Mario B. Pesendorfer

Pollen limitation and flower abortion in a wind‐pollinated, masting tree

Pollen limitation is a key assumption of theories that explain mast seeding, which is common among wind-pollinated and woody plants. In particular, the pollen coupling hypothesis and pollination Moran effect hypothesis assume pollen limitation as a factor that synchronizes seed crops across individuals. The existence of pollen limitation has not, however, been unambiguously demonstrated in wind-pollinated, masting trees. We conducted a two-year pollen supplementation experiment on a masting oak species, Quercus lobata. Supplemental pollen increased acorn set in one year but not in the other, supporting the importance of pollen coupling and pollination Moran effect models of mast seeding. We also tracked the fate of female flowers over five years and found that the vast majority of flowers were aborted for reasons unrelated to pollination, even in the presence of excess pollen. Pollen limitation can reduce annual seed set in a wind-pollinated tree, but factors other than pollen limitation cause the majority of flower abortion.

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.

Competing for seed dispersal: evidence for the role of avian seed hoarders in mediating apparent predation among oaks

Summary In communities of large-seeded tree species, generalist seed hoarders often link the temporally variable dynamics of various species in ways that can result in indirect ecological interactions. A special case of such interactions is “apparent predation” – when variation in seed production of one tree species changes the outcome of the interaction between seed hoarders and another tree species from seed dispersal mutualism to predation. We investigated how two species of avian hoarders responded to differences in acorn availability among individual valley oaks (Quercus lobata) and across a landscape dominated by blue oaks (Q. douglasii) to determine whether seed dispersal and predation dynamics result in apparent predation between these oak species. California scrub-jays (Aphelocoma californica), high-quality seed dispersers with large spatial scales of resource perception, showed strong numerical and functional responses to individually large Q. lobata acorn crops when landscape-level acorn crops were low, but the numerical response was not observed when Q. douglasii produced a good acorn crop the subsequent year. In contrast, acorn woodpeckers (Melanerpes formicivorus), which are significant acorn predators, exhibited a consistent functional response to individual Q. lobata acorn crops independent of landscape-wide availability. Consequently, Q. lobata experienced increased predation and a near absence of high-quality seed dispersal when Q. douglasii acorns were abundant. Our results suggest that apparent predation among oak species can arise from contrasting responses of generalist seed predators and dispersers to acorn availability on different spatial scales. Thus, the spatial scale of resource perception, which can differ greatly among generalist seed hoarders, may provide a proximate mechanism underlying indirect, temporally-variable ecological affects such as apparent predation among sympatric species of large-seeded trees. This article is protected by copyright. All rights reserved.

Testing the Terminal Investment Hypothesis in California Oaks

The terminal investment hypothesis—which proposes that reproductive investment should increase with age-related declines in reproductive value—has garnered support in a range of animal species but has not been previously examined in long-lived plants, such as trees. We tested this hypothesis by comparing relative acorn production and radial growth among 1,000+ mature individuals of eight species of California oaks (genus Quercus) followed for up to 37 years, during which time 70 trees died apparently natural deaths. We found no significant differences in the radial growth, acorn production, or index of reproductive effort, taking into consideration both growth and reproduction among dying trees relative to either conspecific trees at the same site that did not die or growth and reproduction from earlier years for the focal trees that did eventually die. Furthermore, we found no consistent trade-off between growth and reproduction among trees that died, nor did dying trees significantly alter their relative investment in reproduction even as they underwent physical decline. Trees approaching the end of their lives are often in poor physical condition but do not appear to differentially invest more of their diminished resources in reproduction compared with healthy trees.

Drivers of synchrony of acorn production in the valley oak (Quercus lobata) at two spatial scales

We investigated spatial synchrony of acorn production by valley oaks (Quercus lobata) among individual trees at the within-population, local level and at the among-population, statewide level spanning the geographic range of the species. At the local level, the main drivers of spatial synchrony were water availability and flowering phenology of individual trees, while proximity, temperature differences between trees, and genetic similarity failed to explain a significant proportion of variance in spatial synchrony. At the statewide level, annual rainfall was the primary driver, while proximity was significant by itself but not when controlling for rainfall; genetic similarity was again not significant. These results support the hypothesis that environmental factors, the Moran effect, are key drivers of spatial synchrony in acorn production at both small and large geographic scales. The specific environmental factors differed depending on the geographic scale, but were in both cases related to water availability. In addition, flowering phenology, potentially affecting either density-independent pollination failure (the pollination Moran effect) or density-dependent pollination efficiency (pollen coupling), plays a key role in driving spatial synchrony at the local geographic scale.

Spatially biased dispersal of acorns by a scatter-hoarding corvid may accelerate passive restoration of oak habitat on California’s largest island

Abstract Scatter hoarding by corvids (crows, jays, magpies, and nutcrackers) provides seed dispersal for many large-seeded plants, including oaks and pines. When hoarding seeds, corvids often choose nonrandom locations throughout the landscape, resulting in differential survival of seeds. In the context of habitat restoration, such disproportional storing of seeds in areas suitable for germination and establishment can accelerate expansion and recovery of large-seeded tree populations and their associated ecosystems. Here, we investigate the spatial preferences of island scrub jays Aphelocoma insularis during scatter hoarding of acorns (Quercus spp.) on Santa Cruz Island. We use a large behavioral data set on the birds’ behavior in combination with seedling surveys and spatial analysis to determine whether 1) island scrub jays disproportionally cache seeds in specific habitat types, and 2) whether the preferred habitat type is suitable for oak regeneration. Our results show that the jays nonrandomly cache acorns across the landscape; they use chaparral and coastal sage scrub disproportionally while avoiding open and grassy areas. The areas used most often for caching were also the areas with the highest oak seedling densities. We discuss the potential role of these findings for the recovery of Santa Cruz Island’s oak habitat since the 1980s.

Oak habitat recovery on California's largest islands: Scenarios for the role of corvid seed dispersal

Seed dispersal by birds is central to the passive restoration of many tree communities. Reintroduction of extinct seed dispersers can therefore restore degraded forests and woodlands. To test this, we constructed a spatially explicit simulation model, parameterized with field data, to consider the effect of different seed dispersal scenarios on the extent of oak populations. We applied the model to two islands in Californias Channel Islands National Park (USA), one of which has lost a key seed disperser. 2. We used an ensemble modelling approach to simulate island scrub oak (Quercus pacifica) demography. The model was developed and trained to recreate known population changes over a 20-year period on 250-km(2) Santa Cruz Island, and incorporated acorn dispersal by island scrub-jays (Aphelocoma insularis), deer mice (Peromyscus maniculatus) and gravity, as well as seed predation. We applied the trained model to 215-km(2) Santa Rosa Island to examine how reintroducing island scrub-jays would affect the rate and pattern of oak population expansion. Oak habitat on Santa Rosa Island has been greatly reduced from its historical extent due to past grazing by introduced ungulates, the last of which were removed by 2011. 3. Our simulation model predicts that a seed dispersal scenario including island scrub-jays would increase the extent of the island scrub oak population on Santa Rosa Island by 281% over 100 years, and by 544% over 200 years. Scenarios without jays would result in little expansion. Simulated long-distance seed dispersal by jays also facilitates establishment of discontinuous patches of oaks, and increases their elevational distribution. 4. Synthesis and applications. Scenario planning provides powerful decision support for conservation managers. We used ensemble modelling of plant demographic and seed dispersal processes to investigate whether the reintroduction of seed dispersers could provide cost-effective means of achieving broader ecosystem restoration goals on Californias second-largest island. The simulation model, extensively parameterized with field data, suggests that re-establishing the mutualism with seed-hoarding jays would accelerate the expansion of island scrub oak, which could benefit myriad species of conservation concern.

Effects of mistletoe (Phoradendron villosum) on California oaks

Mistletoes are a widespread group of plants often considered to be hemiparasitic, having detrimental effects on growth and survival of their hosts. We studied the effects of the Pacific mistletoe, Phoradendron villosum, a member of a largely autotrophic genus, on three species of deciduous California oaks. We found no effects of mistletoe presence on radial growth or survivorship and detected a significant positive relationship between mistletoe and acorn production. This latter result is potentially explained by the tendency of P. villosum to be present on larger trees growing in nitrogen-rich soils or, alternatively, by a preference for healthy, acorn-producing trees by birds that potentially disperse mistletoe. Our results indicate that the negative consequences of Phoradendron presence on their hosts are negligible—this species resembles an epiphyte more than a parasite—and outweighed by the important ecosystem services mistletoe provides.