fraction (EEF) in four of eight experiments. Surprisingly, preferences were not correlated to seed mass, EEF mass or nitrogen concentration. Predation was lowest among toxic species ( Berberis vulgaris, Euonymus europaeus, Sambucus nigra ) and among species with woody endocarps ( Cornus sanguinea, Crataegus spp.).     

Functional extinction of a desert rodent: implications for seed fate and vegetation dynamics

Population declines of once‐abundant species have often preceded understanding of their roles within ecosystems. Consequently, important drivers of environmental change may remain undiagnosed because we simply do not know how species that are now rare or extinct shaped ecosystems in the past. Australia's desert rodents are thought to have little numerical impact on seed fate and vegetation recruitment when compared with ants or with desert rodents on other continents. However most research on granivory by Australian desert rodents has occurred in areas where rodents were rare or functionally extinct. Here we ask if the paradigm that rodents are relatively un‐important granivores in Australian deserts is an artefact of their historical decline. In the Strzelecki Desert, the endangered rodent, Notomys fuscus is rare where introduced mesopredators are abundant but common where dingoes (an apex predator) suppress mesopredator populations. We used foraging trays to compare rates of seed removal for a common shrub (Dodonaea viscosa angustissima hopbush) between areas where N. fuscus, hopbush shrubs and their seedlings were rare and common and found that seed removal was consistently higher where rodents were common and hopbush rare. By excluding ants and rodents from foraging trays we show that ants removed more seeds than rodents where rodents were rare but rodents removed far more seeds than ants where rodents were common. By manipulating rodents’ access to the soil seed‐bank we show that hopbush seeds persisted in greater numbers where rodents were excluded than where they had access. Our results support the hypothesis that granivory by rodents may once have been a far more important process influencing the fate of seeds and shaping plant communities in arid Australia and suggest that dingo extirpation has cascading effects on shrub seeds. Our study highlights that functional extinction of rodents may be an under‐appreciated driver of vegetation change.     

The challenge of interpreting primate diets: mangabey foraging on Blighia unijugata fruit in relation to changing nutrient content

Primates often make foraging selections that are not apparent. For example, they may eagerly consume a particular plant part and species in some instances, but reject it at other times. Blighia unijugata (Baker) fruit is one of the most frequently eaten foods of mangabeys (Lophocebus albigena) in Lwamunda Forest Reserve, Uganda; however, its use varies strikingly depending on the fruit's developmental stage. We conducted feeding observations to investigate the nutritional criteria that mangabeys may have used for the consumption of specific fruit parts by conducting analysis of fruit parts eaten and rejected at different developmental stages. When seeds had low condensed tannins, mangabeys ate them, but seeds were rejected when tannin levels increased. In the first five stages of fruit development, tannin levels in arils declined and the frequency of consumption of the aril increased. Although fat content in seeds increased with maturity, it did not appear to influence seed consumption, but fat content was related to frequency of consumption of the aril. Considering that primates are often making food selections among many species/part combinations, our results illustrate the value of using nutritional analyses to understand foraging decisions. Furthermore, they demonstrate how very specific trade‐offs between consumption of critical nutrients and antifeedants can drive which foods and parts are eaten.     

Tree-to-tree variation in seed size and its consequences for seed dispersal versus predation by rodents

Individual variation in seed size and seed production is high in many plant species. How does this variation affect seed-dispersing animals and, in turn, the fitness of individual plants? In this study, we first surveyed intraspecific variation in seed mass and production in a population of a Chinese white pine, Pinus armandii. For 134 target trees investigated in 2012, there was very high variation in seed size, with mean seed mass varying among trees almost tenfold, from 0.038 to 0.361 g. Furthermore, 30 of the 134 trees produced seeds 2 years later, and for these individuals there was a correlation in seed mass of 0.59 between years, implying consistent differences among individuals. For a subset of 67 trees, we monitored the foraging preferences of scatter-hoarding rodents on a total of 15,301 seeds: 8380 were ignored, 3184 were eaten in situ, 2651 were eaten after being cached, and 395 were successfully dispersed (cached and left intact). At the scale of individual seeds, seed mass affected almost every decision that rodents made to eat, remove, and cache individual seeds. At the level of individual trees, larger seeds had increased probabilities of both predation and successful dispersal: the effects of mean seed size on costs (predation) and benefits (caching) balanced out. Thus, despite seed size affecting rodent decisions, variation among trees in dispersal success associated with mean seed size was small once seeds were harvested. This might explain, at least in part, the maintenance of high variation in mean seed mass among tree individuals.