Oaks, acorns, and the geographical ecology of acorn woodpeckers

We investigated the geographical ecology of acorn woodpeckers (Melanerpes formicivorus) using 30 years of Audubon Christmas Bird Counts and data on the diversity and abundance of oaks. Spatial autocorrelation in acorn woodpecker population densities is not significantly greater than zero both in either the southwestern United States, where populations are often locally isolated, or along the Pacific Coast, where they are more evenly distributed. In both regions, the effective distributional limit of acorn woodpeckers is set not by the limits of oaks but by sites where oak diversity drops to a single species. This result is consistent with acorn production patterns in central coastal California demonstrating that variability in overall acorn production and the probability of acorn crop failure decline with increasing oak species number but drop most markedly when two, compared to one, species of oaks are present together. Along the Pacific Coast, acorn woodpecker densities increase and population variability decreases with increasing abundance and diversity of oaks; however, analyses indicate that overall population size in this region is primarily determined by resource abundance while population stability is determined by resource diversity. Comparable patterns are not obvious in the Southwest, where acorn woodpecker densities are much lower than along the Pacific Coast. This may be due to a combination of greater competition for resources and oak communities that differ both qualitatively and quantitatively in their productivity compared to those along the Pacific Coast.     

Relation of ramet size to acorn production in five oak species of xeric upland habitats in south-central Florida

This study examined variation in two components of acorn production. Percentage of bearing ramets (stems) and number of acorns per bearing ramet were examined in five clonal oaks in three xeric habitats of south-central peninsular Florida in relation to ramet size within and between species and vegetative associations. Counts of acorns on two white oaks (Quercus chapmanii and Q. geminata) and three red oaks (Q. inopina, Q. laevis, and Q. myrtifolia) were conducted annually from 1969 to 1996 (except in 1991) on permanent grids in southern ridge sandhill, sand pine scrub, and scrubby flatwoods associations at the Archbold Biological Station, Florida, USA. Percentage of bearing individuals and mean number of acorns per bearing individual increased with increasing ramet size for all species across all vegetation associations. However, in Q. geminata and Q. myrtifolia, acorn production declined in the largest size class (>3.2 m), implying that larger individuals of these clonal species may become senescent. All oak species in sand pine scrub, which had a nearly closed overstory, had lower frequencies of bearing oaks and mean numbers of acorns compared with similar-sized individuals of the same species in the more open-canopied southern ridge sandhill and scrubby flatwoods associations, suggesting light limitation. The annual production of acorns by a given oak species was correlated across vegetative associations and annual acorn production of oak species was correlated for species within the same section. Intermediate-size class oaks contributed the most acorns per unit area, suggesting that stands managed with short fire-return times will provide fewer acorns to wildlife than stands managed to produce more variable distributions of oak size classes. However, our study suggests that long-unburned stands, such as those studied here, will maintain relatively constant levels of acorn production as a consequence of ramet replacement within the clones of these shrubby oaks to create a variable distribution of size classes. Of the oak species studied, Q. myrtifolia had the highest acorn production and the smallest acorns, while Q. laevis had the lowest acorn production and the largest acorns, suggesting an allocation trade-off between acorn numbers and size.     

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.     

Re-evaluation of the relationship between rodent populations and acorn masting: a review from the aspect of nutrients and defensive chemicals in acorns

The responses of rodent populations to acorn masting were examined by reviewing 17 studies from the aspect of acorn nutrients and defensive chemicals. Oak species were grouped into three types based on their acorn nutritional characteristics by cluster analysis: Type 1 acorns (two North American red oaks, subgenus Erythrobalanus) were high in tannins and high in fat and proteins (and consequently rich in metabolizable energy); Type 2 acorns (two Japanese evergreen oaks, Cyclobalanopsis; three Japanese deciduous oaks, Lepidobalanus; one North American white oak, Lepidobalanus) were high in tannins but low in fat and proteins; and Type 3 acorns (one Cyclobalanopsis species; seven Lepidobalanus species) were low in tannins and had intermediate levels of fat and proteins. These three acorn groups were nutritionally, and thereby ecologically, not equivalent. Rodents, in general, responded differently to acorn masting depending on their feeding habits and the nutritional characteristics of acorns. Granivorous rodents showed positive responses to masting of Type 1 and 3 acorns, whereas rodents with feeding habits intermediate between granivory and herbivory showed positive responses to masting of Type 3 acorns. In addition, for herbivorous rodents, the responses to masting of any types of acorns have not been reported. The present findings emphasize that the relationship between rodents and acorn masting should not easily be generalized, because there are large variations in characteristics of both acorns and rodents. The viewpoint presented in this review could offer more convincing interpretations to the contradictory observations, found in the studies reviewed, on the response of rodent populations to acorn masting.     

The coexistence of acorns with different maturation patterns explains acorn production variability in cork oak

In dry areas such as Mediterranean ecosystems, fluctuations in seed production are typically explained by resource (water) availability. However, acorn production in cork oak (Quercus suber) populations shows a very low relationship to weather. Because cork oak trees produce acorns with different maturation patterns (annual and biennial), we hypothesized that acorn production in coexisting individuals with a different dominant acorn maturation type should respond differently to climatic factors and that disaggregating the trees according to their acorn-maturation pattern should provide a more proximal relation to weather factors. We assessed acorn production variability in fragmented cork oak populations of the eastern Iberian Peninsula by counting the total number of acorns in 155 trees during an 8-year period. An initial assessment of acorn production variability in relation to weather parameters yielded very low explained variance (7%). However, after the trees were grouped according to their dominant acorn maturation pattern, weather parameters were found to account for 44% of the variability in acorn crops, with trees with annual acorns exhibiting mast fruiting in years with reduced spring frost and shorter summer droughts and trees with biennial acorns showing the opposite pattern. Thus, conditions that negatively affect annual production could be beneficial for biennial production (and vice versa). The results highlight the importance of the resource-matching hypothesis for explaining acorn production in Quercus suber and suggest that different seed maturation types within a population may allow the species to deal with highly variable weather conditions. They also emphasize the importance of understanding acorn maturation patterns for interpreting masting cycles.     

Effectiveness of rodents as local seed dispersers of Holm oaks

In this study we assessed the effectiveness of rodents as dispersers of Quercus ilex in a patchy landscape in southeastern Spain. We experimentally followed the fates of 3,200 marked and weighed acorns from dispersal through the time of seedling emergence over three years. Rodents handled about 99% of acorns, and dispersed 67% and cached 7.4% of the dispersed acorns. Most caches were recovered and consumed, and only 1.3% of the original experimental acorns were found alive in caches the following spring. Dispersal distances were short (mean = 356.2 cm, median = 157 cm) and strongly right-skewed. Heavier acorns were dispersed further and were more likely to be cached and survive than lighter acorns. All caches were in litter or soil, and each contained a single acorn. Rodents moved acorns nonrandomly, mostly to oaks and pines. Most surviving acorns were either in oaks, a poor microhabitat for oak recruitment, or shrubs, a suitable microhabitat for oak recruitment. Our results suggest that rodents, by burying a relatively high proportion of acorns singly in shrubs and pines, act as moderately effective dispersers of Q. ilex. Nonetheless, this dispersal comes at a very heavy cost.     

Acorn production prediction models for five common oak species of the eastern United States

Acorn production varies considerably among oak (Quercus) species, individual trees, years, and locations, which directly affects oak regeneration and populations of wildlife species that depend on acorns for food. Hard mast indices provide a relative ranking and basis for comparison of within- and between-year acorn crop size at a broad scale, but do not provide an estimate of actual acorn yield—the number of acorns that can potentially be produced on a given land area unit based on the species, number, and diameter at breast height (dbh) of oak trees present. We used 10 years of acorn production data from 475 oak trees to develop predictive models of potential average annual hard mast production by five common eastern oak species, based on tree diameter and estimated crown area. We found a weak (R² = 0.08–0.28) relationship between tree dbh and acorn production per unit crown area for most species. The relationship between tree dbh and acorn production per tree was stronger (R² = 0.33–0.57). However, this is because larger-dbh trees generally have larger crowns, not because they have a greater capacity to produce more acorns per unit crown area. Acorn production is highly variable among individual trees. We estimated that dbh of at least 60 dominant or codominant oak trees per species should be randomly sampled to obtain an adequate representation of the range of dbhs (≥12.7 cm dbh) in a given forest area, and achieve precise estimates when using these equations to predict potential acorn production. Our predictive models provide a tool for estimating potential acorn production that land managers and forest planners can apply to oak inventory data to tailor estimates of potential average annual acorn production to different forest management scenarios and multiple spatial scales. © 2011 The Wildlife Society.     

Acorn and aquatic invertebrate biomass in Mississippi Alluvial Valley hardwood bottomlands

The Mississippi Alluvial Valley (MAV) is an internationally important migration and wintering region for Nearctic waterfowl. Most of the MAV is a lowland forested floodplain that contains vast stands of red oaks (Quercus spp.). These trees produce acorns and, when forests flood, diverse communities of aquatic invertebrates emerge, providing diverse nutritious foods for wintering ducks. The MAV is within the Lower Mississippi Valley Joint Venture (LMV JV) region of the North American Waterfowl Management Plan, but no combined MAV-wide estimates of acorn and invertebrate biomass exist to determine foraging carrying capacity for conservation planning or actions by the LMV JV or other partners in regions containing southern red oaks. We sampled acorns that fell to the ground or were submersed under shallow water deemed accessible to foraging ducks and aquatic invertebrates in the MAV of Louisiana, Mississippi, Missouri, and Tennessee, USA, during fall-winter 2009–2011. In good and poor masting years, acorn abundance was non-linearly related to the percentage of the forest canopy made up of red oaks and peaked in late autumn or winter when most other waterfowl resources are depleted or decomposed. This finding is novel and represents a deviation from how the LMV JV has traditionally assumed food resources exist for waterfowl in hardwood bottomlands. We used a daily ration model to estimate energy use days (EUDs) from combined acorn and invertebrate biomasses relative to red oak canopy coverage. For good and poor acorn masting years at the mean MAV-wide red oak canopy coverage of 45%, EUD = 2,273.1 days/ha and 161.2 days/ha, respectively. The LMV JV currently uses EUD = 385–502 days/ha for forests with 40–50% red oak canopy coverage. Because acorns and aquatic macro-invertebrates are a food resource that persists through winter and reaches peak abundance later in winter, we contend conservation planners have undervalued the potential of bottomland hardwoods to provide energy for wintering ducks.     

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     

Two-year cycles of synchronous acorn and leaf production in biennial-fruiting evergreen oaks of subgenus Cyclobalanopsis (Quercus, Fagaceae)

Many masting species switch resources between vegetative growth and reproduction in mast and non-mast years. Although masting of oak species is well known, there have been few investigations of the relationship between vegetative growth and reproduction based on long-term monitoring data, especially in evergreen oaks of subgenus Cyclobalanopsis. We investigated annual variations over 13?years in acorn and leaf production of three evergreen oak species in subgenus Cyclobalanopsis, genus Quercus (Fagaceae)??Q. acuta, Q. salicina and Q. sessilifolia??in western Japan. In these species, the maturation of acorns occurs in the second autumn after flowering, which is known as a biennial-fruiting habit. We found a pattern of acorn production and masting in alternate years that was synchronized in all three species. Masting was not correlated with temperature and precipitation. Annual leaf-fall also showed 2-year cycle in the three oak species; peak years were synchronized between species and peak leaf-fall alternated with acorn production in all three species. Furthermore, there was a significant negative correlation between acorn and leaf production in all three species. Data showing 2-year cycles of acorn and leaf production and the negative correlation between them supports the hypothesis of resource switching between vegetative growth and reproduction. The 2-year cycle might be the basic, intrinsic rhythm of resource allocation in biennial-fruiting Cyclobalanopsis species.     

How does oak mast seeding affect the feeding behavior of sympatric red and roe deer?

Oak reproduction is characterized by mast seeding with high inter-annual fluctuations in fruit production. Such resource pulses can greatly affect ecosystem functioning and may cause seed consumers to alter their mobility, demography, or diet. Consequences of mast seeding for seed consumers remain poorly understood as their long timescale makes them difficult to study. We investigated impacts of oak mast seeding on the feeding behavior of two sympatric European deer species: red deer (Cervus elaphus) and roe deer (Capreolus capreolus). We analyzed their rumen content over a 31-year period in tandem with 10 years of data on oak fructification (i.e. 8 years of field monitoring and two modelled years). Acorn production is strongly correlated with consumption by both deer species. In years of high fructification, acorns represent more than 50% and 35% of red and roe deer diet, respectively, confirming assumptions that deer favor acorns when these are available. Red deer eat more acorns than roe deer both between and within years. High acorn production in mast years appears to saturate the capacity of deer to consume acorns. As the proportion of acorns increase in their diet, red deer eat more grasses and less conifer browse. No dietary shift was found for roe deer. By inducing dietary shifts in consumers, oak mast seeding can have cascading effects on ecosystem processes, notably on the damages on conifers caused by red deer and the consequences for forest dynamics.     

Rodent acorn selection in a Mediterranean oak landscape

Quercus suber, Quercus ilex and Quercus coccifera (Cork, Holm and Kermes oaks, respectively) are common evergreen oak species that coexist in the landscapes of the western part of the Mediterranean basin. Rodents are the main acorn predators and thus one of the main factors for understanding recruitment patterns in oaks. In this paper we analyse to what extent mice prefer acorns from one oak species over another in three oak species studied using acorn removal experiments and video tape recordings. Twenty labelled acorns from each of the three Quercus species (60 acorns) were placed in 40 cm×40 cm quadrats on each plot. Because selection might vary as a result of the vegetation context, we performed the trials in the five main vegetation types within the study area (four replicates in each vegetation type) in order to control for habitat influences on rodent acorn preferences (a total of 20 plots). The removal of 1,200 acorns occurred within 68 days. Mice removed 98.7% of the acorns. Q. ilex acorns were preferred over Q. suber and Q. coccifera in all vegetation types except in pine forest, where no acorn preferences were detected. Acorn removal rates differed with vegetation type, correlating positively with shrub cover. The distance at which acorns were displaced by rodents (mean =4.6 m±5.1 SD) did not differ between acorn species, but varied among vegetation types. Bigger acorns of Q. coccifera were selected only after Q. ilex and Q. suber acorns were depleted, while no size selection was detected for the latter two species. Thus, we conclude that rodents show preference for some oak acorns and that landscape context contributes significantly to rodent activities and decisions.     

Not only size matters: Acorn selection by the European jay (Garrulus glandarius)

A strong selection for acorn characteristics is expected to have evolved in the mutualistic relationship between the European jay (Garrulus glandarius) and the oak (Quercus spp.). Bossema's pioneer work suggested that jays do not select acorns randomly, but rather they preferentially select some size and species. Preference for some seeds over others may have implications on plant community dynamics by conferring advantages (or disadvantages) on the selected (avoided) seed characteristics. In this paper we test to what extent jays select acorns by species and/or by size and the relation between these two traits in Mediterranean oak species. The experiments consist of a set of field tests in which acorns from four different coexisting Mediterranean oak species (Quercus ilex, Quercus faginea, Quercus suber, and Quercus coccifera) were placed in artificial feeders accessible to wild jays. The acorns were previously measured to control individual acorn characteristics. Using video-recording techniques, we followed jay activity and the fate of each acorn (sequence of acorn selection and method of transport). Q. ilex acorns were preferred over other acorns, and Q. coccifera acorns were avoided when other acorns were available. Preference for Q. faginea and Q. suber acorns was intermediate, that is, they were preferred over Q. coccifera acorns but not over Q. ilex acorns. Large acorns were also preferred although acorn species selection was stronger than size selection. Jays selected species and size both by visual means and by using acorn area as an indicator of size. Acorns wider than 17–19 mm were carried in the bill because of throat limitation. Our results confirm Bossema's study on temperate oaks and extend it to Mediterranean oak species, revealing implications on mixed oak forest dynamics.     

Environmental correlates of acorn production by four species of Minnesota oaks

We measured acorn production by four species of oaks in central Minnesota over a 17-year period with the goal of understanding the proximate drivers of masting behavior. All four species exhibited significant annual variation and within-population synchrony of acorn production, although masting behavior was more pronounced in the two species that require 1 year to develop acorns (‘1-year’ species) than the two species that require 2 years (‘2-year’ species). There was also strong synchrony between species that require the same number of years to mature acorns, but not between species requiring different numbers of years. Acorn production by three of the four species correlated with spring or summer conditions, while no significant environmental correlate of acorn production was detected for the fourth species. Acorn production by none of the four species correlated significantly with variables calculated from the differences in weather conditions from 1 year to the next. These results, combined with prior studies of oaks, suggest that environmental conditions during key periods of acorn development frequently correlate with acorn production, as expected if such factors bear a direct mechanistic relationship to seed production. On the other hand, the environmental factors involve vary greatly both among species and even among populations of the same species, a result consistent with the hypothesis that environmental correlates are simply cues used by plants to synchronize reproductive investment. In either case, our results do not support the recent proposition that variables based on differences in environmental conditions from 1 year to the next serve as a general cue for masting behavior.     

Acorn production is linked to secondary growth but not to declining carbohydrate concentrations in current-year shoots of two oak species

In trees, reproduction constitutes an important resource investment which may compete with growth for resources. However, detailed analyses on how growth and fruit production interact at the shoot level are scarce. Primary canopy growth depends on the development of current-year shoots and their secondary growth might also influence the number and size of fruits supported by them. We hypothesise that an enhanced thickening of current-year shoots is linked positively to acorn production in oaks. We analysed the effect of acorn production on shoot growth of two co-occurring Mediterranean oak species with contrasting leaf habit (Quercus ilex, Quercus faginea). Length and cross-sectional area of current-year shoots, apical bud mass, number of leaves and acorns, xylem and conductive area, number of vessels of acorn-bearing and non-bearing shoots were measured in summer and autumn. Nitrogen and carbohydrates analyses were also performed in stems and leaves of both shoot types. Stem cross-sectional area increased in acorn-bearing shoots when compared with non-bearing shoots for both species and such surplus secondary growth was observed since summer. In bearing shoots, the total transversal area occupied by vessels decreased significantly from basal to apical positions along the stem as did the xylem area and the number of vessels. Leaves of bearing shoots showed lower nitrogen concentration than those of non-bearing shoots. Carbohydrate concentrations did not differ in stems and leaves as a function of the presence of acorns. Such results suggest that carbohydrates may preferentially be allocated towards reproductive shoots, possibly through enhanced secondary growth, satisfying all their carbon demands for growth and reproduction. Our findings indicate that acorn production in the two studied oaks depends on shoot secondary growth.     

The Geographic Distribution of a Tropical Montane Bird Is Limited by a Tree: Acorn Woodpeckers (Melanerpes formicivorus) and Colombian Oaks (Quercus humboldtii) in the Northern Andes

Species distributions are limited by a complex array of abiotic and biotic factors. In general, abiotic (climatic) factors are thought to explain species’ broad geographic distributions, while biotic factors regulate species’ abundance patterns at local scales. We used species distribution models to test the hypothesis that a biotic interaction with a tree, the Colombian oak (Quercus humboldtii), limits the broad-scale distribution of the Acorn Woodpecker (Melanerpes formicivorus) in the Northern Andes of South America. North American populations of Acorn Woodpeckers consume acorns from Quercus oaks and are limited by the presence of Quercus oaks. However, Acorn Woodpeckers in the Northern Andes seldom consume Colombian oak acorns (though may regularly drink sap from oak trees) and have been observed at sites without Colombian oaks, the sole species of Quercus found in South America. We found that climate-only models overpredicted Acorn Woodpecker distribution, suggesting that suitable abiotic conditions (e.g. in northern Ecuador) exist beyond the woodpecker’s southern range margin. In contrast, models that incorporate Colombian oak presence outperformed climate-only models and more accurately predicted the location of the Acorn Woodpecker’s southern range margin in southern Colombia. These findings support the hypothesis that a biotic interaction with Colombian oaks sets Acorn Woodpecker’s broad-scale geographic limit in South America, probably because Acorn Woodpeckers rely on Colombian oaks as a food resource (possibly for the oak’s sap rather than for acorns). Although empirical examples of particular plants limiting tropical birds’ distributions are scarce, we predict that similar biotic interactions may play an important role in structuring the geographic distributions of many species of tropical montane birds with specialized foraging behavior.     

人工埋藏植物种子对鼠类采食下辽东栎坚果存活及萌发率的影响

辽东栎林（Q. liaotungensis)是北京东灵山地区典型地带植被。过去研究表明,鼠类对辽东栎更新有很大影响。本研究主要调查3个问题：（1）采食辽东栎坚果的鼠类种类;（2）将坚果埋藏于土中能否减少鼠类对坚果的采食,并提高出苗率？（3）地表植被是否影响辽东栎的出苗率？1996年9月在北京东灵山地区山地选取5个类型的样地37块,每样地在地表放置10粒坚果,检查地表种子消失率。使用木板夹来调查采食坚果的鼠类种类,诱饵分别为坚果和花生米。于1997年10月,选取5块样地,间隔25 m,每样地在1m²范围内埋50粒坚果,深度5 cm,次年5月调查出苗情况。研究结果表明,大林姬鼠、社鼠和黑线姬鼠为取食坚果的主要种类,取食比例与其丰富度有关。370粒坚果放在地表后,1d、10d、20d的消失率分别达15.1%、78.1%、和 100%。250粒坚果埋入土层后,出苗率达11.2%,苗平均高度为7.8cm,出苗率与地表植被盖度有一定负相关。本研究认为,人工埋藏植物种子可以减少鼠类捕食坚果,并能增加其出苗率。     

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.     

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.     

Edge effects in a tropical montane forest mosaic: experimental tests of post-dispersal acorn removal

Forest edges created by scattered-patch clear-cutting have become common in tropical montane cloud forests in the highlands of Chiapas, Mexico. It was hypothesised that forest edges may influence regeneration of oak species, which are canopy dominants in these forests, by affecting the activities of small mammal species. Acorns of different oak species varying in germination timing were offered to predators and/or dispersers at different positions along replicated forest edges during 2 consecutive years. We investigated the effects of (1) edge type (hard and soft), (2) distance from the edge (0, 15, 30, 45 and 60 m inside forest fragments) and (3) oak species, on the rate of acorn removal mainly by small mammals. During a non-masting year, acorn removal was affected by the interaction of edge type and distance from the edge (P<0.05), with acorn removal being highest near hard edges compared to adjacent forest interiors. As predicted, acorn removal was greater along soft (100%) than along hard edges (82%), but this pattern was recorded only during the non-masting year. This study partly supports previous studies of rodents preferentially consuming acorns with early germination rather than acorns exhibiting dormancy, however these patterns may change with variation in acorn abundance. These results suggest that patch clear-cutting affects regeneration processes within forest fragments by influencing the activities of small mammals, but the nature of this effect also depends on acorn abundance and the characteristics of the forest edge created.