Guild structures of seed insects in relation to acorn development in two oak species

We investigated patterns of acorn growth in Quercus variabilis Blume and Quercus serrata Thunb., seasonal trends in emergence of seed insects found in the acorns, oviposition periods of the insects and falling periods of insect-infested acorns. In Q. variabilis, two insect guilds were associated with acorn development: (i) the immature acorn feeding (IAF) guild [Curculionidae sp., Poecilips cardamomi (Schaufuss), and Characoma ruficirra (Hampson)]; and (ii) the mature acorn feeding (MAF) guild [Curculio robustus (Roelofs), Curculio sikkimensis (Heller), and Cydia glandicolana (Danilevsky)]. In Q. serrata, there were three guilds: (i) the pistillate flower feeding (PFF) guild (cynipid wasp); (ii) the IAF guild [sap absorption by Mechoris ursulus (Roelofs)]; and (iii) the MAF guild (M. ursulus, C. sikkimensis, Cydia danilevskyi (Kuznetzov), C. glandicolana and Autostichidae sp.). The succession of guilds during acorn development may be a consequence of the use by different species of the limited food resource. The lack of a PFF guild in Q. variabilis that was found in our field site would have a positive effect on IAF guilds in utilizing the acorns.     

Rainfall and host reproduction regulate population dynamics of a specialist seed predator

1. Mast seeding is a widespread resource pulse caused by synchronized and intermittent production of a large seed crop by plant populations. The effects of masting on wildlife have been well documented in granivorous vertebrates, but less is known about its impact on population dynamics of insects. 2. This study investigated, over 6 years, variation in abundance of a specialist weevil (Curculio elephas) preying on holm oak (Quercus ilex) acorns. 3. An immediate bottom-up effect of seed production on weevil larval abundance was detected, which was driven by an increase in realised fecundity and aggregation at seed-rich trees. Moreover, trees producing on average more and larger acorns sustained larger weevil populations. However, no correlation was found between current and previous year adult abundance, suggesting that C. elephas did not capitalise on the reproductive bottom-up effect. 4. It was rainfall, not masting, that most strongly shaped the temporal variation in insect abundance. Rainfall facilitates emergence after diapause at underground earth cells and was also responsible for among-tree synchronisation in adult weevil population dynamics. 5. In spite of their trophic specialisation, not only food availability, but also weather affects weevil numbers. The present results indicate that moving beyond bottom-up effects is required to better understand complex systems that involve masting plants and insects that consume their seeds.     

Prolonged diapause of specialist seed-feeders makes predator satiation unstable in masting of<Emphasis Type="Italic"> Quercus crispula</Emphasis>

Quercus crispula (=Q. mongolica var. grosseserrata) is the predominant tree species in cool temperate, mixed broadleaf/conifer forests in northern Japan. We compared 11 years of data on acorn production in a population of Q. crispula, with data on seed-insect populations, to try to answer the following questions: (1) Does Q. crispula show a regular pattern of masting? (2) How long do principal seed predators remain in diapause? (3) How do the seed predators affect the pattern of predator satiation? Q. crispula showed a tendency to alternate bearing, with significant synchrony between individual trees. The principal acorn-feeding insects (Curculio spp. weevils), which infested 25%–70% of matured acorns, generally exhibited a prolonged diapause of 2 years. No significant negative relationship was found between the rate of injury by the weevils and the density of mature acorns, indicating that simple predator satiation fails due to the synchrony of the life-cycle of acorn-feeding insects and the periodical production of acorns. However, the rate of injury by the weevils was negatively correlated with the relative abundance of mature acorns to the number of weevil larvae that had matured 2 years previously. Thus, the proportion of sound acorns notably increased in a rich crop after a disturbance in alternate bearing. Prolonged diapause of specific seed predators is critical in determining the peak year of sound-seed production.     

Large acorns benefit seedling recruitment by satiating weevil larvae in <Emphasis Type="Italic">Quercus aliena</Emphasis>

Plants can enhance their seed viability and survival through reducing the fitness costs of herbivory by satiating seed predators at either plant or seed level, or both. The former satiates with production of large crops while the latter with large-sized seeds, enhancing the probability of seed survival. We analyzed both types of satiation in the interaction between the Quercus aliena and the weevil Curculio spp in 2 years with different crop size and acorn size. Larger crop size seemed to satiate less effectively weevils due to higher proportions of the seeds were attacked in oaks producing more crops of smaller acorns. However, prematurely abscission of infested acorns in large crop year might compensate this ineffectiveness. Larger acorns in lean year well satiated weevil larvae, as a larger acorn size increased the likelihood of embryo survival and seedling establishment. Although effective satiation by larger acorns would be negated by smaller seed crops, the proportion of attacked acorns that survived increased with large acorn size in lean year. These results highlighted the importance of annual variation in acorn size of Quercus aliena in satiating and defending weevil larval predations, consequently in successful seedling recruitment and establishment.     

Seed growth suppression constrains the growth of seed parasites: premature acorn abscission reduces Curculio elephas larval size

Abstract.  1. The abscission of seeds infested by insects is common in many plants and has been proposed as a defensive mechanism, although its negative consequences for insects have rarely been assessed.
2. We assessed the consequences of seed abscission studying the interactions between the holm oak Quercus ilex and the chestnut weevil Curculio elephas , its main pre-dispersal seed predator. Female weevils oviposit into the acorns and the larvae must complete their development inside a single acorn feeding on the cotyledons. The growth of the infested acorns is suppressed because they are prematurely abscised.
3. Female weevils oviposit along the acorn growing period; hence, the size of the infested acorns increased with date. The growth of the larvae inside the smaller, early infested, acorns was constrained because food (i.e. cotyledons) was frequently depleted. Larval size increased with the date along with the size of the infested acorns, but it declined slightly in the latest dates as a result of the higher conspecific competition provoked by the larger number of larvae per acorn.
4. The present results demonstrate that premature seed abscission by Q. ilex had negative consequences for C. elephas , as a smaller acorn size reduced food availability and constrained larval size, a key insect life history trait. At the same time, it is suggested that the growth suppression of infested acorns may condition the oviposition phenology of these insects according to their body size. These results need to be considered in further research in the context of the evolutionary significance of premature seed abscission as a defensive mechanism.     

Mismatch between the timing of oviposition and the seasonal optimum. The stochastic phenology of Mediterranean acorn weevils

1. The timing of reproduction is predicted to match the period of maximum food availability. In this sense, the case of many phytophagous insects in temperate regions is very illustrative, as their larvae usually depend on a resource only available for a limited period of time each year. 2. For 3 years the interactions between the weevil Curculio elephas and the Mediterranean Holm oak Quercus ilex were studied. Weevil larvae grow within the acorns, feeding on the cotyledons. The timing of oviposition will determine food availability for the larvae, as acorns stop growing once they are attacked. 3. Acorn temporal growing patterns did not change between years and food availability for larvae was at its highest in October, when temperature was still suitable for larval development. However, oviposition phenology did change between years. In 2002 females oviposited later, larvae grew within larger acorns, and their body mass was significantly higher than in 2003 or 2004, when females oviposited into early acorns. 4. Thus, weevils do not always adjust oviposition to the best possible feeding conditions for their offspring. Rather, they seem to maximise their own lifetime fitness, ovipositing as soon as they emerge in late summer. Emergence, in turn, depends strongly on stochastic events such as summer storms in the Mediterranean region. 5. Under a climate change perspective, the trend towards higher August rainfall recorded in our study area may alter oviposition phenology, with the subsequent cascade effects on weevil body size and fitness     

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 crop size and pre-dispersal predation determine inter-specific differences in the recruitment of co-occurring oaks

The contribution of pre-dispersal seed predation to inter-specific differences in recruitment remains elusive. In species with no resistance mechanisms, differences in pre-dispersal predation may arise from differences in seed abundance (plant satiation) or in the ability of seeds to survive insect infestation (seed satiation). This study aimed to analyse the impact of pre-dispersal acorn predation by weevils in two co-occurring Mediterranean oaks (Quercus ilex and Quercus humilis) and to compare its relevance with other processes involved in recruitment. We monitored the patterns of acorn production and acorn infestation by weevils and we conducted experimental tests of acorn germination after weevil infestation, post-dispersal predation and seedling establishment in mixed forests. Monitoring and experimental data were integrated in a simulation model to test for the effects of pre-dispersal predation in recruitment. In both oaks pre-dispersal acorn infestation decreased with increasing acorn crop size (plant satiation). This benefited Q. ilex which exhibited stronger masting behaviour than Q. humilis, with almost a single and outstanding reproductive event in 6 years. Acorn infestation was more than twice as high in Q. humilis (47.0%) as in Q. ilex (20.0%) irrespective of the number of seeds produced by each species. Although germination of infested acorns (seed satiation) was higher in Q. humilis (60%) than in Q. ilex (21%), this could barely mitigate the higher infestation rate in the former species, to reduce seed loss. Conversely to pre-dispersal predation, no inter-specific differences were observed either in post-dispersal predation or seedling establishment. Our results indicate that pre-dispersal predation may contribute to differences in seed supply, and ultimately in recruitment, between co-existing oaks. Moreover, they suggest that seed satiation can barely offset differences in seed infestation rates. This serves as a warning against overemphasising seed satiation as a mechanism to overcome seed predation by insects. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

槲栎超寄生橡子内象甲幼虫的适应对策

栎属植物的橡子常常受到象甲的侵害, 对橡子存活产生影响, 但有关橡子对象甲幼虫适合度影响的研究尚未见报道。本研究旨在通过对槲栎Quercus aliena种子雨进程以及象甲幼虫逃逸过程的调查, 研究二者之间的进化适应关系。结果表明： 当年槲栎的种子产量为51.92±29.26粒/m², 虫蛀率达到42.4%。虫蛀的橡子中65%以上被象甲超寄生, 且显著大于完好的橡子。在种子下落过程中, 早期从橡子中逃逸的象甲幼虫要比后期逃逸的大。对某一超寄生的橡子而言, 首先逃逸出的象甲幼虫要比随后逃逸的幼虫大。另外, 从早期脱落的橡子中逃逸的幼虫明显小于后期脱落的橡子。象甲幼虫的干重随着橡子内象甲幼虫数量的增加而显著下降。象甲幼虫寄生数量与橡子的长径和短径呈现明显的正相关关系。尽管超寄生降低了象甲虫的适合度, 但雌性成体象甲产卵过程中偏好选择大橡子在一定程度上可以减少这种不利影响。因此, 小橡子可能更容易逃脱象甲的超寄生而获得较大的存活机会。     

Seed weevils living on the edge: pressures and conflicts over body size in the endoparasitic Curculio larvae

Abstract 1. Body size in parasitic insects can be subjected to contrasting selective pressures, especially if they complete their development within a single host. On the one hand, a larger body size is associated with a higher fitness. On the other hand, the host offers a discrete amount of resources, thus constraining the evolution of a disproportionate body size. 2. The present study used the weevil Curculio elephas as a study model. Larvae develop within a single acorn, feeding on its cotyledons, and larval body size is strongly related to individual fitness. 3. The relationship between larval and acorn size was negatively exponential. Larval growth was constrained in small acorns, which did not provide enough food for the weevils to attain their potential size. Larval size increased and levelled off in acorns over a certain size (inflexion point), in which cotyledons were rarely depleted. When there were more than one larva per acorn, a larger acorn was necessary to avoid food depletion. 4. The results show that C. elephas larvae are sometimes endoparasitic, living on the edge of host holding capacity. If they were smaller they could avoid food depletion more easily, but the fitness benefits linked to a larger size have probably promoted body size increase. The strong negative effects of conspecific competition may have possibly influenced female strategy of laying a single egg per seed. 5. Being larger and fitter, but always within the limits of the available host sizes, may be one main evolutionary dilemma in endoparasites.     

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.     

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.     

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     

Complex selection on life-history traits and the maintenance of variation in exaggerated rostrum length in acorn weevils

Trophic interactions can trigger the development of exaggerated specialized characters and promote morphological diversification. For example, acorn weevils (genus Curculio) present strikingly long rostrums, which are used by females to perforate oviposition holes through the seed coat. Species exhibiting longer rostrums are known to exploit larger acorns, and therefore rostrum length is thought to be subject to selection to match the preferred acorn type. However, rostrum length is strongly correlated with body size, and morphological divergence could result from either selection on rostrum length for optimal food exploitation or from other pressures acting on body size. We collected infested acorns at oak forests where the large Curculio elephas and the small-bodied Curculio glandium co-occur. There were no interspecific differences in adult female body size to rostrum length allometric relationships, and rostrum length is equally correlated with body size in either species. MtDNA-based species identification showed that C. glandium larvae were present within acorns of all sizes, whereas C. elephas larvae were restricted to acorns above a minimum size, irrespective of oak species. Hence, exploitation of large acorns can hardly have triggered rostrum enlargement, as the small sized C. glandium adults (with short rostrums) could perforate and oviposit in both small and large acorns. Rather, increased rostrum length is probably a by-product of the larger body sizes of individuals emerging from bigger acorns, which allow increased larval size and enhance larval survival likelihood. Summarizing, when exaggerated feeding traits co-vary with other body features, interspecific morphological variability may result from contrasting selective pressures acting on these correlated characters.     

Effects of acorn masting on population dynamics of three forest-dwelling rodent species in Hokkaido,Japan

The effects of the abundance of acorns of the oak, Quercus crispula, on the population dynamics of three rodent species (Apodemus speciosus, A. argenteus, and Clethrionomys rufocanus) were analyzed using time series data (1992–2006). The data were obtained in a forest in northern Hokkaido, Japan, by live trapping rodents and directly counting acorns on the ground. Apodemus speciosus generally increased in abundance following acorn masting. However, the clear effect of acorn abundance was not detected for the other two rodent species. Acorns of Q. crispula contain tannins, which potentially have detrimental effects on herbivores. Apodemus speciosus may reduce the damage caused by acorn tannins with tannin-binding salivary proteins and tannase-producing bacteria, whereas such physiological tolerance to tannins is not known in the other two rodent species. The differences in the effects of acorns between the three species may be due to differences in their physiological tolerance to tannins.     

Satiation of predispersal seed predators: the importance of considering both plant and seed levels

Plants can reduce the fitness costs of granivory by satiating seed predators. The most common satiation mechanism is the production of large crops, which ensures that a proportion of the seeds survive predation. Nevertheless, satiation of small granivores at the seed level may also exist. Larger seeds would satiate more efficiently, enhancing the probability of seed survival after having been attacked. However, a larger seed size could compromise the efficiency of satiation by means of large crops if there were a negative relationship between seed size and the number of seeds produced by an individual plant. We analyze both types of satiation in the interaction between the holm oak Quercus ilex and the chestnut weevil Curculio elephas. Both crop size and acorn size differed strongly in a sample of 32 trees. Larger crop sizes satiated weevils, and higher proportions of the seeds were not attacked as crop size increased. Larger seeds also satiated weevil larvae, as a larger acorn size increased the likelihood of embryo survival. Seedling size was strongly related to acorn size and was reduced by weevil attack, but seedlings coming from large weeviled acorns were still larger. The number and the size of the acorns produced by individual trees were negatively related. Larger proportions of the crop were infested in oaks producing less numerous crops of larger acorns. However, contrary to expectations, these trees did not satiate more effectively at the seed level either. Effective satiation by larger acorns was precluded by larger multi-infestation rates associated to smaller seed crops, in such a way that the proportion of attacked seeds that survived did not vary among trees with different acorn sizes. These results highlight the need of considering satiation by means of large crops and large seeds in studies of predispersal seed predation. Long-term monitoring on individual oaks will help to assess whether there is a trade-off between the number and the size of the acorns and, if it existed, how it could condition the fitness consequences of both types of satiation.     

Strategies of emergence in the chestnut weevil Curculio elephas (Coleoptera: Curculionidae)

In the chestnut weevil Curculio elephas, adult emergences spread over 3 or 4 years due to prolonged larval diapause in some individuals. Weevils with an extended diapause emerge, on the average, 1–10 days before those with simple diapause, but whatever the age of insects, emergences occur always from mid-August to early October. When the summer is dry, some adults cannot emerge because of the hardness of the soil. Emergence sucess of adults is smaller in females than in males. The result is that the sex ratio is female-biased before emergence and male-biased after. Summer drought cannot be predicted by the chestnut weevil, and when the soil is dry 27–78% of females cannot emerge and do not reproduce. The year after a summer drought, many reproducing females may emerge from larvae with prolonged diapause. These results suggest an evolutionary influence on the variability in diapause duration. Computer simulations and observations do not support the hypothesis that the main cause of variation in diapause length is the existence of several distinct genotypes within populations. On the contrary, our data strengthen the hypothesis for coin-flipping plasticity discussed in a previous paper.     

Multi-trophic effects of ungulate intraguild predation on acorn weevils

Predators and parasitoids may contribute to controlling the population sizes of phytophagous insects, and this has been shown to benefit plants. Phytophagous insects may also be killed by other herbivores (intraguild predation), usually larger-sized vertebrates that ingest insects accidentally while feeding on common food sources. We studied the intraguild predation on acorn weevils by ungulates and assessed the consequences for weevil populations. Infested acorns are prematurely abscised and the weevil larvae finish their development inside the acorns after being dropped. Our results show that weevil larvae were killed by ungulates eating the infested acorns on the ground. Ungulates did not discriminate between infested and sound acorns, and the probability of a larva being incidentally eaten was inversely related to acorn availability. Thus, predation risk was enhanced by the premature drop of infested acorns when acorn availability on the ground was low. Predation rates on infested acorns were much higher where ungulates were present, and acorn infestation rates were significantly lower. However, ungulates did not provide the oaks any net benefit, since the reduction of infestation rates was not enough to compensate for the large amounts of sound acorns eaten by ungulates. Seed predation is usually studied as a progressive loss of seeds by pre- and post-dispersal predators, but the interactions between them are usually not considered. We show that intraguild predation on insects by large ungulates had an effect on the structure of the foraging guild, as the proportion of acorns predated by insects decreased; however, replicating the same experimental design in different ecological scenarios would increase the strength of these results. In conclusion, the present study shows the importance of considering the multi-trophic interactions between seed predators in order to have a complete picture of granivory.     

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.     

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.