The Proximate Basis of the Oak Dispersal Syndrome: Detection of Seed Dormancy by Rodents

Previously we have shown how a range of physical and chemicalcharacteristics of acorns influences the behavioral decisionsof food-hoarding rodents which in turn affects the dispersal,establishment and spatial arrangement of oaks. One such behaviorinvolves the selective caching of acorns of red oaks (subgenus:Erythrobalanus) over those of white oaks (Quercus) because ofreduced perishability that results from delayed germinationof acorns in the red oak group. In this study, we sought toidentify the specific proximate cues (visual and olfactory)that eastern gray squirrels (Sciurus carolinensis) use whenmaking these decisions. In two series of field experiments,we presented individual, free-ranging animals with pairs ofexperimentally altered acorns (that differed with respect toa single chemical or visual characteristic) and recorded theirfeeding and caching responses. Squirrels cached artificial acornswith pericarps (shells) of red oak acorns and ate those withshells of white oak regardless of the internal chemical compositionof either type of acorn. Only when the shells of artificialacorns were first soaked in acetone (to remove potential chemicalodors) did animals eat artificial acorns made with the shellsof red oak acorns. Squirrels also ate one-year old red oak acornsthat had broken dormancy, even when they exhibited no signsof germination. We argue that a chemical cue in the shell ofacorns is important in the detection of seed dormancy and thedecision to cache acorns, and that such a cue might ultimatelycontribute to the differential dispersal of red and white oaksby rodents.     

Dietary circumvention of acorn tannins by blue jays

Blue jays consume large quantities of acorns to fuel energy-demanding caching flights in the fall. Yet blue jays possess no known physiological adaptation to counter the negative effects of a high tannin diet on protein digestion. Dietary experiments were conducted to determine if blue jays could subsist on an acorn-only diet, and if they could not, to determine whether supplements of acorn weevil larvae (Curculio), present inside acorns, enabled them to maintain their mass. Comparative tannin assays also were conducted on Lepidobalanus (low tannin; white oak) and Erythrobalanus (high tannin; pin oak) acorns using radial diffusion assay. Captive jays consumed considerable acorn material, yet were unable to maintain mass on ad lib. acorn-only diets or on an acorn +1.5 g larvae/day supplement. There were no significant differences in mass loss between high and low tannin diets. In contrast, blue jays were able to stabilize mass on a diet of acorns +5.0 g larvae supplement/day. These results suggest that acorn weevil larvae, or perhaps other insects, counteract the effects of acorn tannins in the jay diet allowing jays to subsist largely on acorns during the fall caching season. Oak demographic processes may be partly regulated by a tri-trophic relationship among plant, insect and bird. Acorn weevil larvae, considered damaging to oak populations, may actually facilitate oak recruitment and population vagility in the long-term.     

Acorn Pericarp Removal as a Cache Management Strategy of the Siberian Chipmunk,Tamias sibiricus

Numerous recent studies have revealed a variety of behavioral adaptations of rodents for maximizing returns from cached seeds. Herein we report on a novel behavior by the Siberian chipmunk (Tamias sibiricus) in northeastern China, by which they consistently remove the pericarp (shell) of Quercus mongolica acorns before dispersing and caching these nuts. We investigated the effects of pericarp removal on acorn germination, tannin concentrations, cache pilferage, and insect damage, to determine if and how pericarp removal facilitates cache management by Siberian chipmunks and whether or not such behavior influences seed fates. Chipmunks cached acorns only after the pericarps were removed. Chipmunks preferred pericarp‐removed acorns over intact acorns when removing them from seed stations for both consumption and caching. Pericarp removal did not affect germination or tannin concentration of cached Q. mongolica acorns, suggesting that the behavior is not an adaptation for long‐term storage and tannin decomposition. Acetone treatments of the pericarp and artificial pericarp removal failed to alter pilferage rates by Siberian chipmunks and wood mice (Apodemus peninsulae). Since damage of acorns by weevils often leads to cache losses, we also tested the effects of weevil infestation on cache decision following pericarp removal. Siberian chipmunks removed pericarps and then scatter hoarded significantly more sound than weevil‐infested acorns, strongly suggesting that pericarp removal is used to discriminate between the infested and non‐infested acorns. Thus, we argue that the primary function of this behavior is to ensure successful storage of sound acorns, at least for short‐term storage. Future studies should consider the potential impact of pericarp removal on weevil populations and long‐term patterns of seed survival and establishment from the Siberian chipmunk’s caches.     

Spatial patterns in long-distance dispersal of Quercus ilex acorns by jays in a heterogeneous landscape

In this paper, I analyse the interaction between the holm-oak Quercus ilex , and one of its main dispersers, the European jay Garrulus glandarius , in an heterogeneous Mediterranean landscape. I quantify the spatial dispersal pattern of the seed shadow at two spatial scales, landscape (among patches) and microhabitat (within patches), by directly tracking the movement of seeds. Two main traits of the jay-mediated dispersal of holm-oak acorns across the landscape, the spatial pattern of dissemination and the distance from the source tree, are significantly and directly influenced by jay activity. Jays moved acorns nonrandomly, avoiding one main patch type of the study area to cache acorns, the shrubland-grasslands, and moving most of the acorns to pine stands, whether afforestation or open pinewoods. Within each patch type, jays had also a strong preference for caching acorns in some microhabitats, since>95% of the acorns dispersed by jays were cached beneath pines. The distance of holm-oak acorn dispersal was long in the study site, over 250 m, with some dispersals occurring up to 1 km from the source oaks. The shape of the dispersal kernel function fitted to the dispersal pattern produced by jays differed from those quantified for many other plant species. Jay-mediated dispersal had two components, one local and another produced by long-distance dispersal. Due to the heterogeneity of these Mediterranean environments, this difference in scale overlaps with a difference in habitat composition, short distances events resulting in dispersals within the same oak stands and long distance events resulting in dispersal outside of oak stands, usually to other vegetation units. Jay activity and movement pattern can have thus dramatic effects on both the local regeneration as well as the potential for regional spread of the holm-oak populations.     

Re-caching of acorns by rodents: Cache management in eastern deciduous forests of North America

Scatter-hoarding rodents such as tree squirrels selectively cache seeds for subsequent use in widely-spaced caches placed below the ground surface. This behavior has important implications for seed dispersal, seedling establishment, and tree regeneration. Hoarders manage these caches by recovering and eating some seeds, and moving and re-caching others. This process of re-caching, however, is poorly understood. Here, we use radio-telemetry to evaluate re-caching behavior for the management of acorn caches by rodents in eastern deciduous forests. We also test the hypothesis that as seeds are re-cached, the distance from the source increases. Radio transmitters were implanted in Northern red oak (Quercus rubra) acorns and presented to rodents in a natural setting over 3 seasons. We used radio-telemetry to track and document evidence of recovery and re-caching. We tracked a total of 102 acorns. Of the 39 radio-tagged acorns initially cached, 19 (49%) were cached on two or more occasions; one acorn was cached four times. The hypothesis that rodents move seeds to progressively greater distances from the source is not well-supported, suggesting that acorns are being moved within an individual's home range. Given the species of rodents in the study area, gray squirrels (Sciurus carolinensis) are the most likely to be responsible for the caching and re-caching events. Gray squirrels appear to engage in extensive re-caching during periods of long-term food storage, which has important implications for understanding how caching behavior influences acorn dispersal and oak regeneration.     

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.     

Partial acorn consumption by small rodents: implication for regeneration of white oak, <Emphasis Type="Italic">Quercus mongolica</Emphasis>

Partial consumption of acorns by rodents, birds, and insects has been widely reported in various oak species. However, to what extent these partially eaten acorns contribute to the regeneration of oak trees is poorly understood. To date, there is limited knowledge of the effects of seed availability on partial consumption of acorns. Herein, we released tagged Quercus mongolica acorns in two consecutive years with different seed crops, to explore the probability of partial acorn consumption. We also placed simulated partially consumed acorns in the field to investigate their contribution to regeneration of white oak. Our results showed that more acorns were partially eaten in a good crop year than in poor crop year, reflecting an effect of predator satiation on acorn partial consumption by rodents at the population level. Partially eaten acorns were more likely to be damaged at the basal end, suggesting consistent consumption preferences of small rodents. Although, partially consumed acorns were less likely to be scatter-hoarded by small rodents, they germinated more rapidly than the intact acorns in the field, offsetting the negative effects of the non-buried deposition. Despite lower germination rates, lightly damaged acorns exhibited greater growth of roots and shoots, suggesting a compensatory response to partial acorn consumption. Partial consumption may spread predation pressure on acorns and thus appears to be much better for the plant than total consumption by seed-eating animals. Therefore, partially consumed acorns as dispersal leftovers may play a potential role in natural regeneration of Quercus mongolica, especially in mast years. However, this role and the underlying mechanisms of partial acorn consumption by rodents, birds, and herbivore insects need further investigation.     

Acorn selection by Mexican jays: a test of a tri-trophic symbiotic relationship hypothesis

By caching acorns, jays serve as important dispersal agents for oak (Quercus) species. Yet little is known about which acorn characteristics affect selection by jays. In the traditional model of jay/oak symbiosis, large, brown, ripe acorns free of invertebrate parasites (e.g., Curculio acorn weevils) are selected by jays. Recently, it has been suggested that a tri-trophic relationship between oaks, jays, and weevils may have evolved to counter the negative dietary effects of acorn tannins. Under the tri-trophic model, jays would preferentially select acorns containing weevil larvae. We tested the assumptions that (1) acorns containing curculionid larvae exist in sufficient quantities to support jay populations and (2) jays can detect, and preferentially select, acorns containing weevil larvae, and investigated the cues by which jays select acorns. Captive Mexican jays (Aphelocomaultramarina) were presented Emory oak (Quercusemoryi) acorns in aviary feeding trials. Large, dense, viable acorns free of curculionid larvae were preferentially selected. Contrary to results of previous research, color did not affect selection. Acorn viability increased and curculionid larval occupancy decreased in adjacent savannas and isolated stands relative to existing oak woodland, perhaps favoring oak recruitment into adjacent lower-elevation grasslands. Our results compel us to reject the tri-trophic model for this system, and are consistent with the traditional jay/oak symbiosis model. Relatively long-distance dispersal of viable acorns favors Emory oak replacement, and spatial patterns of acorn viability and curculionid parasitism suggest expansion of Emory oak into adjacent low-elevation semi-arid grasslands. Received: 29 February 1996 / Accepted: 26 September 1996     

Acorns containing deeper plumule survive better: how white oaks counter embryo excision by rodents

Several squirrel species excise the embryo of acorns of most white oak species to arrest germination for long‐term storage. However, it is not clear how these acorns counter embryo excision and survive in the arms race of coevolution. In this study, we simulated the embryo excision behavior of squirrels by removing 4 mm of cotyledon from the apical end of white oak acorns differing in embryo depths to investigate the effects of embryo excision on acorn germination and seedling performance of white oak species. The embryo depth in the cotyledons was significantly different among white oak acorns, with Quercus mongolica containing the embryo most deeply in the acorns. We found that artificial embryo excision significantly decreased acorn germination rates of Quercus variabilis, Quercus acutissima, Quercus aliena, Quercus aliena var. acutiserrata, Quercus serrata. var. brevipetiolata but not Q. mongolica. Artificial embryo excision exerted significant negative impacts on seedling performance of all oak species except Quercus aliena. Our study demonstrates the role of embryo depth of acorns in countering embryo excision by squirrels and may explain the fact that squirrels do not perform embryo excision in acorns of Q. mongolica with deeper embryos. This apparent adaptation of acorns sheds light on the coevolutionary dynamics between oaks and their seed predators.     

Utilization of red oak acorns in non-bumper crop year

Utilization of acorns from individual red oaks Quercus rubra Fagaceae) was examined in the tree canopies during fall and from the ground beneath the canopies during winter in a Missouri oak-hickory forest. The goal of the study was to determine whether vertebrates show preference for acorns from individual trees on the basis of acorn or crop characteristics. Seed production, percentage of crop which was viable, maldeveloped, or infested by insects, average seed weight and tannin content of mature, viable acorns were measured for 11 red oak adults. Behavioral observations were conducted in September and October during the peak period of seed fall. Over winter removal was determined by counting the number of acorns removed from 3x3 m plots underneath the trees' canopies between December and mid-April. Canopy removal of acorns was surprisingly low-an estimated average of 52 acorns per tree compared with a mean estimated crop size of approximately 1,200 mature acorns. Over winter, approximately 50% of the acorns were removed from ground plots (range 7% to 93.1%). Number removed corresponded more to the density of nuts rather than to seed and crop characteristics. The low rate of acorn utilization by canopy visitors contrasts with other studies in the literature and may have been due to 1) the fact that this study was done in a relatively homogeneous forest where high density of conspecifics reduces the impact on any one individual or 2) perhaps an unusually low density of birds and mammals.     

Acorn dispersal estimated by radio-tracking

Bird-dispersed seeds are difficult to track, especially in the case of long-distance dispersal events. To estimate the oak dispersal distance and the seed shadow generated by the European jay (Garrulus glandarius), we inserted radio-transmitters in 239 acorns, placed them in bird-feeders and then located them by radio-tracking. Using this methodology we located the exact caching site of 94 Quercus ilex and 54 Q. suber acorns and determined the caching habitat characteristics (vegetation type, distance, spatial distribution). The results show that: (1) there is no differences in the dispersal distance distribution between the different acorn species or sizes, (2) dispersal distances range from approximately 3 m up to approximately 550 m (mean = 68.6 m; median = 49.2 m), (3) recently abandoned fields and forest tracks were the sites preferred by jays to cache acorns, whereas fields and shrublands were avoided and (4) seed shadows showed acorn aggregation zones (i.e. clusters of caches) close to the feeder as well as isolated caches at longer distances. The results also suggest that radio-transmitters are a cheap and reliable way to determine seed shadows and quantify both seed dispersal and post-dispersal seed predation for medium to large seeds.     

Post-fire recovery of acorn production by four oak species in southern ridge sandhill association in south-central Florida

We examined post-fire recovery of two components of acorn production (percentage of bearing ramets [stems] and number of acorns per bearing ramet) for four species of oaks in southern ridge sandhill vegetation in south-central peninsular Florida. Annual counts of acorns on two white oaks (Quercus chapmanii and Q. geminata) and two red oaks (Q. laevis and Q. myrtifolia) were conducted annually (except in 1991) on two 2.7-ha grids from 1969 to 1998. A prescribed burn was conducted on one of the grids in May 1993. Newly sprouted ramets of both white oaks produced acorns during the first year following the fire, whereas red oaks required 3 yr (Q. myrtifolia) or 4 yr (Q. laevis) to produce acorns. The difference in the timing of post-fire acorn production between the white and red oak species reflected the difference in the number of years from flower bud initiation to mature acorns in the two groups, with the additional year-long lag in Q. laevis probably attributable to the fact that it is typically a tree rather than a shrub species. The data suggested that percentage of bearing ramets in the smallest size class of the two white oak species was markedly lower in the burned than unburned grid in the first year of post-fire acorn production and higher in the fifth year, but these trends were not evident for the red oaks. Among all four species, differences between mean number of acorns in burned and unburned grids were significant in only two cases (the largest size class of both white oak species in the fifth year). There was no evidence of recruitment from acorns on the burned grid, possibly due to the rapid redevelopment of the shrub layer because of low mortality of the extensive clonal root systems. Rapid post-fire recovery of acorn production in xeric fire-prone habitats is presumably the result of selection to increase the probability of recovery and persistence following sufficiently intense fires that result in high oak mortality. The timing and magnitude of post-fire acorn production in sandhill and other xeric Florida associations has a potential impact on a wide variety of insects, birds, and mammals that feed on acorns, as well as on the species with which they interact.     

Effects of forest fragmentation on the oak–rodent mutualism

Land‐use changes are expected to affect plant–disperser conditional mutualisms through changes in animal behavior. We analyzed the oak–rodent conditional mutualism in Mediterranean fragmented forests at two climatically different locations. We quantified fragmentation effects on seed dispersal effectiveness and assessed if such effects were due to changes in habitat structure and intraspecific competition for acorns in fragmented areas. Fragmentation decreased cover from predators within mouse territories as well as intraspecific competition for acorns. This resulted in lower dispersal effectiveness in small forest fragments. Globally, habitat structure was the main driver in mouse foraging decisions. In small fragments, low shelter availability precluded mouse movements, leading to short mobilization distances and low caching rates. However, as the proportion of cover from predators increased, mice were able to modulate their foraging decisions depending on intraspecific competition for acorns, resulting in higher dispersal quality. In addition to fragmentation effects, delayed breeding in the southern locality caused lower number of rodents during the dispersal season, which reduced acorn mobilization rates. Our study shows that seed dispersal patterns in managed systems can be analyzed as the result of management effects on key environmental factors in dispersers’ foraging decisions.     

昆虫与栎树的相互关系及其对栎林更新的影响

本文概述了昆虫和栎树的相互关系以及栎林种群更新研究的进展,从叶片和栎实两个方面分别分析昆虫对栎树的危害、栎树对昆虫的防御以及栎林种群更新中的问题等。（1）危害栎树的昆虫种类丰富,它们以取食叶片和栎实为主,昆虫的寄生几乎使栎实完全失去活力。（2）栎树的防御效应能影响昆虫啃食叶片的程度,被昆早寄生的栎实提前下落以及栎实产量周期性的大小年变化,能降低昆虫寄生和取食栎实所带来的损失。（3）昆虫啃食叶片降低栎树的能量利用和营养物质贮存,昆虫对栎实的寄生和取食给栎树的种子库及苗库带来压力,直接影响到栎林的种群更新,此外,昆虫对栎实的寄生也会影响脊椎动物搬运栎实时的选择取向,间接控制栎实扩散,进而影响栎树的种群更新。     

Predicting forest management effects on oak–rodent mutualisms

Wood mice Apodemus sylvaticus are the main dispersers of acorns in highly managed Mediterranean holm‐oak woodlands. Mice mobilize and cache acorns to store them for winter consumption. They carry acorns away from potential competitors, face predation risks during mobilization, and cache acorns in areas where pilfering risks are low. However, mice can act either as net predators or as moderately efficient dispersers, depending on the way landscape management affects intraspecific competition for acorns and shelter availability. To assess the influence of landscape structure and mouse behavior on acorn dispersal, we developed an agent‐based model (ABM) that translates forest management into changes in key environmental factors driving mouse foraging decisions. The model was able to predict accurately acorn dispersal patterns in a wide range of forest management practices based on information on forest habitat availability, stem density and shrub cover. Sensitivity analysis revealed that caching rates emerged from the interplay between intraspecific competition for seeds and predation risk accepted during mobilization. It also showed that intraspecific competition for acorns decreased with increasing habitat loss (due to positive edge effects on acorn production) while landscape resistance to mouse movements increased. As a result, the net benefits of caching declined and acorn predation became the dominant strategy. Finally, we assessed the effects of shrub encroachment as a management practice to enhance dispersal services in savanna‐like landscapes (dehesas). The model predicted non‐linear responses with a 65% threshold of shrub cover needed to achieve relatively high levels of acorn dispersal. This value may not be compatible with the traditional exploitation of dehesas (livestock rearing). Our study shows that integrated approaches that combine environmental change driven by management with behavioral responses of dispersers improve our understanding of the causes of recruitment bottlenecks, and are useful tools for evaluating conservation strategies aimed at enhancing dispersal services.     

Spatial and temporal patterns of seed predation on three tree species in an oak-pine forest

We experimentally examined spatial and temporal patterns of seed predation on three tree species in an oak-pine forest in southern Maine. USA, Rodents were the principal seed predators. Rates of seed loss varied with tree species and exposure to three different suites of potential seed predators (all vertebrate seed predators, medium-sized rodents [primarily squirrels] and small rodents [primarily mice and voles]), Acer rubrum seeds were removed more slowly (mean of 17.7% within 2 d across all habitats and treatments in 1991) than seeds of either Quereus rubra or Pinus strobus (e,g., > 99% removed within 2 d across all habitats and treatments for each species in 1991), Levels of final removal (cumulative removal at final census) varied with year but not with microhabitat; i.e., seed predation in four types of forest gap was not significantly different than in intact forest. Both field data and experimental feeding trials with captive Peromyscus leucopus and Clethrionomys gapperi suggest that a significant proportion of while pine seeds is eaten at time of detection, while red oak acorns are more likely to be cached. Captive Peromyscus leueopus cached and ate significantly more red oak acorns than Clethrionomys gapperi. These results suggest that red oak mast may be more important in overwintering success in Feromyscus than in Clethrionomys. Nevertheless, both species did consume red oak acorns, suggesting that in southern Maine these species are not substantially inhibited by high tannin levels in red oak acorns, as has been suggested by researchers elsewhere.     

Rodents change acorn dispersal behaviour in response to ungulate presence

Small rodents are prominent seed predators, but they also favour plant recruitment as seed dispersers. The direct interactions of ungulates on plants are more one‐sided and negative, as they mainly reduce plant recruitment through predation on seeds and seedlings. The effects of small rodents and ungulates on plant recruitment have been considered and studied as independent episodes within plant regeneration cycles. However, ungulate–rodent interactions and their potential effects on plant regeneration have not been considered so far. A number of studies have recently documented ungulate effects on the abundance, diversity and spatial distribution of small rodents. Here, we hypothesize that ungulates may also affect rodent seed dispersal behaviour. We monitored acorn dispersal by small rodents (Mus spretus and Apodemus sylvaticus) in oak woodlands with and without exclosures for large ungulates, mainly red deer, Cervus elaphus, and wild boar, Sus scrofa. The study was carried out in a typical Mediterranean Holm oak, Quercus ilex, forest throughout the acorn fall season in 2003 and 2004. We found that, in both years, the proportion of acorns cached and not recovered in the short‐term was, on average, lower in the presence (1.4%) than in the absence (19.9%) of ungulates. Acorn dispersal distances were not affected by ungulate presence in either year. However, ungulates had an effect on the spatial distribution of dispersed seeds; rodents apparently avoided shrubs as caching sites in both years. This result was interpreted as a behavioural response to reduce the risk of cache pilferage by conspecifics, which are closely associated with shrubs in presence, but not in absence, of ungulates. Potential effects of different densities of rodents or predators were discarded, as none of them differed between the areas with and without ungulates. The present study found significant interactions between heterospecific seed and seedling consumers that had been considered as independent episodes within tree regeneration cycles. As a result of such interactions, ungulates may have negative indirect effects on oak recruitment by reducing (1) acorn caching frequency, and (2) the proportion of acorns cached under shrubs, key nurse‐plants for the establishment of Holm oak seedlings in Mediterranean areas.     

外果皮厚度和种子大小对五种栎属橡子扩散的影响

动物对种子的扩散和贮藏是一个复杂的生态学过程,常常受到种子特征的影响。有关种子特征如何影响动物对种子扩散,许多研究结果并非完全一致。我们于2009 年9 月在黑龙江东方红林场野外和围栏内释放五种栎属橡子(Quercus mongolica,Q.serrata var. brevipetiolata,Q. aliena,Q.variabilis 和Q. liaotungensis),研究种子特征对鼠类(Apodemus peninsulae, Clethrionomys rufocanus 和Tamias sibiricus)扩散和埋藏橡子的影响。野外释放结果表明:橡子大小和外果皮厚度显著影响鼠类对橡子的扩散和埋藏。鼠类偏向扩散和埋藏种皮厚的大橡子,种皮薄的小橡子则多被原地取食。种皮厚的大橡子扩散距离显著高于种皮薄的小橡子。然而,只有外果皮的厚度显著影响围栏内花鼠对橡子的扩散和埋藏,橡子大小并非主要的影响因素。种子特征影响种子扩散的效应可能在种群和群落水平上存在差异。     

Tracking animal-mediated seedling establishment from dispersed acorns with the aid of the attached cotyledons

Various methods have been used to track seed dispersal of large-seeded species; however, the influence of different seed tracking methods on ecological outcomes of seed dispersal by animals is not well evaluated. Acorn removal by food hoarding animals and the following seedling establishment of Mongolian oak (Quercus mongolica) were investigated in Xiaoxing’anling Mountain, Heilongjiang, northeastern China, by using four different marking methods: plastic tagging, nail insertion, hole drilling, and isotope labeling. The acorn removal speed differed among marking methods, with plastic-tagged acorns being removed more slowly than those marked with nails, holes, and isotope. By checking the attached cotyledons and performing isotope analyses, more seedlings were found to establish from nailed acorns and isotope-soaked acorns than from drilled acorns and plastic-tagged acorns. Plastic-tagged acorns were transported closer than those marked with nails, holes, and isotope. Moreover, seedlings were often found clustered in caches containing acorns marked with plastic tags. Low level of cotyledon predation by animals makes it possible to directly identify focal seedlings of white oaks based on the attached cotyledons. Considering cotyledon predation by animals, coupling minor modification of cotyledons with isotope labeling appears to be an easy way to explore the actual pattern of seed dispersal of large-seeded trees, e.g., oaks.     

Scatter‐hoarding and cache pilfering of rodents in response to seed abundance

Seed caching and reciprocal cache pilferage play an important role in the coexistence of food‐hoarding animals. Understanding what affects seed caching and how cache pilferage occurs is an important question in seed dispersal ecology. However, tracking seed fate and cache pilferage presents substantial practical difficulties. Siberian chipmunks Tamias sibiricus always remove the entire pericarp when scatter‐hoarding acorns of Mongolian oak Quercus mongolica, whereas wood mice Apodemus peninsulae often store whole acorns in their caches. These differences in behavior provide an opportunity to investigate unilateral cache pilferage of T. sibiricus from A. peninsulae in response to seed abundance. In this study, tagged acorns were released at the peak and end periods of seed rain from Q. mongolica. This allowed us to investigate seed caching and unilateral cache pilferage at different seed abundances. We found that a higher proportion of acorns were cached at lower level of seed abundance (toward the end of seed rain), mainly because T. sibiricus rather than A. peninsulae scatter‐hoarded significantly more acorns at this time. Cache distances decreased with increasing seed abundance, indicating that acorns were cached further away and into smaller caches at lower seed abundance. Unexpectedly, unilateral cache pilferage by T. sibiricus was not significantly influenced by seed abundance—remaining at around 28% during both periods of high and low seed abundance.