Acorn Crop Size Research Articles

Modelling Annual Southern Appalachian Acorn Production Using Visual Surveys

ABSTRACTAcorn production varies considerably among species, locations, and years with potential effects on populations of acorn‐consuming wildlife, oak (Quercus spp.) regeneration, and forest ecology. Methods to estimate annual acorn crop size include acorn‐trapping (quantitative) or visual surveys (indices or ranks). Forest managers need a tool for converting visual survey data to quantitative estimates of acorn yield within specific years and areas. I used 7 years (2006–2010, 2012, 2016) of visual acorn survey data with acorn trap data from the same individual trees of 5 common eastern oak species in the Pisgah National Forest, North Carolina, USA, to determine whether a known within‐year relationship between the proportion of oaks bearing acorns (PBA) and acorn density using trap data (PBA‐trap) could be used to predict quantitative (no. of acorns per oak), within‐year acorn crops using visual survey data (PBA‐visual). At the individual tree level, visually determined percent crown with acorns (PCA‐visual) and number of acorns/m2 trap or per crown were correlated for all oak subgroups (species, subgenera, and all oaks combined). At the population level, mean PCA‐visual was correlated with the mean number of acorns/m2 trap or per crown for black oak (Q. velutina). Both PBA‐visual and mean PCA‐visual, and PBA‐visual and PBA‐trap were correlated for all oak subgroups. At the population level, PBA‐visual was a strong predictor of the mean number of acorns/m2 trap or per crown for most oak subgroups. Model results can be used as an index of crop size by comparing number of acorns/m2 trap or per crown among years or applied to oak inventory data to quantitatively estimate annual acorn crop sizes at the stand level or landscape level. This method enables users to estimate and compare the number of acorns produced within specific years and landscapes, rather than rely on qualitative hard‐mast indices. Published 2020. This article is a U.S. Government work and is in the public domain in the USA.

Drivers of tree fecundity in pedunculate oak (Quercus robur) refugial populations at the species' southwestern range margin.

The current low latitudinal range margins of many extra-tropical plant species consist of small and scattered populations that persist locally in microrefugia. It remains poorly understood how their refugial distribution affects mating patterns and reproductive success. Here we examine flower and acorn production and their determinants in refugial populations of the widespread European forest tree pedunculate oak (Quercus robur). We monitored male flower, female flower and acorn production in 159 adult trees from 12 oak stands over 2years. We related these and derived parameters to a series of ecological and genetic predictor variables extrinsic (stand size, density and isolation as well as elevation, topography and forest cover) or intrinsic (size, phenology and several genotypic measures) to the target tree. Tree fertility was unrelated to extrinsic factors but determined by tree size, although we detected size-independent variation in reproductive investment. Female flower number accurately predicted acorn crop size. Fruit set differed between years, evidencing the existence of pollen limitation at the landscape but not at the local scale. Fruit setalso tended to increase with the number of mates of the target tree. We detected no other evidence for genetic constraints on mating. Reproduction was triggered by a combination of small-scale and landscape-scale drivers. Although short-distance mating prevailed, limited pollen flow did not appear to significantly constrain reproductive success. The high intrinsic ability of populations to maintain their reproductive capacity may help explain their successful long-term persistence in an adverse broader environment.

Beyond predator satiation: Masting but also the effects of rainfall stochasticity on weevils drive acorn predation

AbstractEscaping seed predation is a classic “economy of scale” hypotheses (predator satiation hypothesis, Psh) to explain the selection for the synchronous production of massive and nil seed crops (masting) in plants. The Psh postulates that predator satiation occurs through a combination of (1) “functional satiation,” as not all seeds can be consumed during a massive crop, and (2) “numerical satiation,” as predator populations collapse during poor crop years. Many studies advocate for the Psh, but few have investigated the importance of masting compared to other factors for the control of predation extent. Namely, environmental cues prompting masting could also determine predator's success and, ultimately, influence directly and independently seed predation intensity. We explored this question in Mediterranean oaks, as they exhibit strong masting behavior; acorns are heavily predated upon by weevils; and rainfall stochasticity drives masting and the emergence of adult weevils from the soil. Results of two mid‐term studies (4 and 11 yr) showed that acorn production and predation were highly variable across years, while the abundance of adult weevils was positively related to autumn rainfall and to the number of infested acorns the previous years. Ultimately, acorn predation was negatively influenced by inter‐annual fluctuation of seed production (masting) yet, mainly and positively, prompted by autumn rainfall and acorn crop size (only in one site). Our results highlight the relevance of masting to reduce seed predation. Yet evidences that rainfall stochasticity directly determines the success of weevils, and it independently influences seed predation extent, indicate that environmental cues prompting masting may also fine‐tune the output of this reproductive behavior. Additionally, local differences suggest that the relevance of masting may change with tree characteristics (low vs. high seed production) and landscape structure (isolated vs. dense forests). We also discuss what can be the effects of increasing drought in Mediterranean areas for this antagonistic interaction, triggered by rainfall.

Context-dependent seed dispersal by a scatter-hoarding corvid.

Corvids (crows, jays, magpies and nutcrackers) are important dispersers of large-seeded plants. Studies on captive or supplemented birds suggest that they flexibly adjust their scatter-hoarding behaviour to the context of social dynamics and relative seed availability. Because many corvid-dispersed trees show high annual variation in seed production, context-dependent foraging can have strong effects on natural corvid scatter-hoarding behaviour. We investigated how seed availability and social dynamics affected scatter-hoarding in the island scrub jays (Aphelocoma insularis). We quantified rates of scatter-hoarding behaviour and territorial defence of 26 colour-marked birds over a three-year period with variable acorn crops. We tested whether caching parameters were correlated with variation in annual seed production of oaks as predicted by the predator dispersal hypothesis, which states that caching rates and distances should vary with seed abundance in ways that benefit tree fitness. We also tested whether antagonistic interactions with conspecifics would affect scatter-hoarding adversely, as found in experimental studies. Caching behaviour varied with acorn availability. Caching distances correlated positively with annual acorn crop size, increasing by as much as 40% between years. Caching rates declined over time in years with small acorn crops, but increased when crops were large. Acorn foraging and caching rates were also negatively correlated with rates of territorial aggression. Overall foraging rates, however, were not associated with aggression, suggesting that reduced dispersal rates were not simply due to time constraints. Our field results support laboratory findings that caching rates and distances by scatter-hoarding corvids are context-dependent. Furthermore, our results are consistent with predictions of the predator dispersal hypothesis and suggest that large seed crops and social interactions among scatter-hoarders affect dispersal benefits for oaks and other masting tree species.

The effect of within-year variation in acorn crop size on seed harvesting by avian hoarders.

Spatial and temporal variation in resource distribution affect the movement and foraging behavior of many animals. In the case of animal-dispersed trees, numerous studies have addressed masting-the synchronized variation in seed production between years-but the fitness consequences of spatial variation in seed production within a year are unclear. We investigated the effects of variable acorn production in a population of valley oaks (Quercus lobata) on the composition and behavior of the avian-disperser community. We found that western scrub-jays (Aphelocoma californica), high-quality dispersers that store seeds in the ground, were attracted to, and exhibited increased per capita dispersal rates from, trees with large acorn crops. In contrast, acorn woodpeckers (Melanerpes formicivorus), low-quality dispersers that store acorns in trees where they are unlikely to germinate, increased per capita hoarding rates but did not attend trees with large seed crops in higher numbers, suggesting that the two species responded to resources on different spatial scales. Antagonistic interactions within and between species increased with the number of birds attending a tree, resulting in a potential cost for foraging birds, but did not reduce dispersal rates. Using a simulation model, we estimated that trees with large initial crops experienced a greater proportion (77 %) of high-quality seed dispersal events than trees with small crops (62 %). Our findings provide support for a mechanistic link between seed production and foraging behavior of seed dispersers as predicted by the predator dispersal hypothesis for the functional consequences of variable seed production in hoarder-dispersed trees.

Temporal trends in the enhanced vegetation index and spring weather predict seed production in Mediterranean oaks

The extremely year-to-year variable production of seeds (masting) is an extended plant reproductive behaviour important for forest dynamics and food webs. The dependence of these episodes of massive seed production on recently or long-term photosynthesised carbohydrates, however, remains controversial. In this paper, we explore whether vegetation (tree canopy) changes, detected using EVI as a proxy of leaf area and photosynthetic capacity, can provide a reliable estimation of seed production. To complete this analysis, we also explored the effect of weather both in the trends of EVI and in acorn crop size. To this end, we compared the trends of the EVI and acorn production over 10 years (2000–2009) in five stands of Quercus ilex L. in Barcelona (Catalonia, NE Spain). We found that acorn production was mainly driven by a combination of: (i) a minimum initial threshold in the EVI values, (ii) an increase in EVI in the 9 ± 4 months prior to reproduction, and (iii) appropriate weather conditions (low water stress) during spring. These results indicated, apparently for the first time, that reproduction in masting species could be detected and partly predicted by remotely sensed vegetative indices. Our results suggested that this particular reproductive behaviour in Mediterranean oaks was driven by a combination of two factors, i.e. good and improving vegetation conditions, as shown by a minimum initial threshold and the increase in EVI needed for large seed crops, and the need of wet weather conditions during spring. Moreover, our results fully supported recent studies that have associated short-term photosynthate production with seed production.

Cues versus proximate drivers: testing the mechanism behind masting behavior

Masting, the intermittent and synchronized production of seeds, is a common and important phenomenon throughout the plant kingdom. Surprisingly, the proximate mechanisms by which populations of masting plants synchronize their seed sets have been relatively unexplored. We examined how temperature influences the acorn crop of the valley oak Quercus lobata, a masting species common in California, USA, over 33 years in order to assess whether temperature acts directly on acorn crop as a cue or whether it acts instead through intermediate steps indicative of a direct mechanistic connection to acorn production. Compared to several alternatives, the difference in temperature during the spring flowering period over the prior two years (Δt) was a good predictor of annual acorn crop in valley oak, as proposed recently by Kelly et al. Significantly, Δt correlates positively with temperatures the previous April, a likely driver of pollination success in valley oak, and negatively with the previous year's acorn crop, which is in turn negatively correlated with the current year's acorn crop, presumably due to resource limitation. Thus, the success of Δt is not as a cue but rather explained by its close relationship to the proximate drivers that have a direct, mechanistic relationship with acorn crop size.

Masting in oaks: Disentangling the effect of flowering phenology, airborne pollen load and drought

Quercus species exhibit an extreme inter-annual variability in seed production often synchronized over large geographical areas (masting). Since this reproductive behavior is mostly observed in anemophilous plants, pollination efficiency is suggested as one hypothesis to explain it, although resource-based hypotheses are also suggested as alternatives. We analyzed the effect of flowering phenology, airborne pollen presence and meteorological conditions in the pattern of acorn production in mixed evergreen-deciduous oak forests (Quercus ilex and Quercus pubescens) in NE Spain for twelve years (1998–2009). In both oaks, higher temperatures advanced the onset of flowering and increased the amount of airborne pollen. Nevertheless, inter-annual differences in pollen production did not influence acorn crop size. Acorn production was enhanced by a delay in flowering onset in Q. ilex but not in Q. pubescens. This suggests that in perennial oaks a larger number of photosynthates produced before flowering could benefit reproduction while the lack of effects on deciduous oaks could be because these species flush new leaves and flowers at the same time. Notwithstanding this effect, spring water deficit was the most relevant factor in explaining inter-annual variability in acorn production in both species. Considering that future climate scenarios predict progressive warmer and dryer spring seasons in the Mediterranean Basin, this might result in earlier onsets of flowering and higher water deficits that would constrain acorn production.

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

AbstractAcorn 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.

Effects of asynchronous acorn production by co-occurring Quercus trees on resource utilization by acorn-feeding insects

To clarify the effects of asynchronous seed production among tree species on the population of seed predators, we investigated the relationship between the annual variation in production of mature acorns and the insect damage in those acorns of two sympatric oak species, Quercus variabilis Blume and Quercus serrata Thunb. ex Murray, over 4 years at two study sites. The annual variation in acorn production was noticeable, with a coefficient of variation (CV) at the two sites of 1.05 and 0.80 for Q. variabilis and 0.87 and 0.73 for Q. serrata. Annual fluctuation in acorn production by Q. serrata was synchronized between the two sites. Since annual fluctuation in acorn production was not synchronized between the two species, the CVs for the total acorn production by both oak species (0.83 and 0.62 at the two sites) were lower than those for Q. variabilis and Q. serrata alone. The rate of predation by the specialist predators (Curculio weevils) on the acorns of both species was not related to the annual acorn crop size. Prolonged diapause of Curculio weevils might stabilize their populations. The rate of acorn predation by the generalist predators (tortricid moths) was also not related to the annual crop size. Asynchronous acorn production by the two oak species would help to stabilize the population.

Increased drought reduces acorn production in Quercus ilex coppices: thinning mitigates this effect but only in the short term

Summary In order to explore the effects of climate change on Mediterranean regenerating forests, we experimentally assessed the effects of increased drought on the reproductive attributes of Quercus ilex over a 4-year period (2005 – 2008). We also investigated whether traditional thinning (selection of one to a few stems per stump) could mitigate the consequences of increased drought in oak coppices. Increased drought reduced the number of reproductive trees, mean number of female fl owers produced and acorn crop size, although most of these effects appeared only in the last 2 years of the experiment. In a different way, thinning enhanced all reproductive attributes, but its main effects were transient and covered only 1 or 2 years after the application of the treatments. Our results indicate that a moderate reduction in rainfall (15 per cent ) reduces the reproductive ability of Q. ilex . This may have long-term negative consequences for recruitment as well as for the fauna feeding on acorns. Although traditional thinning may mitigate the consequences of increased drought, it has a remarkably short-term effect. This highlights the need to re-examine traditional forestry practices as potential adaptive strategies for coping with climate change in Mediterranean regenerating forests.

Spatial patterns of acorn dispersal by rodents: do acorn crop size and ungulate presence matter?

Seed dispersal is qualitatively effective when it increases recruitment probability. A poorly studied factor likely affecting recruitment is the spatial distribution of dispersed seeds. Seed-caching animals are thought to disperse seeds in a way that reduces clumping and density to impede cache pilfering. Furthermore, dispersal might differ depending on whether the seed is immediately consumed versus cached for later consumption, and might differ depending on the ecological context. The main objectives of this study were to determine: 1) the spatial pattern of seed dispersal by rodents in a heterogeneous environment; 2) whether the patterns differ among years and among acorn competitor exclosure treatments, and 3) whether rodents create different spatial patterns of dispersal for acorns that are cached versus consumed immediately following dispersal. We studied the degree of spatial aggregation of acorn dispersal by rodents using two different estimators derived from the Ripley K and the Diggle G functions. We also analyzed various metrics of dispersal distances. For both analyses we used observed acorn dispersal patterns in two years differing in crop size and inside versus outside exclosures restricting access to acorn-consuming ungulates. During 2003, a year with a larger crop size, maximum seed dispersal distances were less, and the pattern of dispersed seeds was more clumped, than in 2004, a year with a smaller crop size. Median dispersal distances did not differ between years. In the presence of ungulates, seed dispersal was marginally sparser than in their absence. Cached acorns were dispersed more sparsely than acorns eaten immediately. These results have important implications for the quality of seed dispersal for oak recruitment that are likely relevant to other systems as well.

Pre‐dispersal acorn predation in mixed oak forests: interspecific differences are driven by the interplay among seed phenology, seed size and predator size

Summary1. Pre‐dispersal seed predation (PSP) often occurs in multi‐host–predator systems (e.g. several plant species exposed to a common array of granivorous insects). However, whether the interaction among seed phenology, seed size and predator size accounts for interspecific differences in PSP remains elusive.2. We studied PSP in a mixed‐oak forest with two oaks (the larger‐seeded Quercus humilis and the smaller‐seeded Q. ilex), both depredated by two acorn weevils (the smaller Curculio glandium and the larger C. elephas). We intensively monitored acorn production and infestation phenology and we identified the weevil species depredating acorns by means of DNA taxonomy.3. The minimum acorn size required for infestation was lower for C. glandium than for C. elephas, in accordance with their different body sizes. This resulted in an earlier infestation phenology in C. glandium and the ability of this species to infest both smaller and larger acorns. Above a minimum acorn size threshold, no selection for larger acorns by weevils was observed.4. Initial acorn crop size was similar in the two oaks. Nonetheless, the earlier acorn phenology and the production of larger acorns in Q. humilis favoured the earlier infestation by C. glandium and the predation by both small and large weevils. Smaller acorns of Q. ilex almost excluded infestation by the larger C. elephas.5. Although larger acorns of Q. humilis could better survive infestation (preserve the embryo), higher PSP in this species finally resulted in a lower mature acorn crop size than in Q. ilex.6. Synthesis. In a multi‐host–predator system, smaller‐seeded species may benefit from a reduced PSP because they exclude larger granivorous insects, but also by means of a ‘free‐rider effect’, if larger‐seeded heterospecifics earlier reach a critical size to be depredated. These results also highlight the benefits of a small body size in granivorous insects to depredate seeds earlier and to forage on a wider range of seed sizes. Whether the advantage of ‘being small’ in this antagonistic plant–animal interaction is offset by other processes, or whether it results in a pressure towards seed and insect size reduction, deserves further attention.

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.

No Trade‐Off between Seed Size and Number in the Valley OakQuercus lobata

We examined the relationship between acorn mass and number in valley oaks (Quercus lobata) over 4 years in central coastal California. Despite considerable variation in acorn size among both trees and years, trees produced acorns of the same size relative to other trees in different years. Across years, the relationship between acorn mass and acorn crop size was generally positive, even after controlling for environmental conditions and differences in individual tree size and quality. Life-history trade-offs in valley oaks are primarily between current and future reproduction and indirectly between concurrent growth and reproduction, not between seed size and number, and are probably related to this species' mast-seeding behavior. Phenotypic trade-offs in long-lived plants such as oaks exhibit complex patterns of life-history covariation and deserve greater attention, both theoretically and empirically.

Influence of physiographic and climatic factors on spatial patterns of acorn production in Maryland and Virginia, USA

AbstractAim The aims of this study were to identify the effects physiographic differences have on the spatial synchrony of acorn production within red (Quercus rubra) and white (Quercus alba) oak subgenus groups, to identify climatic factors associated with acorn production patterns, and to assess if and how these relationships vary across a distinct physiographic boundary.Location Maryland and Virginia in the eastern United States.Methods Using data from 36 survey sites in Virginia (1973–2000) and Maryland (1977–2000), we described the spatial synchrony of annual acorn production based on Mantel tests and modified correlograms at three spatial scales: (1) the entire study area, (2) within two ecological regions (mountain and Piedmont), and (3) between these ecoregions. Using climate data obtained from the National Climate Data Centre, we described the relationship between climate and acorn production within each ecological region using linear regression.Results Spatial synchrony in the white oak group acorn production was strong among sites closest together, and declined with distance. The average April temperature during the year of acorn drop was positively related to acorn crop size within the mountain and Piedmont ecoregions. Spatial synchrony in the red oak group was strong among sites closest together and declined with distance in the mountain region. Synchrony was weaker in the Piedmont than in the mountain ecoregion and declined slightly with increasing distance. Between the mountain and Piedmont ecoregions, synchrony was not detected among the closest sites, but was detected with increasing strength as the distance between site pairs increased. In the Piedmont ecoregion, the number of April freeze events and total July precipitation, both 2 years prior to acorn drop, were negatively related to red oak acorn production. In the mountains, average July temperature 2 years prior to acorn drop was positively related to acorn production.Main conclusions Physiographic differences affected the spatial acorn production synchrony in the red oak but not in the white oak subgenus group. Climatic factors related to annual acorn production variability differed between subgenus groups and physiographic regions. The physiographic differences between the mountain and Piedmont ecoregions may define spatial patterns of acorn production synchrony in some oak species and mediate climatic influences on acorn production.

MASTING MEDIATED BY SUMMER DROUGHT REDUCES ACORN PREDATION IN MEDITERRANEAN OAK FORESTS

Temporally variable production of seed crops by perennial plants (masting) has been hypothesized to be a valuable mechanism in the reduction of seed predation by satiating and starving seed consumers. To achieve these benefits, coexisting species subjected to the same predator would benefit from a similar pattern of seeding fluctuation over time that could lead to a reduction in predation at the within-species level. We tested for the existence of an environmental factor enforcing synchrony in acorn production in two sympatric Mediterranean oaks (Quercus ilex and Q. humilis) and the consequences on within-species and between-species acorn predation, by monitoring 15 mixed forests (450 trees) over seven years. Acorn production in Q. ilex and Q. humilis was highly variable among years, with high population variability (CVp) values. The two species exhibited a very different pattern across years in their initial acorn crop size (sum of aborted, depredated, and sound acorns). Nevertheless, interannual differences in summer water stress modified the likelihood of abortion during acorn ripening and enforced within- and, particularly, between-species synchrony and population variability in acorn production. The increase in CVp from initial to mature acorn crop (after summer) accounted for 33% in Q. ilex, 59% in Q. humilis, and 60% in the two species together. Mean yearly acorn pre-dispersal predation by invertebrates was considerably higher in Q. humilis than in Q. ilex. Satiation and starvation of predators was recorded for the two oaks, and this effect was increased by the year-to-year variability in the size of the acorn crop of the two species combined. Moreover, at a longer time scale (over seven years), we observed a significant reduction in the mean proportion of acorns depredated for each oak and the variability in both species' acorn production combined. Therefore, our results demonstrate that similar patterns of seeding fluctuation over time in coexisting species mediated by an environmental cue (summer drought) may contribute to the reduction of the impact of seed predation at a within-species level. Future research should be aimed at addressing whether this process could be a factor assisting in the coexistence of Q. ilex and Q. humilis.

Pollen-limited reproduction in blue oak: implications for wind pollination in fragmented populations.

Human activities are fragmenting forests and woodlands worldwide, but the impact of reduced tree population densities on pollen transfer in wind-pollinated trees is poorly understood. In a 4-year study, we evaluated relationships among stand density, pollen availability, and seed production in a thinned and fragmented population of blue oak (Quercus douglasii). Geographic coordinates were established and flowering interval determined for 100 contiguous trees. The number of neighboring trees within 60m that released pollen during each tree's flowering period was calculated and relationships with acorn production explored using multiple regression. We evaluated the effects of female flower production, average temperature, and relative humidity during the pollination period, and number of pollen-producing neighbors on individual trees' acorn production. All factors except temperature were significant in at least one of the years of our study, but the combination of factors influencing acorn production varied among years. In 1996, a year of large acorn crop size, acorn production was significantly positively associated with number of neighboring pollen producers and density of female flowers. In 1997, 1998, and 1999, many trees produced few or no acorns, and significant associations between number of pollen-producing neighbors and acorn production were only apparent among moderately to highly reproductive trees. Acorn production by these reproductive trees in 1997 was significantly positively associated with number of neighboring pollen producers and significantly negatively associated with average relative humidity during the pollination period. In 1998, no analysis was possible, because too few trees produced a moderate to large acorn crop. Only density of female flowers was significantly associated with acorn production of moderately to highly reproductive trees in 1999. The effect of spatial scale was also investigated by conducting analyses with pollen producers counted in radii ranging from 30m to 80m. The association between number of pollen-producing neighbors and acorn production was strongest when neighborhood sizes of 60m or larger were considered. Our results suggest that fragmentation and thinning of blue oak woodlands may reduce pollen availability and limit reproduction in this wind-pollinated species.

Acorn production by oaks in central coastal California: influence of weather at three levels

Mean annual acorn crop size of oaks in central coastal California correlated with weather for four of five species. For the deciduous Quercuslobata and Quercusdouglasii, larger crops were produced in years when April temperatures were warmest, while for the evergreen Quercusagrifalia and Quercuschrysolepis, crops were larger when there was more rain one and two years prior to the year when acorns were produced. No correlations were found between weather and mean annual acorn production by Quercuskelloggii. Within years, individual Q. douglasii that flowered during periods of greater solar radiation produced more acorns, and during 1 of 5 years individual Q. lobata and Q. agrifolia produced larger crops when they flowered during warm, dry periods. These results indicate that conditions favorable for pollination and fertilization strongly enhance mean annual acorn production in Q. lobata and Q. douglasii and may have an effect on differences in acorn crop size among individual Q. lobata, Q. douglasii, and Q. agrifolia. Mean annual acorn crops of these three species were significantly concordant among three sites in central coastal California 300 km apart. Weather conditions were also significantly correlated, especially annual rainfall and mean April temperatures. Thus, the environmental factors that have the strongest influence on mean annual acorn production within a site are also those that are the most geographically concordant. However, we found no relationship between environmental factors and differences in mean annual acorn production by these same three species at the three sites.

Ecology of Mast‐Fruiting in Three Species of North American Deciduous Oaks

We conducted an 8—yr study of flower and acorn production in three species of oak in east—central Missouri: white oak (Quercus alba), red oak (Q. rubra), and Black oak (Q. velutina). The goal was to evaluate whether mast—fruiting, the synchronous production of large seed crops followed by small seed crops, is simply a response to weather conditions or is actually an evolved reproductive strategy. In this paper, we address four specific questions: (1) Are annual fluctuations in acorn crop size the result of synchronous production of acorns by individuals? (2) Are large acorn crops the result of large flower crops and/or high survival of those flowers to mature fruit? (3) To what extent do weather variables account for variation in acorn production among years? (4) Does acorn crop size correlate negatively with prior acorn production? Red oak and white oak showed a greater degree of mast—fruiting than did black oak. Within a species, individuals tended to produce large acorn crops in the same years, but each species differed in which years they produced large crops. The size of a given acorn crop was determined by both flower abundance and survival of flowers to fruit. Principal components and multiple—regression analyses were used to describe the relationship between weather variables and acorn production. The first principal component explained the largest amount of the variation in black oak (R2 = 0.55) and the red oak (R2 = 0.89). In white oak, two principal components combined to explain 77% of the variation in acorn production. The weather variables that were associated with these principal components included spring temperature (positive effect) and summer drought (negative effect). Past acorn production had a major impact on the size of the current acorn crop, with each species showing a different pattern. In black oak, the current acorn crop was negatively correlated with the crop 3 yr prior but positively correlated with the crop 2 yr prior. In red oak, acorn crop size 1, 2, and 3 yr prior had negative correlations with current acorn crop, while acorn crop size 4 yr prior was positively correlated. In white oak, there were negative correlations between acorn crop size and crop size 1, 2, and 4 yr prior but a positive correlation with the acorn crop size 3 yr prior. These data are consistent with the hypothesis that mast—fruiting species must store resources during some years in order to produce a mast crop. We discuss the possibility that these three species may have inherent cycles of reproduction that are modified by the impact of weather conditions, black oak with 2—yr cycles, white oak with 3—yr cycles, and red oak with 4—yr cycles. We conclude that the patterns of acorn production for black oak, red oak, and white oak are not simply responses to weather events but are also a function of prior reproductive events. This suggests that masting is an evolved reproductive strategy.