Variation in Avian Predation Pressure as a Driver for the Diversification of Periodical Cicada Broods.

AbstractPeriodical cicadas live 13 or 17 years underground as nymphs, then emerge in synchrony as adults to reproduce. Developmentally synchronized populations called broods rarely coexist, with one dominant brood locally excluding those that emerge in off years. Twelve modern 17-year cicada broods are believed to have descended from only three ancestral broods following the last glaciation. The mechanisms by which these daughter broods overcame exclusion by the ancestral brood to synchronously emerge in a different year, however, are elusive. Here, we demonstrate that temporal variation in the population density of generalist predators can allow intermittent opportunities for new broods to invade, even though a single brood remains dominant most of the time. We show that this mechanism is consistent, in terms of the type and frequency of brood replacements, with the distribution of periodical cicada broods throughout North America today. Although we investigate one particularly charismatic case study, the mechanisms involved (competitive exclusion, Allee effects, trait variation, predation, and temporal variability) are ubiquitous and could contribute to patterns of species diversity in a range of systems.

Periodical Cicada Emergences Affect Masting Behavior of Oaks.

AbstractOaks (Quercus spp.) are masting species exhibiting highly variable and synchronized acorn production. We investigated the hypothesis that periodical cicadas (Magicada spp.), well known to have strong effects on the ecosystems in which they occur, affect acorn production of oaks through their xylem feeding habits as nymphs, the oviposition damage they inflict as adults during emergences, or the nutrient pulse resulting from the decomposition of their bodies following breeding. We found negative effects on acorn production during emergence years and the year following emergences and enhanced acorn production 2 years after emergence. We also found evidence indicating a significant effect of cicada emergences on spatial synchrony of acorn production by trees growing within the range of the same cicada brood compared with different broods. These results demonstrate that periodical cicadas act as a trophic environmental "veto" depressing acorn production during and immediately following emergences, after which the nutrient pulse associated with the cicada's demise enhances oak reproduction.

The effect of climate change on avian offspring production: A global meta-analysis.

Climate change affects timing of reproduction in many bird species, but few studies have investigated its influence on annual reproductive output. Here, we assess changes in the annual production of young by female breeders in 201 populations of 104 bird species (N = 745,962 clutches) covering all continents between 1970 and 2019. Overall, average offspring production has declined in recent decades, but considerable differences were found among species and populations. A total of 56.7% of populations showed a declining trend in offspring production (significant in 17.4%), whereas 43.3% exhibited an increase (significant in 10.4%). The results show that climatic changes affect offspring production through compounded effects on ecological and life history traits of species. Migratory and larger-bodied species experienced reduced offspring production with increasing temperatures during the chick-rearing period, whereas smaller-bodied, sedentary species tended to produce more offspring. Likewise, multi-brooded species showed increased breeding success with increasing temperatures, whereas rising temperatures were unrelated to reproductive success in single-brooded species. Our study suggests that rapid declines in size of bird populations reported by many studies from different parts of the world are driven only to a small degree by changes in the production of young.

Lifetime inclusive fitness effects of cooperative polygamy in the acorn woodpecker.

Although over 50 y have passed since W. D. Hamilton articulated kin selection and inclusive fitness as evolutionary explanations for altruistic behavior, quantifying inclusive fitness continues to be challenging. Here, using 30 y of data and two alternative methods, we outline an approach to measure lifetime inclusive fitness effects of cooperative polygamy (mate-sharing or cobreeding) in the cooperatively breeding acorn woodpecker Melanerpes formicivorus. For both sexes, the number of offspring (observed direct fitness) declined while the number of young parented by related cobreeders (observed indirect fitness effect) increased with cobreeding coalition size. Combining these two factors, the observed inclusive fitness effect of cobreeding was greater than breeding singly for males, while the pattern for females depended on whether fitness was age-weighted, as females breeding singly accrued greater fitness at younger ages than cobreeding females. Accounting for the fitness birds would have obtained by breeding singly, however, lifetime inclusive fitness effects declined with coalition size for males, but were greater for females breeding as duos compared to breeding singly, due largely to indirect fitness effects of kin. Our analyses provide a road map for, and demonstrate the importance of, quantifying indirect fitness as a powerful evolutionary force contributing to the costs and benefits of social behaviors.

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.

Aún existen importantes vacíos en la comprensión de la respuesta reproductiva de las plantas al cambio medioambiental, en parte, porque su monitoreo en especies de plantas longevas requiere una observación directa durante muchos años, y estos conjuntos de datos rara vez han estado disponibles. Aquí presentamos a MASTREE +, una base de datos que recopila series de tiempo de la reproducción de las plantas de todo el planeta, poniendo a disposición estos datos de libre acceso para la comunidad científica. MASTREE + incluye 73.828 puntos de observación de la reproducción anual georreferenciados (ej. conteos de semillas y frutos) en poblaciones de plantas perennes en todo el mundo. Estas observaciones consisten en 5971 series temporales a nivel de población provenientes de 974 especies en 66 países. La mediana de la duración de las series de tiempo es de 10 años (media = 12.4 años) y el conjunto de datos incluye 1.122 series de al menos dos décadas (≥20 años de observaciones). Para un subconjunto de especies bien estudiadas, MASTREE +incluye un amplio conjunto de series temporales replicadas en gradientes geográficos y climáticos. Describimos el conjunto de datos de acceso abierto disponible como un archivo.csv y presentamos una aplicación web asociada para la exploración de datos. MASTREE+ proporcionará la base para mejorar la comprensión sobre la respuesta reproductiva de plantas longevas al cambio medioambiental. Además, MASTREE+ facilitará los avances en la investigación de la ecología y la evolución de las estrategias reproductivas en plantas perennes y el papel de la reproducción vegetal como determinante de la dinámica de ecosistemas.

North American tree migration paced by climate in the West, lagging in the East.

Tree fecundity and recruitment have not yet been quantified at scales needed to anticipate biogeographic shifts in response to climate change. By separating their responses, this study shows coherence across species and communities, offering the strongest support to date that migration is in progress with regional limitations on rates. The southeastern continent emerges as a fecundity hotspot, but it is situated south of population centers where high seed production could contribute to poleward population spread. By contrast, seedling success is highest in the West and North, serving to partially offset limited seed production near poleward frontiers. The evidence of fecundity and recruitment control on tree migration can inform conservation planning for the expected long-term disequilibrium between climate and forest distribution.

A brief history of masting research.

Although it has long been recognized that seed production by many forest trees varies greatly from year to year, masting (along with 'mast fruiting', 'mast seeding' and 'masting behaviour') as a concept referring to such variability is a relatively recent development. Here, I provide a brief history of masting research, highlighting some of the early contributions by foresters, zoologists and others that paved the way for the burgeoning number of studies currently being conducted by researchers around the world. Of particular current interest is work attempting to understand the proximate mechanisms, evolutionary drivers and community effects of this important ecological phenomenon as well as the ways that climate change may influence masting behaviour in the future. This article is part of the theme issue 'The ecology and evolution of synchronized seed production in plants'.

Is there tree senescence? The fecundity evidence.

Despite its importance for forest regeneration, food webs, and human economies, changes in tree fecundity with tree size and age remain largely unknown. The allometric increase with tree diameter assumed in ecological models would substantially overestimate seed contributions from large trees if fecundity eventually declines with size. Current estimates are dominated by overrepresentation of small trees in regression models. We combined global fecundity data, including a substantial representation of large trees. We compared size-fecundity relationships against traditional allometric scaling with diameter and two models based on crown architecture. All allometric models fail to describe the declining rate of increase in fecundity with diameter found for 80% of 597 species in our analysis. The strong evidence of declining fecundity, beyond what can be explained by crown architectural change, is consistent with physiological decline. A downward revision of projected fecundity of large trees can improve the next generation of forest dynamic models.

Lifetime reproductive benefits of cooperative polygamy vary for males and females in the acorn woodpecker (Melanerpes formicivorus).

Cooperative breeding strategies lead to short-term direct fitness losses when individuals forfeit or share reproduction. The direct fitness benefits of cooperative strategies are often delayed and difficult to quantify, requiring data on lifetime reproduction. Here, we use a longitudinal dataset to examine the lifetime reproductive success of cooperative polygamy in acorn woodpeckers (Melanerpes formicivorus), which nest as lone pairs or share reproduction with same-sex cobreeders. We found that males and females produced fewer young per successful nesting attempt when sharing reproduction. However, males nesting in duos and trios had longer reproductive lifespans, more lifetime nesting attempts and higher lifetime reproductive success than those breeding alone. For females, cobreeding in duos increased reproductive lifespan so the lifetime reproductive success of females nesting in duos was comparable to those nesting alone and higher than those nesting in trios. These results suggest that for male duos and trios, reproductive success alone may provide sufficient fitness benefits to explain the presence of cooperative polygamy, and the benefits of cobreeding as a duo in females are higher than previously assumed. Lifetime individual fitness data are crucial to reveal the full costs and benefits of cooperative polygamy.

Continent-wide tree fecundity driven by indirect climate effects.

Indirect climate effects on tree fecundity that come through variation in size and growth (climate-condition interactions) are not currently part of models used to predict future forests. Trends in species abundances predicted from meta-analyses and species distribution models will be misleading if they depend on the conditions of individuals. Here we find from a synthesis of tree species in North America that climate-condition interactions dominate responses through two pathways, i) effects of growth that depend on climate, and ii) effects of climate that depend on tree size. Because tree fecundity first increases and then declines with size, climate change that stimulates growth promotes a shift of small trees to more fecund sizes, but the opposite can be true for large sizes. Change the depresses growth also affects fecundity. We find a biogeographic divide, with these interactions reducing fecundity in the West and increasing it in the East. Continental-scale responses of these forests are thus driven largely by indirect effects, recommending management for climate change that considers multiple demographic rates.

Tracking the warriors and spectators of acorn woodpecker wars.

au please provide in brief summary.

Climate Dipoles as Continental Drivers of Plant and Animal Populations.

Ecological processes, such as migration and phenology, are strongly influenced by climate variability. Studying these processes often relies on associating observations of animals and plants with climate indices, such as the El Niño-Southern Oscillation (ENSO). A common characteristic of climate indices is the simultaneous emergence of opposite extremes of temperature and precipitation across continental scales, known as climate dipoles. The role of climate dipoles in shaping ecological and evolutionary processes has been largely overlooked. We review emerging evidence that climate dipoles can entrain species dynamics and offer a framework for identifying ecological dipoles using broad-scale biological data. Given future changes in climatic and atmospheric processes, climate and ecological dipoles are likely to shift in their intensity, distribution, and timing.

Mast seeding patterns are asynchronous at a continental scale.

Resource pulses are rare events with a short duration and high magnitude that drive the dynamics of both plant and animal populations and communities1. Mast seeding is perhaps the most common type of resource pulse that occurs in terrestrial ecosystems2, is characterized by the synchronous and highly variable production of seed crops by a population of perennial plants3,4, is widespread both taxonomically and geographically5, and is often associated with nutrient scarcity6. The rare production of abundant seed crops (mast events) that are orders of magnitude greater than crops during low seed years leads to high reproductive success in seed consumers and has cascading impacts in ecosystems2,7. Although it has been suggested that mast seeding is potentially synchronized at continental scales8, studies are largely constrained to local areas covering tens to hundreds of kilometres. Furthermore, summer temperature, which acts as a cue for mast seeding9, shows patterns at continental scales manifested as a juxtaposition of positive and negative anomalies that have been linked to irruptive movements of boreal seed-eating birds10,11. Here, we show a breakdown in synchrony of mast seeding patterns across space, leading to asynchrony at the continental scale. In an analysis of synchrony for a transcontinental North America tree species spanning distances of greater than 5,200 km, we found that mast seeding patterns were significantly asynchronous at distances of greater than 2,000 km apart (all P < 0.05). Other studies have shown declines in synchrony across distance, but not asynchrony. Spatiotemporal variation in summer temperatures at the continental scale drives patterns of synchrony in mast seeding, and we anticipate that this affects the spatial dynamics of numerous seed-eating communities, from insects to small mammals to the large-scale migration patterns of boreal seed-eating birds.

Biogeography and phylogeny of masting: do global patterns fit functional hypotheses?

Interannual variability of seed crops (CVp) has profound consequences for plant populations and food webs, where high CVp is termed 'masting'. Here we ask: is global variation in CVp better predicted by plant or habitat differences consistent with adaptive economies of scale, in which flower and seed benefits increase disproportionately during mast years; or by passive mechanisms, in which seed production responds to variation in resource availability associated with climate variability? To address this question, we compiled a dataset for phylogenetic comparative analysis of long-term fruit/seed production for plants comprising 920 time series spanning 311 plant species. Factors associated with both adaptive benefits of CVp (wind pollination and seed dispersal) and climatic variability (variability of summer precipitation) were among the best predictors of global variation in CVp. We observed a hump-shaped relationship between CVp and latitude and intermediate phylogenetic and geographic signals in CVp. CVp is patterned nonrandomly across the globe and over the plant tree of life, where high CVp is associated with species benefiting from economies of scale of seed or flower production and with species that experience variable rainfall over summer months when seeds usually mature.

From theory to experiments for testing the proximate mechanisms of mast seeding: an agenda for an experimental ecology.

Highly variable and synchronised production of seeds by plant populations, known as masting, is implicated in many important ecological processes, but how it arises remains poorly understood. The lack of experimental studies prevents underlying mechanisms from being explicitly tested, and thereby precludes meaningful predictions on the consequences of changing environments for plant reproductive patterns and global vegetation dynamics. Here we review the most relevant proximate drivers of masting and outline a research agenda that takes the biology of masting from a largely observational field of ecology to one rooted in mechanistic understanding. We divide the experimental framework into three main processes: resource dynamics, pollen limitation and genetic and hormonal regulation, and illustrate how specific predictions about proximate mechanisms can be tested, highlighting the few successful experiments as examples. We envision that the experiments we outline will deliver new insights into how and why masting patterns might respond to a changing environment.

Wandering woodpeckers: foray behavior in a social bird.

In many cooperatively breeding taxa, nonbreeding subordinates, or helpers, use extra-territorial forays to discover dispersal opportunities. Such forays are considered energetically costly and foraying birds face aggression from conspecific members of the territories they visit. In contrast, breeders in cooperatively breeding taxa are expected to foray seldomly. We used novel tracking technologies to follow 62 acorn woodpeckers (Melanerpes formicivorus), a cooperatively breeding bird, to study extra-territorial foray behavior. Both helpers and breeders engaged in extra-territorial forays routinely and often several times per day. Helpers forayed earlier in the day and invested more time when foraying to high-quality territories. Unexpectedly, breeders forayed as often and as far as helpers. Breeders from high-quality territories forayed closer to their home territories than breeders from low-quality territories, reflecting a potential trade-off between foraying and territory defense. Such a routine pattern of extra-territorial forays in both helpers and breeders suggests that the motives behind forays differ by sex and social status and involve more than simply searching for dispersal opportunities.

Habitat Saturation Results in Joint-Nesting Female Coalitions in a Social Bird.

Joint nesting by females and cooperative polyandry-cooperatively breeding groups with a male-biased breeder sex ratio-are little-understood, rare breeding systems. We tested alternative hypotheses of factors potentially driving these phenomena in a population of joint-nesting acorn woodpeckers (Melanerpes formicivorus). During periods of high population density and thus low independent breeding opportunities, acorn woodpecker females formed joint-nesting coalitions with close kin. Coalitions were typically associated with groups with a male bias. We found strong evidence for both inter- and intrasexual conflict, as joint nesting conferred a fitness benefit to some males, a significant fitness cost to females, and no gain in per capita reproductive output for either sex. Such conflict, particularly the cost to females, may be an important reason why joint nesting is rare among cooperatively breeding taxa.

Weather cues associated with masting behavior dampen the negative autocorrelation between past and current reproduction in oaks.

PREMISE OF THE STUDY: The influence of weather conditions on masting and the ecological advantages of this reproductive behavior have been the subject of much interest. Weather conditions act as cues influencing reproduction of individual plants, and similar responses expressed across many individuals lead to population-level synchrony in reproductive output. In turn, synchrony leads to benefits from economies of scale such as enhanced pollination success and seed predator satiation. However, there may also be individual-level benefits from reproductive responses to weather cues, which may explain the origin of masting in the absence of economies of scale. In a previous study, we found support for a mechanism whereby individual responses to weather cues attenuate the negative autocorrelation between past and current annual seed production-a pattern typically attributed to resource limitation and reproductive tradeoffs among years. METHODS: Here we provide a follow-up and more robust evaluation of this hypothesis in 12 species of oaks (Quercus spp.), testing for a negative autocorrelation (tradeoff) between past and current reproduction and whether responses to weather cues associated with masting reduce the strength of this negative autocorrelation. KEY RESULTS: Our results showed a strong negative autocorrelation for 11 of the species, and that species-specific reproductive responses to weather cues dampened this negative autocorrelation in 10 of them. CONCLUSIONS: This dampening effect presumably reflects a reduction in resource limitation or increased resource use associated with weather conditions, and suggests that responses to weather cues conferring these advantages should be selected for based on individual benefits.

Wild acorn woodpeckers recognize associations between individuals in other groups.

According to the social intelligence hypothesis, understanding the cognitive demands of the social environment is key to understanding the evolution of intelligence. Many important socio-cognitive abilities, however, have primarily been studied in a narrow subset of the social environment-within-group social interactions-despite the fact that between-group social interactions often have a substantial effect on fitness. In particular, triadic awareness (knowledge about the relationships and associations between others) is critical for navigating many types of complex social interactions, yet no existing study has investigated whether wild animals can track associations between members of other social groups. We investigated inter-group triadic awareness in wild acorn woodpeckers (Melanerpes formicivorus), a socially complex group-living bird. We presented woodpeckers with socially incongruous playbacks that simulated two outsiders from different groups calling together, and socially congruous playbacks that simulated two outsiders from the same group calling together. Subjects responded more quickly to the incongruous playbacks, suggesting that they were aware that the callers belonged to two different groups. This study provides the first demonstration that animals can recognize associations between members of other groups under natural circumstances, and highlights the importance of considering how inter-group social selection pressures may influence the evolution of cognition.