Predicting forest management effects on oak–rodent mutualisms

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

Effects of forest fragmentation on the oak–rodent mutualism

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

Tree-shrub associations in grazed woodlands: first rodents,then cattle?

Spatial associations of tree saplings with spiny or toxic plants in grazed woodlands are generally explained by associational resistance, i.e., protection against grazing via a well-defended neighbor. In this study, we tested whether directed seed dispersal and post-dispersal seed removal by wood mice are additional explanations for the observed spatial association between thorny shrubs and trees, hence before associational resistance. We performed three studies in grazed woodlands in the Netherlands to test this idea. Our first seed dispersal experiment with tagged acorns showed indeed that wood mice disperse acorns directed towards shrubs. The majority of these dispersed acorns were, however, consumed. Our second experiment revealed that post-dispersal removal of cached acorns was higher under shrubs than in grassland and under trees, but also indicated the importance of within-shrub position: shrub centers were faster depleted than shrub edges. Also the number of freshly emerged seedlings and older saplings differed between micro-habitats, being higher under trees and at shrub inner-edges than at shrub outer-edges, shrub centers, and in grassland. The spatial associations with shrubs got stronger with the age of saplings, which probably reflect accumulated grazing effects over time, being higher in the open than in shrubs. We conclude that directed acorn dispersal and post-dispersal acorn removal by wood mice are two additional explaining mechanisms for the spatial associations between thorny shrubs and trees in grazed woodlands, before associational resistance. Our findings change the view that large herbivores are the sole main drivers behind tree recruitment patterns in grazed woodlands.     

Context-dependent post-dispersal predation of acorns in a California oak community

Seed dispersal and predation play important roles in plant life history by contributing to recruitment patterns in the landscape. Mast-seeding – extensive synchronized inter-annual variability in seed production – is known to influence the activity of acorn consumers at source trees, but little is known about its effect on post-dispersal predation. We conducted a planting experiment over three years to investigate the relationship between habitat-level post-dispersal predation and landscape-wide acorn production of three sympatric oak species (Quercus spp.). We measured post-dispersal predation in three oak-dominated habitats – savanna (under Q. lobata), forest edge (under Q. agrifolia), and woodland (under Q. douglasii) – as well as in chaparral and open fields. Overall, landscape-level predation was similarly high among study years, averaging 61.4%. Neither species nor mass of planted acorns affected predation. Habitat had a significant effect on post-dispersal predation risk with acorns disappearing most rapidly in chaparral and least rapidly in woodlands. However, a significant interaction between year and habitat (Z = −4.5, P < 0.001) showed that the hierarchy of predation risk among habitats was inconsistent among years. Using annual acorn census data from local populations of each oak species, we found that predation risk in oak-dominated habitats was significantly and positively related to acorn production of the overstory species (Z = −9.53, P = 0.009). Our findings add to growing evidence that seed dispersal, predation, and regeneration are context-dependent on annual variation in community-level seed production, and we discuss the potential consequences of these dynamics on oak recruitment and animal behavior.     

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

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

Application of radiotelemetry to the survey of acorn dispersal byApodemus mice

We examined the applicability of radiotelemetry to studies of acorn dispersal byApodemus mice and compared its efficiency with the of this spool-and-line method. Installation of a transmitter (2.2 g) onto acorns did not interfere with the transporting and feeding behavior of the mice. We were able to detect all transmitter-installed acorns and follow the daily changes in the sites in which they were hoarded, while we missed 59% of the spool-tied acorns due to mice breaking the threads. Mice carried transmitter-installed acorns farther than spool-tied ones. The radiotelemetry method is superior to the spool-and-line method and useful for the study of hoarding behavior in rodents.     

昆虫蛀蚀对鼠类介导下的锐齿槲栎种子扩散的影响

森林鼠类的种子贮藏行为对植物的扩散和自然更新有着非常重要的影响。然而,鼠类是否具有鉴别虫蛀种子的能力还存在一定的争议。此外,鼠类的鉴别能力是否受到食物丰富度变化的影响也未见相关报道。采用标签标记法,2011年秋季(9—11月,食物丰富季节)和2012年春季(4—6月,食物匮乏季节)分别在秦岭南坡的佛坪国家级自然保护区内,调查了森林鼠类对完好和虫蛀锐齿槲栎(Quercus aliena)种子的选择差异。结果显示:1)在秋季,尽管2种类型种子的存留动态没有显著差异,但是在后期虫蛀种子的存留时间相对更长;而在春季2种类型种子的存留动态则极为显著,几乎所有的完好种子(99%)在释放后的第3天就被鼠类全部扩散,虫蛀种子的存留时间则相对较长。2)在秋季,鼠类更喜好扩散后取食完好种子;而在春季,鼠类则喜好在原地取食绝大部分的种子,并且优先取食完好种子。3)在秋季,鼠类贮藏了更多的完好种子;而在春季,尽管完好种子在释放后第1天便达到贮藏高峰,然而由于后期的大量被捕食,2种类型种子在贮藏动态上没有显示出显著差异。研究结果表明秦岭地区森林鼠类可以准确区分完好与虫蛀种子,但是食物丰富度会影响鼠类对种子的选择策略。在食物丰富的秋季,鼠类更多地选择贮藏完好种子;而在食物相对匮乏的春季,鼠类更倾向于同时取食2种类型种子。森林鼠类通过对2种类型种子的鉴别和选择,影响不同种子的命运,从而可能对种子的扩散和自然更新产生重要影响。     

Spatial and elevational variation in fruiting phenology of a deciduous oak (Quercus crispula) and its effect on foraging behavior of the Asiatic black bear (Ursus thibetanus)

In habitats with elevational gradients, differences in the fruiting phenology of a single key food resource may affect the feeding behavior of an animal. The objectives of the present study were to assess (1) whether or not fruiting phenology and characteristics of Quercus crispula acorns differed with changes in altitude (900?C1,400?m asl) and area; (2) when bears foraged acorns in relation to their phenological development; and (3) where bears engaged in acorn foraging behavior with respect to acorn phenology. No difference in the fruiting phenology of acorns at various altitudes and locations was found, with the exception of changes in color and abscission period. Acorn abscission period occurred later at a site with an elevation of 1,200?m in Tochigi and at another with an elevation of 1,400?m in Gunma, making the available energy of acorns in the tree canopy (AET) greater and available for a longer time period at those two sites. Foraging of acorns by bears was observed at sites of moderate to high altitude between late September and mid-October. A threshold date when acorns became suitable for foraging by bears could not be identified, as the size and nutritional value of acorns increased continuously. Foraging activity of bears observed at moderate and high altitude sites corresponded with locations where AET was available in greater amounts and for a longer period of time for some sites; however, the small sample size precluded accurate assessment.     

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

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

北京东灵山落叶阔叶林中辽东栎种子雨

在北京东灵山地区的一个落叶阔叶林中调查了辽东栎（Ｑｕｅｒｃｕｓ ｌｉａｏｔｕｎｇｅｎｓｉｓ Ｋｏｉｄｚ．）的种子雨。对于选定的４棵辽东栎中的３棵,树冠下的种子雨分布格局符合二次分布,具有很高的决定系数。由设置在树冠下的种子捕捉器收集的坚果数量来估计整棵树的种子雨。４棵树的种子雨中有活力的种子很少,变化范围从２６到２５９个。每棵树的树冠下的种子雨密度变化范围从０．７６到７．２６个／ｍ＾２。林中平均种     

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

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

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     

Determinant factors in the seedling establishment of Pasania edulis (Makino) Makino

Pasania edulis (Makino) Makino is one of the dominant Fagucea tree species in evergreen broad-leaved forests in southern Japan, and its regeneration success may have a major impact on the dynamics of evergreen broad-leaved forests. We conducted a field survey on the population process from acorn production to seedling establishment of P. edulis in an evergreen broad-leaved forest in Kagoshima, southern Japan, from 1995 to 2009. The acorn crop varied greatly among the 14 cohorts, with mast cropping being recorded every 3–4 years. The mortality rate of acorns was very high for all 14 cohorts (99.3–100 %). Important mortality factors were failure to mature (empty acorns), attack by Curculio weevil on trees, predation of dropped acorns by Apodemus mice and large or medium-sized mammals (wild boar (Sus scrofa leucomystax), badger (Meles meles anakuma), and raccoon dog (Nyctereutes procyonoides viverrinu)), and attacks on germinated acorns by the acorn borer (Coccotrypes graniceps). Among these factors, predation by Apodemus mice was the greatest contributor to annual fluctuations in total mortality until seedling establishment. Large or medium-sized mammals and the acorn borer also caused severe damage to dropped acorns in some years, but contributed little to annual fluctuations in total mortality. For successful regeneration, mast cropping was essential. However, a small population of Apodemus mice, that is, a low predation pressure, was also required during mast years.     

Predation-risk effects of predator identity on the foraging behaviors of frugivorous bats

Predators directly and indirectly affect the density and the behavior of prey. These effects may potentially cascade down to lower trophic levels. In this study, we tested the effects of predator calls (playbacks of bird vocalizations: Tyto alba, Speotyto cunicularia, and Vanellus chilensis), predator visual stimuli (stuffed birds) and interactions of visual and auditory cues, on the behavior of frugivore phyllostomid bats in the field. In addition, we tested if the effects of predation risk cascade down to other trophic levels by measuring rates of seed dispersal of the tree Muntingia calabura. Using video recording, we found that bats significantly decreased the foraging frequency on trees when a visual cue of T. alba was present. However, no stimuli of potential predatory birds, including vocalization of T. alba, affected bat foraging frequency. There was a change in bat behavior during 7 min, but then their frequency of activity gradually increased. Consequently, the presence of T. alba decreased by up to ten times the rate of seed removal. These results indicate that risk sensitivity of frugivorous phyllostomid bats depends on predator identity and presence. Among the predators used in this study, only T. alba is an effective bat predator in the Neotropics. Sound stimuli of T. alba seem not to be a cue of predation risk, possibly because their vocalizations are used only for intraspecific communication. This study emphasizes the importance of evaluating different predator stimuli on the behavior of vertebrates, as well as the effects of these stimuli on trait-mediated trophic cascades.     

What Cues Do Ungulates Use to Assess Predation Risk in Dense Temperate Forests?

Anti-predator responses by ungulates can be based on habitat features or on the near-imminent threat of predators. In dense forest, cues that ungulates use to assess predation risk likely differ from half-open landscapes, as scent relative to sight is predicted to be more important. We studied, in the Białowieża Primeval Forest (Poland), whether perceived predation risk in red deer (Cervus elaphus) and wild boar (Sus scrofa) is related to habitat visibility or olfactory cues of a predator. We used camera traps in two different set-ups to record undisturbed ungulate behavior and fresh wolf (Canis lupus) scats as olfactory cue. Habitat visibility at fixed locations in deciduous old growth forest affected neither vigilance levels nor visitation rate and cumulative visitation time of both ungulate species. However, red deer showed a more than two-fold increase of vigilance level from 22% of the time present on control plots to 46% on experimental plots containing one wolf scat. Higher vigilance came at the expense of time spent foraging, which decreased from 32% to 12% while exposed to the wolf scat. These behavioral changes were most pronounced during the first week of the experiment but continuous monitoring of the plots suggested that they might last for several weeks. Wild boar did not show behavioral responses indicating higher perceived predation risk. Visitation rate and cumulative visitation time were not affected by the presence of a wolf scat in both ungulate species. The current study showed that perceived predation risk in red deer and wild boar is not related to habitat visibility in a dense forest ecosystem. However, olfactory cues of wolves affected foraging behavior of their preferred prey species red deer. We showed that odor of wolves in an ecologically equivalent dose is sufficient to create fine-scale risk factors for red deer.     

Rodents change acorn dispersal behaviour in response to ungulate presence

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

Space competition in the acorn woodpecker: Power struggles in a cooperative breeder

During a 5-year study of the cooperatively living acorn woodpecker Melanerpes formicivorus, I observed 26 irruptions of agonistic behaviour involving large numbers of intruders invading a territory. Evidence from several sources suggests that these invasions are usually contests among potential immigrants for the opportunity to replace a missing group member (hence, ‘power struggles’). Thus, power struggles indicate intense competition among individuals to transfer out of their natal group. Such dispersal is not necessarily excluded by the hypothesis that individuals gain directly by living in cooperative groups through either increased foraging efficiency or better predator defence. However, birds involved in power struggles expend considerable energy and incur considerable risks. These costs are likely to be an acceptable part of dispersal only if cooperative living is a result of birds being forced to remain in groups as a result of resource localization.     

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.     

Foraging patterns of acorn woodpeckers (Melanerpes formicivorus) on valley oak (Quercus lobata N��e) in two California oak savanna-woodlands

Landscape characteristics and social behavior can affect the foraging patterns of seed-dependent animals. We examine the movement of acorns from valley oak (Quercus lobata) trees to granaries maintained by acorn woodpeckers (Melanerpes formicivorus) in two California oak savanna-woodlands differing in the distribution of Q. lobata within each site. In 2004, we sampled Q. lobata acorns from 16 granaries at Sedgwick Reserve in Santa Barbara County and 18 granaries at Hastings Reserve in Monterey County. Sedgwick has lower site-wide density of Q. lobata than Hastings as well as different frequencies of other Quercus species common to both sites. We found acorn woodpeckers foraged from fewer Q. lobata seed source trees (K(g) = 4.1 ± 0.5) at Sedgwick than at Hastings (K(g) = 7.6 ± 0.6) and from fewer effective seed sources (N(em)* = 2.00 and 5.78, respectively). The differences between sites are due to a greater number of incidental seed sources used per granary at Hastings than at Sedgwick. We also found very low levels of seed source sharing between adjacent granaries, indicating that territoriality is strong at both sites and that each social group forages on its own subset of trees. We discovered an interesting spatial pattern in the location of granaries. At Sedgwick, acorn woodpeckers situated their granaries within areas of higher-than-average tree density, while at Hastings, they placed them within areas of lower-than-average tree density, with the outcome that granaries at the two sites were located in areas of similar valley oak density. Our results illustrate that landscape characteristics might influence the number of trees visited by acorn woodpeckers and the locations of territories, while woodpecker social behavior, such as territoriality, shapes which trees are visited and whether they are shared with other social groups.     

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.