Bonal R., Hernández M., Espelta J.M., Muñoz A., Aparicio J.M. (2015) Unexpected consequences of a drier world: Evidence that delay in late summer rains biases the population sex ratio of an insect. Royal Society Open Science. 2: 0-0.LinkDoi: 10.1098/rsos.150198
The complexity of animal life histories makes it difficult to predict the consequences of climate change on their populations. In this paper, we show, for the first time, that longer summer drought episodes, such as those predicted for the dry Mediterranean region under climate change, may bias insect population sex ratio. Many Mediterranean organisms, like the weevil Curculio elephas, become active again after summer drought. This insect depends on late summer rainfall to soften the soil and allow adult emergence from their underground refuges. We found that, as in many protandric species, more C. elephas females emerged later in the season. Male emergence timing was on average earlier and also more dependent on the beginning of late summer rainfall. When these rains were delayed, the observed weevil sex ratio was biased towards females. So far, the effects of global warming on animal sex ratios has been reported for temperature-dependent sex determination in reptiles. Our results show that rainfall timing can also bias the sex ratio in an insect, and highlight the need for keeping a phenological perspective to predict the consequences of climate change. We must consider not just the magnitude of the predicted changes in temperature and rainfall but also the effects of their timing. © 2015 The Authors.
Fernandez-Martinez M., Garbulsky M., Penuelas J., Peguero G., Espelta J.M. (2015) Temporal trends in the enhanced vegetation index and spring weather predict seed production in Mediterranean oaks. Plant Ecology. 216: 1061-1072.LinkDoi: 10.1007/s11258-015-0489-1
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. © 2015, Springer Science+Business Media Dordrecht.
Sunyer P., Boixadera E., Munoz A., Bonal R., Espelta J.M. (2015) The interplay among acorn abundance and rodent behavior drives the spatial pattern of seedling recruitment in mature Mediterranean oak forests. PLoS ONE. 10: 0-0.LinkDoi: 10.1371/journal.pone.0129844
The patterns of seedling recruitment in animal-dispersed plants result from the interactions among environmental and behavioral variables. However, we know little on the contribution and combined effect of both kinds of variables. We designed a field study to assess the interplay between environment (vegetation structure, seed abundance, rodent abundance) and behavior (seed dispersal and predation by rodents, and rooting by wild boars), and their contribution to the spatial patterns of seedling recruitment in a Mediterranean mixed-oak forest. In a spatially explicit design, we monitored intensively all environmental and behavioral variables in fixed points at a small spatial scale from autumn to spring, as well as seedling emergence and survival. Our results revealed that the spatial patterns of seedling emergence were strongly related to acorn availability on the ground, but not by a facilitationeffect of vegetation cover. Rodents changed seed shadows generated by mother trees by dispersing most seeds from shrubby to open areas, but the spatial patterns of acorn dispersal/predation had no direct effect on recruitment. By contrast, rodents had a strong impact on recruitment as pilferers of cached seeds. Rooting by wild boars also reduced recruitment by reducing seed abundance, but also by changing rodent's behavior towards higher consumption of acorns in situ. Hence, seed abundance and the foraging behavior of scatter-hoarding rodents and wild boars are driving the spatial patterns of seedling recruitment in this mature oak forest, rather than vegetation features. The contribution of vegetation to seedling recruitment (e.g. facilitation by shrubs) may be context dependent, having a little role in closed forests, or being overridden by directed seed dispersal from shrubby to open areas. We warn about the need of using broad approaches that consider the combined action of environment and behavior to improve our knowledge on the dynamics of natural regeneration in forests. © 2015 Sunyer et al.
Munoz A., Bonal R., Espelta J.M. (2014) Acorn - weevil interactions in a mixed-oak forest: Outcomes for larval growth and plant recruitment. Forest Ecology and Management. 322: 98-105.LinkDoi: 10.1016/j.foreco.2014.02.039
Weevils are the most important pre-dispersal acorn predators in the Mediterranean region, where oaks often form mixed forests and different weevil species can coexist. The performance of weevil larvae depends in great extent on their feeding activities inside the infested acorns that, in turn, are known to reduce the viability of acorns. In this paper, we have analysed the interactions among the weevil community and four oak species (Quercus pyrenaica, Quercus suber, Quercus faginea and Quercus ilex) coexisting in a Mediterranean mixed-oak forest. DNA sequencing of weevil larvae revealed four different weevil species (Curculio elephas, Curculio glandium, Curculio pellitus and Curculio venosus) infesting the acorns of the four oak species. Oak species differed in acorn size, and weevil species also differed in body size. Weevil species showed some degree of specificity among the four oak species, but specificity was not related to variations in acorn size. By contrast, larval development and seedling recruitment were mostly driven by inter-specific differences in larval and acorn size. Larger seeded species suffered less seed damages by weevils (i.e. embryo predation and cotyledon consumption), thus reducing the impacts of acorn infestation in seedling emergence and seedling size. Larval development for the largest weevil species C. elephas was constrained by cotyledon depletion in all acorn species. Yet, this pattern was not observed for other weevil species. Larval size of the same weevil species also varied among different oak species after controlling for the amount of cotyledon eaten by larvae, thus, variation of other acorn traits among acorn species (e.g. chemical composition) may also have consequences for the performance of weevil larvae. It is likely that other variables operating at population level, such as temporal and spatial changes in acorn production or phenological variations of weevils and oaks, are also implicated in the complex functioning of these outstanding mixed-oak forests where natural regeneration seems to be threatened. © 2014 Elsevier B.V.
Peguero G., Bonal R., Espelta J.M. (2014) Variation of predator satiation and seed abortion as seed defense mechanisms across an altitudinal range. Basic and Applied Ecology. 15: 269-276.LinkDoi: 10.1016/j.baae.2014.03.006
Predator satiation and seed abortion have been reported as effective mechanisms reducing pre-dispersal seed predation, however, whether they may act simultaneously and whether their contribution to seed defense may spatially vary has been barely addressed. Across the altitudinal range of the dry tropical tree Acacia pennatula we investigated the importance of seed production and seed abortion as defense mechanisms against its pre-dispersal seed predators (Mimosestes spp.). Additionally, we measured the potential relationship between the number of seeds that escaped predation and plant recruitment. Predator satiation was effective since greater fruit production was associated with a lower proportion of predated seeds, while high seed abortion rates were related to increases in larval mortality. Although both mechanisms were present simultaneously, their relative contribution varied considerably across the altitudinal range: predator satiation was favored in the middle parts of the range, where seed production is much higher, whereas seed abortion was particularly relevant at the peripheral sites and especially high at the upper margin. The number of seeds that escaped predation was related to seedling density at plot level, indicating the demographic significance of these defense mechanisms against pre-dispersal seed predation. Overall, these results highlight the importance of considering spatial variability when analyzing seed defense traits and they also suggest considering predator satiation and seed abortion as two complementary mechanisms to reduce seed loss. © 2014 Gesellschaft für Ökologie.
Peguero G., Espelta J.M. (2014) Endozoochory and Fire as Germination Triggers in Neotropical Dry Forests: An Experimental Test. Biotropica. 46: 83-89.LinkDoi: 10.1111/btp.12076
Endozoochory and fire are crucial ecological factors determining germination success and recruitment in many plant species. Fire is a well-known germination trigger while endozoochory may allow seed dispersal along with an increase in germination. Their interaction has rarely been addressed, however, even though both factors are pervasive in human-transformed ecosystems like most Neotropical Dry Forests (NDF). For three common Mesoamerican tree species (Acacia pennatula, Enterolobium cyclocarpum, and Guazuma ulmifolia), we used feeding trials to assess the preference of cattle, which are their main seed dispersal agent. We also experimentally tested the interaction between gut passage and fire as triggers of germination. The fruits of the three species were eaten by cattle, but the small seeds of G. ulmifolia were ingested 10-fold more than those of the other species. While gut passage did not have any effect on germination, heat-shocks above 90 °C increased the number of germinating seeds by 15 percent. These results suggest that cattle may be a key dispersal vector in NDF, but that fire may be an important germination trigger. Physical dormancy in these species may have been selected for by extinct megaherbivores because it was a key trait ensuring seed survival after gut passage. However, in light of the recent expansion of cattle-ranching and fire occurrence in NDF, it has become a useful exaptation facilitating the colonization of disturbed areas. © 2013 The Association for Tropical Biology and Conservation.
Ravera F., Tarrason D., Espelta J.M. (2014) Land use change trajectories, conservation status and social importance of dry forests in Nicaragua. Environmental Conservation. : 0-0.LinkDoi: 10.1017/S0376892914000186
Interdisciplinary studies have proved the interconnectedness of history and ecology relevant to forest conservation proposals and management policies. Engaging local views and concerns in the evaluation and monitoring process can lead to more robust knowledge in the pursuit of effective conservation. This study aimed to assess the degree to which land use change trajectories influence the state of tropical dry forest conservation, as evaluated by scientists and local people. Focusing on northern Nicaragua, the research identified three historical trajectories for types and magnitude of forest disturbance. The assessment process included inventorying sites under different trajectories and integrating ecological and social indicators (namely local perceptions of biodiversity value and concern over species threat). The different land use change trajectories had no influence on the present structure of the dry forest, but strongly affected species diversity, composition and their social importance. The study provided evidence of positive species selection by farmers, which suggested a feedback loop between ecological conditions, social value and awareness of conservation. Copyright © Foundation for Environmental Conservation 2014.
Sunyer P., Espelta J.M., Bonal R., Munoz A. (2014) Seeding phenology influences wood mouse seed choices: The overlooked role of timing in the foraging decisions by seed-dispersing rodents. Behavioral Ecology and Sociobiology. 68: 1205-1213.LinkDoi: 10.1007/s00265-014-1731-x
Scatter-hoarding rodents influence the population dynamics of plants by acting as seed predators and dispersers. Therefore, rodent foraging preferences for certain seed traits (species, size, condition) have been extensively studied. However, to what extent these preferences are fixed or they track the temporal changes on seed characteristics due to phenological differences has been seldom explored. We studied the temporal variability in seed preferences by wood mouse (Apodemus sylvaticus), according to phenological changes in seed characteristics of two co-occurring oaks (Quercus ilex and Quercus pubescens). The phenology of acorn abundance and the acorn predation/dispersal patterns by rodents were monitored over an entire seeding season. Results revealed temporal changes in rodent preferences for acorns of the two oaks, matching their different seeding phenology (earlier in Q. pubescens and later in Q. ilex). On the other hand, whatever the species considered, rodents preferred larger and sound acorns along the entire season, although the dispersal of infested ones increased slightly during the peaks of acorn drop. The observed influence of seeding phenology on seed choices by rodents warns about inferring definite conclusions regarding their foraging behavior when arising from short-term experiments. Indeed, this study reveals that foraging preferences may be highly dynamic and context-dependent for some seed traits (e.g., species and condition), rather than fixed behavioral patterns. Plasticity in rodent foraging choices may allow them to successfully exploit different oaks with uncoupled seeding phenologies, while potentially favoring their coexistence. © 2014 Springer-Verlag Berlin Heidelberg.
Peguero G., Espelta J.M. (2013) Evidence for insect seed predator dynamics mediated by vertebrate frugivores. Revista Chilena de Historia Natural. 86: 161-167.LinkDoi: 10.4067/S0716-078X2013000200005
Vertebrate frugivores have been suggested to reduce seed predation, indirectly controlling populations of insect seed predators (ISP) by means of consuming many individuals when feeding on fruits. The possibility has not been explored, however, that this 'frugivore predation' may differentially affect ISP according to species-specific differences in larval development time within the fruit. In the dry tropical forest trees Acacia pennatula and Guazuma ulmifolia we compared seed predation and the absolute and relative abundances of bruchid beetle species (Bruchinae) in two sites, one with large frugivores (cattle) and the other cattle-free for a considerable time. In the site with cattle we found a notable overall reduction in the proportion of seeds predated (ca. 15 %) with respect to cattle-free site, and changes in the ISP community as well, in particular a reduced absolute and relative abundance of the bruchid species with the longest larval development time. Our results suggest that the interplay between evolutionary processes (resulting in variation in insect life-histories) and complex ecological interactions (inadvertent consumption by larger animals) may contribute to the coexistence of different insect species feeding upon the same host plant. © Sociedad de Biología de Chile.
Peguero G., Lanuza O.R., Savé R., Espelta J.M. (2013) Allelopathic potential of the neotropical dry-forest tree Acacia pennatula Benth.: Inhibition of seedling establishment exceeds facilitation under tree canopies. Plant Ecology. 213: 1945-1953.LinkDoi: 10.1007/s11258-011-0014-0
Secondary succession after land abandonment in tropical dry forests has been suggested to be favoured by the facilitation effects for seedling establishment exerted by pioneer trees isolated in these savannah-like landscapes. However, it has also been noticed that these pioneer species may sometimes have an encroaching effect and arrest succession for several decades. We investigated in this study whether allelopathy can play a role in limiting seedling establishment of co-occurring tree species under the canopy of Acacia pennatula by means of lab bioassays and field experiments in north-west Nicaragua. Leaf extracts of A. pennatula did not affect seed germination but reduced the general growth and especially the development of the root compartment in seedlings, shifting their biomass allocation model to a reduced root/shoot ratio. Survival of planted seedlings under the canopy of A. pennatula was about 20-30% lower than outside, and this reduction was particularly pronounced as the dry season progressed, despite the milder conditions (e. g. higher soil moisture) being experienced in the inner positions under the canopy. Altogether, our results suggest that, rather than facilitating, A. pennatula may inhibit the establishment of seedlings under its canopy probably by means of an allelopathic interference in the development of the root system with critical negative consequences for young seedlings in terms of overcoming the dry season. This article warns about overemphasizing the nucleation effect that remnant and isolated trees may have to facilitate secondary succession in these highly disturbed savannah-like tropical dry forests. © 2011 Springer Science+Business Media B.V.
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