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

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

Context-dependent seed dispersal by a scatter-hoarding corvid

1. 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 behavior 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 behavior. 2. We investigated how seed availability and social dynamics affected scatter-hoarding in the island scrub jays (Aphelocoma insularis). We quantified rates of scatter-hoarding behavior and territorial defense of 26 colormarked birds over a three-year period with variable acorn crops. 3. 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. 4. Caching behavior 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. 5. 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

Declining fruit production before death in a widely distributed tree species, Sorbus aucuparia L.

International audienceAbstractKey messageTrees are commonly thought to increase their seed production before death. We tested this terminal investment hypothesis using long-term data on rowan trees (Sorbus aucuparia) and found no support. Rather, seed production declined significantly before death, which points to the potential detrimental effects of reproductive senescence on regeneration in stands of old trees.ContextAging poses a fundamental challenge for long-lived organisms. As mortality changes with with age due to actuarial senescence, reproductive senescence may also lead to declines in fertility. However, life history theory predicts that reproductive investment should increase before mortality to maximize lifetime reproductive success, a phenomenon termed terminal investment.AimsTo date, it is unclear whether long-lived, indeterminantly growing trees experience reproductive senescence or display terminal investment.MethodsWe investigated fruit production of rowan (Sorbus aucuparia L.), widely distributed trees that live up to 150 years, as they approached death.ResultsIn our study population in Poland’s Carpathian Mountains, 79 trees that died produced up to 20% fewer fruits in the years before their demise compared to 199 surviving trees of the same population.ConclusionThe pattern of reproductive investment in S. aucuparia is suggestive of age-independent reproductive senescence rather than terminal investment. These findings highlight that the understanding of the generality of life history strategies across diverse taxa of perennial plants is still in its infancy

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 1,0001 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

Unraveling a Paradox of Habitat Relationships: Scale-Dependent Drivers of Temporal Occupancy-Abundance Relationships in a Cooperatively Breeding Bird

Context Spatial occupancy and local abundance of species often positively covary, but the mechanisms driving this widespread relationship are poorly understood. Resource dynamics and habitat changes have been suggested as potential drivers, but long-term studies relating them to abundance and occupancy are rare. In this 34-year study of acorn woodpeckers (Melanerpes formicivorus), a cooperatively breeding species, we observed a paradoxical response to changes in habitat composition: despite a reduction in the availability of high-quality breeding habitat, the population increased considerably. Objectives We investigated the role of annual variation in food availability and long-term changes in habitat composition as predictors of population dynamics. Methods Using model selection, we contrasted competing hypotheses on the effects of changing resource availability on occupancy and social group size across three spatial scales: territory, neighborhood, and landscape. Results The increase in abundance was largely determined by the formation of new social groups, driven by a landscape-level expansion of canopy cover and its interaction with neighborhood-level acorn abundance, indicative of long-term increases in overall acorn productivity. Group size increased with neighborhood acorn crop two years earlier but groups were smaller in territories with more canopy cover. Conclusions Our results indicate that scale-dependent processes can result in paradoxical relationships in systems with spatial and temporal resource heterogeneity. Moreover, the findings support the role of resources in driving changes in abundance and occupancy at a landscape scale, suggesting that colonization of marginal habitat drives the positive occupancy-abundance relationship in this cooperatively breeding species

The ecology and evolution of synchronized reproduction in long-lived plants

Populations of many long-lived plants exhibit spatially synchronized seed production that varies extensively over time, so that seed production in some years is much higher than on average, while in others, it is much lower or absent. This phenomenon, termed masting or masting seeding, has important consequences for plant reproductive success, ecosystem dynamics and plant–human interactions. Inspired by recent advances in the field, this special issue presents a series of articles that advance the current understanding of the ecology and evolution of masting. To provide a broad overview, we reflect on the state-of-the-art of masting research in terms of underlying proximate mechanisms, ontogeny, adaptations, phylogeny and applications to conservation. While the mechanistic drivers and fitness consequences of masting have received most attention, the evolutionary history, ontogenetic trajectory and applications to plant–human interactions are poorly understood. With increased availability of long-term datasets across broader geographical and taxonomic scales, as well as advances in molecular approaches, we expect that many mysteries of masting will be solved soon. The increased understanding of this global phenomenon will provide the foundation for predictive modelling of seed crops, which will improve our ability to manage forests and agricultural fruit and nut crops in the Anthropocene. This article is part of the theme issue ‘The ecology and evolution of synchronized seed production in plants’

Caching territoriality and site preferences by a scatter-hoarder drive the spatial pattern of seed dispersal and affect seedling emergence

1. For plants with seeds dispersed by scatter-hoarders, decision-making by animals when caching determines the spatial pattern of seed dispersal and lays the initial template for recruitment, driving the regeneration of many species. However, the mechanism by which animal behaviour shapes seed distributions in spatially complex landscapes is not well understood. We investigated caching territoriality and site preferences to determine the spatial pattern of seed caching at different scales and whether scatter-hoarding behaviour drives the spatial distribution of seedling emergence. 2. We used radio-tracking and automatic wildlife cameras to monitor holm oak (Quercus ilex) acorn caching by Eurasian magpies (Pica pica), who are effective scatter-hoarders in agroforestry systems. We assessed the effect of caching territories, distance to seed source, habitat, subhabitat, microsites, and caching material in the spatial pattern of acorn dispersal by magpies. In addition, we analysed the relationship between the density of cached acorns and of emerged seedlings in different habitats. 3. Breeding magpies cached the acorns inside their caching territories, where they preferred tilled areas over oak plantations and mostly avoided old fields. These differences in habitat preference were maximized at relatively short to medium dispersal distances, where most acorns were cached, and decreased or disappeared at long-distances. Within tree plantations, magpies preferred high plant-productivity sites over low productivity ones. At the finest spatial scale, magpies preferred structures built by animals, such as rabbit grit mounds and latrines and ant litter mounds, to cache the acorns. In many sites, magpies selected uncommon materials such as stones and litter to cover caches. In the subsequent spring, seedling emergence was positively correlated with acorn cache density. 4. Synthesis. Scatter-hoarding is a hierarchical process in which caching sites are selected using different criteria at different spatial scales driven by territoriality and site preferences. Territoriality constrained dispersal distance and the habitats available for acorn caching. Magpie territoriality therefore indirectly drives oak seedling emergence and can determine oak recruitment and forest regeneration.CGL2014-459 53308-P (Ministerio Ciencia y Tecnología) PID2019-106806GB-I00 (Ministerio Ciencia e Innovación) S2013/MAE-2719 and S2018/EMT-4338 (Remedinal, CAM) CCG2014/BIO-02 & GP2019-6 (UAH) BES-2015-075276(FPI, MCT

The Maintenance of Traditions in Marmosets: Individual Habit, Not Social Conformity? A Field Experiment

Social conformity is a cornerstone of human culture because it accelerates and maintains the spread of behaviour within a group. Few empirical studies have investigated the role of social conformity in the maintenance of traditions despite an increasing body of literature on the formation of behavioural patterns in non-human animals. The current report presents a field experiment with free-ranging marmosets (Callithrix jacchus) which investigated whether social conformity is necessary for the maintenance of behavioural patterns within groups or whether individual effects such as habit formation would suffice.Using a two-action apparatus, we established alternative behavioural patterns in six family groups composed of 36 individuals. These groups experienced only one technique during a training phase and were thereafter tested with two techniques available. The monkeys reliably maintained the trained method over a period of three weeks, despite discovering the alternative technique. Three additional groups were given the same number of sessions, but those 21 individuals could freely choose the method to obtain a reward. In these control groups, an overall bias towards one of the two methods was observed, but animals with a different preference did not adjust towards the group norm. Thirteen of the fifteen animals that discovered both techniques remained with the action with which they were initially successful, independent of the group preference and the type of action (Binomial test: exp. proportion: 0.5, p<0.01).The results indicate that the maintenance of behavioural patterns within groups 1) could be explained by the first rewarded manipulation and subsequent habit formation and 2) do not require social conformity as a mechanism. After an initial spread of a behaviour throughout a group, this mechanism may lead to a superficial appearance of conformity without the involvement of such a socially and cognitively complex mechanism. This is the first time that such an experiment has been conducted with free-ranging primates

MASTREE+ : time-series of plant reproductive effort from six continents

Significant gaps remain in understanding the response of plant reproduction to environmental change. This is partly because measuring reproduction in long-lived plants requires direct observation over many years and such datasets have rarely been made publicly available. Here we introduce MASTREE+, a data set that collates reproductive time-series data from across the globe and makes these data freely available to the community. MASTREE+ includes 73,828 georeferenced observations of annual reproduction (e.g. seed and fruit counts) in perennial plant populations worldwide. These observations consist of 5971 population-level time-series from 974 species in 66 countries. The mean and median time-series length is 12.4 and 10 years respectively, and the data set includes 1122 series that extend over at least two decades (≥20 years of observations). For a subset of well-studied species, MASTREE+ includes extensive replication of time-series across geographical and climatic gradients. Here we describe the open-access data set, available as a.csv file, and we introduce an associated web-based app for data exploration. MASTREE+ will provide the basis for improved understanding of the response of long-lived plant reproduction to environmental change. Additionally, MASTREE+ will enable investigation of the ecology and evolution of reproductive strategies in perennial plants, and the role of plant reproduction as a driver of ecosystem dynamics