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     

Prolonged diapause and seed predation by the acorn weevil,Curculio robustus,in relation to masting of the deciduous oak Quercus acutissima

Many researchers have studied the relationship between masting by trees and seed predation by insects. Most of these studies have been plant centered, with little focus on the insect perspective. To estimate the effect of mast seeding on insect seed predators, the life‐history traits of these insects must also be considered because some seed insects can survive lean years by prolonged diapause. In this study, I examined larval infestation of acorns and life‐history traits of the acorn weevil, Curculio robustus (Roelofs) (Coleoptera: Curculionidae), in relation to acorn production of the deciduous oak Quercus acutissima Carruthers (Fagaceae) in a coppice stand in central Japan in 2004–2009. Curculio robustus females oviposit into Q. acutissima acorns, inside which the larvae develop. Mature larvae leave acorns and burrow into the soil, where they overwinter. Although germination did occur in acorns infested by weevil larvae, the percentage of germination was lower in acorns damaged by many larvae. Acorn production in Q. acutissima varied considerably among years. Both the number of C. robustus larvae infesting acorns and the percentage of acorns infested were affected by the amount of acorns produced by Q. acutissima, and two successive lean years appeared to have a considerable impact on C. robustus population size. Consequently, only a small fraction of the acorns produced were lost to predation in a mast year after two successive lean years. However, C. robustus could survive the two successive lean years because of prolonged larval diapause, probably leading to a marked decrease in population size. These findings suggest that masting in Q. acutissima succeeds as a predator satiation strategy in response to acorn damage by C. robustus, and that C. robustus has developed prolonged diapause as a counter‐adaptation.     

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.     

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.     

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.     

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.     

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     

Zero‐sum landscape effects on acorn predation associated with shifts in granivore insect community in new holm oak (Quercus ilex) forests

Aim

Landscape attributes can determine plant–animal interactions via effects on the identity and abundance of the involved species. As most studies have been conducted in a context of habitat loss and fragmentation, we know very little about interaction assembly in new habitats from a landscape approach. This study aimed to test the effect of forest age and connectivity on acorn predation by a guild of predator insects differing in dispersal ability and resilience mechanisms: two weevils (Curculio elephas and C. glandium) and one moth (Cydia fagiglandana) in expanding Quercus ilex forests.

Location

Barcelona, Spain.

Methods

We assessed the proportion of infested acorns and identified the predator at the species level in five patches of connected old forests, connected new forests and isolated new forests. Effects of habitat age and connectivity at three scales (tree, patch and landscape) were analysed using generalized linear mixed‐effects models.

Results

Predation by weevils was positively associated with old connected forests, while moths, with better dispersal ability, were able to predate upon all patches equally. Moreover, C. elephas, the weevil with lower dispersal ability, exhibited colonization credits in the new isolated patches. In spite of these changes in the guild of seed predators, the proportion of infested acorns was non‐significantly different among forests.

Main conclusions

The guild of seed predators may vary depending on forest age and connectivity. However, because those with higher dispersal ability may replace less mobile species, this resulted in zero‐sum effects of landscape attributes on acorn predation (i.e., similar predation rates in well‐connected old forests vs. isolated new forests).

     

Ecomorphological adaptation of acorn weevils to their oviposition site

Comparisons between closely related species in different habitats provide a window into understanding how biotic factors shape evolutionary pathways. Weevils in the genus Curculio have radiated extensively across the Palearctic, where similar ecomorphs have evolved independently on different hosts. We examined ecological and morphological data for 31 Curculio species using multivariate statistics to determine which morphological traits covary and which correlate with the host seed size. A subset of 15 taxa for which phylogenetic relationships were known were used for comparative analyses and inferring historical patterns of trait evolution. The morphological analyses suggest rostrum size increased proportionately to body size in both males and females and that both rostrum and body size correlate with host seed size but that rostrum shape does not correlate with any of the seed traits used in the analyses. Host shifts from small seeds to considerably larger seeds or vice versa have occurred several times independently and historical trait evolution indicates that these host shifts were accompanied by morphological changes in rostrum size. These patterns suggest that seed size is an important selective agent for changes in rostrum length along with body size and thus may be a key factor promoting morphological differentiation in the genus Curculio.     

Acorn preference under field and laboratory conditions by two flightless Iberian dung beetle species (Thorectes baraudi and Jekelius nitidus): implications for recruitment and management of oak forests in central Spain

1. Recently, a mutualistic relationship has been described between some dung beetles (Thorectes lusitanicus and Mycotrupes lethroides) and oak species (Quercus suber, Q. canariensis, and Q. rubra), which could be crucial for ensuring seedling recruitment and sustaining the equilibrium of oak populations. For T. lusitanicus, a diet based on acorns during the reproductive period improved resistance to low‐temperature conditions and improved ovarian development. 2. In this paper, we conducted field and laboratory experiments to investigate the interaction between two potential acorn‐eating beetles, Thorectes baraudi and Jekelius nitidus, with Quercus suber. We determined the feeding preferences of both beetle species and estimated the rates of acorn manipulation by beetles according to habitat structure and several characteristics of the acorn, such as seed size and acorn infestation by weevils. 3. Results demonstrated the positive interaction between the dung beetle Thorectes baraudi and Quercus trees. Thorectes baraudi was clearly more attracted to volatiles of acorns than to dung. Jekelius nitidus, on the contrary, was either not or anecdotally attracted to acorns. On the contrary, in the case of Jekelius nitidus, the acorn attraction could be considered anecdotal or even accidental. Our field results demonstrated the acorn burying behaviour of T. baraudi in the oak forests of the Cabañeros National Park (Spain), suggesting a potential role of this beetle species as an active secondary acorn disperser. 4. This unexpected behaviour could be particularly important in Mediterranean oak forests and savannahs, where most Quercus species are strongly recruitment limited because of serious overgrazing problems.     

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.     

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.     

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.     

A Rapid Hard-Mast Index From Acorn Presence-Absence Tallies

Abstract: We used 21 years of acorn data from visual surveys of oak (Quercus spp.) trees (n = 20,113) conducted in western North Carolina, USA, to develop predictive equations for hard-mast indices (HMIs) based on the proportion of trees bearing acorns (PBA). We calculated PBA using visual estimates of the percentage of oak crown with acorns (PCA), assigning acorn presence if PCA ≥ 33.5%.The proportion of trees bearing acorns and average PCA were highly correlated, and PBA alone was a successful predictor of HMIs. Precise estimates of PBA (therefore HMIs) at a 95% confidence level required 139-385 sample trees for each oak subgroup or spatial scale of interest, and the sample size required varied with the true PBA. Substituting this faster, simpler visual survey method for others involving labor-intensive counts of twigs and acorns means that more trees can be sampled (if needed) with less time and effort for improved PBA (therefore HMI) precision. We offer a reliable method for predicting HMIs that are comparable to past HMI estimates for states using the Whitehead (1969) method, thus providing continuity in tracking long-term acorn production patterns. We also developed categories for subjectively ranking acorn crop sizes based on the range of PBA observed during 21 years. We recommend that PBA be adopted by resource management agencies as a standard, stand-alone index of acorn production to ensure comparable data among years and across the eastern United States. A standardized protocol for assigning acorn presence or absence must be used for consistent, comparable regional use of PBA in predicting HMIs or by itself as an index of acorn production.     

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.     

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.     

Sweep-Net Sampling Acorns in Forested Wetlands

ABSTRACT We are unaware of any previous studies to evaluate using a sweep net to estimate abundance of red oak acorns (Quercus spp.) after they fall from tree crowns, sink to the ground in flooded bottomlands (i.e., sound acorns), and become potential food for animals or propagules for seedlings. We placed known numbers of white-painted red oak acorns of 3 size classes and used a sweep net to recover them in a flooded hardwood bottomland in Noxubee National Wildlife Refuge, Mississippi, USA. We recovered large acorns 1.96 and 1.32 times more often than small and medium acorns, respectively. Mean recovery rate of all marked acorns across size and density classes was 34.0 ± 7.0% (SE, n = 9). Thus, sweep-net sampling for sound acorns in flooded oak bottomlands may yield negatively biased estimates of acorn abundance, and investigators should consider using correction factors.     

Adaptive divergence of scaling relationships mediates the arms race between a weevil and its host plant

Coevolution of exaggerated morphologies between insects and plants is a well-known but poorly understood phenomenon in evolutionary biology. In the antagonistic interaction between a seed-predatory insect, the camellia weevil (Curculio camelliae), and its host plant, Japanese camellia (Camellia japonica), we examined the evolutionary trajectory of an exaggerated offensive trait of the weevil (rostrum length) in terms of scaling relationship. Sampling throughout Japan revealed that the ratio of the rostrum length to overall body size was correlated with the ratio of the pericarp thickness to overall fruit size across the localities. We found a geographical interpopulation divergence in a parameter pertaining to the allometric equation of rostrum length (the coefficient a in y=axb, where y and x denote rostrum and body lengths, respectively), and the pattern of geographical differentiation in the allometric coefficient was closely correlated with the variation in the pericarp thickness of Japanese camellia. Our results provide a novel example of a geographically diverged scaling relationship in an insect morphology resulting from a coevolutionary arms race with its host plant.     

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.     

Phenotypic variation in seedlings of a “keystone” tree species (Quercus douglasii): the interactive effects of acorn source and competitive environment

Blue oak (Quercus douglasii) is a deciduous tree species endemic to California that currently exhibits poor seedling survival to sapling age classes. We used common garden techniques to examine how genetic variation at regional and local scales affected phenotypic expression in traits affecting oak seedling growth and survival. Between-population variation was examined for seedlings grown from acorns collected from a northern, mesic population and a southern, xeric population. Within-population variation was examined by comparing seedlings from different maternal families within the mesic population. Acorns were planted into neighborhoods of an annual dicot (Erodium botrys), an annual grass (Bromus diandrus), and a perennial bunchgrass (Nassella pulchra). By varying the species composition of herbaceous neighborhoods into which acorns were planted, the interactive effects of competition and acorn germplasm source on phenotypic expression could also be examined. Potential maternal effects, expressed as variation in acorn size, were assessed by weighing each acorn before planting. Probability of seedling emergence increased significantly with acorn size in the xeric population but not in the mesic population. Similarly, the effect of acorn size on seedling leaf area, stem weight, and root weight was also population-dependent. At a within-population level, acorn size effects on seedling traits varied significantly among maternal families. In addition to acorn size effects, rates of oak seedling emergence were also dependent on an interaction of population source and competitive environment. Interactions between maternal family and competitive environment in the expression of seedling leaf characters suggest the possibility of genetic variation for plasticity in traits such as specific leaf area. Using carbon isotope discrimination () as an index of relative water-use efficiency (WUE), higher water use efficiency was indicated for oak seedlings grown in the annual plant neighborhoods compared to seedlings grown in the bunchgrass neighborhood. This trend may represent an adaptive plastic response because, compared to the bunchgrass neighborhood, soil water depletion was more rapid within annual plant neighborhoods.