Functional Traits Predict Outcomes of Current and Novel Competition Under Warmer Climate.

Functional traits offer a potential avenue to generalize and forecast the impacts of changing competition on plant communities, including changing outcomes of competition among species that currently interact (current competition) or that will interact in the future following range shifts (novel competition). However, it remains unclear how well traits explain variation in the outcomes of current and novel competition as well as the underlying processes determining coexistence or competitive exclusion, under changing climate. Here, we interacted pairs of high and low-elevation species in three sites across an elevation gradient in the Swiss Alps. For each species pair, we quantified the population-level outcomes of competition (invasion growth rates), relative fitness differences, and niche overlap and related these to 15 functional traits that were measured in each site. Most traits were significantly associated with invasion growth rates at the low elevation, where species had greater relative fitness differences, but these associations were much weaker towards higher elevations. This appears to be because traits, particularly those associated with light competition, captured species' relative fitness differences at lower elevations, but not at the high elevation site, highlighting that the predictive ability of traits can depend on environmental context. The amplified relative fitness differences towards lower elevations suggest that climate warming may increase the likelihood of competitive exclusion. In addition, novel competitors tended to show greater niche overlap than current competitors, leading to stronger overall competitive effects. However, in general, trait differences predicted competitive outcomes of novel and current competitors similarly well, suggesting that traits can predict interactions between species that do not yet interact. Our study reinforces the importance of considering changing interactions for predicting species responses to climate change and provides experimental evidence supporting the usefulness of functional trait differences in forecasting the impacts of future plant interactions under changing climate.

Lags in the response of mountain plant communities to climate change.

Rapid climatic changes and increasing human influence at high elevations around the world will have profound impacts on mountain biodiversity. However, forecasts from statistical models (e.g. species distribution models) rarely consider that plant community changes could substantially lag behind climatic changes, hindering our ability to make temporally realistic projections for the coming century. Indeed, the magnitudes of lags, and the relative importance of the different factors giving rise to them, remain poorly understood. We review evidence for three types of lag: "dispersal lags" affecting plant species' spread along elevational gradients, "establishment lags" following their arrival in recipient communities, and "extinction lags" of resident species. Variation in lags is explained by variation among species in physiological and demographic responses, by effects of altered biotic interactions, and by aspects of the physical environment. Of these, altered biotic interactions could contribute substantially to establishment and extinction lags, yet impacts of biotic interactions on range dynamics are poorly understood. We develop a mechanistic community model to illustrate how species turnover in future communities might lag behind simple expectations based on species' range shifts with unlimited dispersal. The model shows a combined contribution of altered biotic interactions and dispersal lags to plant community turnover along an elevational gradient following climate warming. Our review and simulation support the view that accounting for disequilibrium range dynamics will be essential for realistic forecasts of patterns of biodiversity under climate change, with implications for the conservation of mountain species and the ecosystem functions they provide.

Global climate change increases risk of crop yield losses and food insecurity in the tropical Andes.

One of the greatest current challenges to human society is ensuring adequate food production and security for a rapidly growing population under changing climatic conditions. Climate change, and specifically rising temperatures, will alter the suitability of areas for specific crops and cultivation systems. In order to maintain yields, farmers may be forced to change cultivation practices, the timing of cultivation, or even the type of crops grown. Alternatively, farmers can change the location where crops are cultivated (e.g., to higher elevations) to track suitable climates (in which case the plants will have to grow in different soils), as cultivated plants will otherwise have to tolerate warmer temperatures and possibly face novel enemies. We simulated these two last possible scenarios (for temperature increases of 1.3°C and 2.6°C) in the Peruvian Andes through a field experiment in which several traditionally grown varieties of potato and maize were planted at different elevations (and thus temperatures) using either the local soil or soil translocated from higher elevations. Maize production declined by 21%-29% in response to new soil conditions. The production of maize and potatoes declined by >87% when plants were grown under warmer temperatures, mainly as a result of the greater incidence of novel pests. Crop quality and value also declined under simulated migration and warming scenarios. We estimated that local farmers may experience severe economic losses of up to 2,300 US$ ha-1  yr-1 . These findings reveal that climate change is a real and imminent threat to agriculture and that there is a pressing need to develop effective management strategies to reduce yield losses and prevent food insecurity. Importantly, such strategies should take into account the influences of non-climatic and/or biotic factors (e.g., novel pests) on plant development.

Metazoan parasites of Micropterus salmoides (Lacépède 1802) (Perciformes, Centrarchidae): a review with evidences of spillover and spillback.

Among the topics related to invasion science, the least studied are parasite co-introduction and spillback. This leads to an uncertainty in invasion ecology theories and applications to management. Therefore, the present study brings a systematic review of published information on the metazoan parasite fauna of Micropterus salmoides, a widely introduced fish, with the aim of comparing information about the composition and richness of the associated parasite communities in its native and introduced regions. This review demonstrates that there were twice as many studies of M. salmoides in its native region in comparison with introduced regions, although most of the studies focused on the analysis of a single species or taxon of parasite. This bias impacts the number of parasite species observed and, consequently, the apparent importance of enemy release in introduced regions. The composition of the parasite community in the two regions showed high similarity, which indicates the introduction and acquisition of parasites in introduced regions. Otherwise, there was no pattern related to the geographic distance, highlighting the influence of the propagule pressure and vector strength on the introduction of novel parasites. This illustrates the importance of vector strength on fish-parasite co-introduction and the necessity of new research examining host-parasite interactions with the parasite community of the invaded ecosystems. We still do not know the major influences of the composition of the parasite fauna of M. salmoides or how we can manage to develop a more restrictive vector pathway of introduction. The future of our ecosystems depends on how to account for current and future interactions among novel interactions, habitat, and climate change.

Taxonomic similarity, more than contact opportunity, explains novel plant-pathogen associations between native and alien taxa.

Novel associations between plants and pathogens can have serious impacts on managed and natural ecosystems world-wide. The introduction of alien plants increases the potential for biogeographically novel plant-pathogen associations to arise when pathogens are transmitted from native to alien plant species and vice versa. We quantified biogeographically novel associations recorded in New Zealand over the last 150 yr between plant pathogens (fungi, oomycetes and plasmodiophorids) and vascular plants. We examined the extent to which taxonomic similarity, pathogen traits, contact opportunity and sampling effort could explain the number of novel associates for host and pathogen species. Novel associations were common; approximately one-third of surveyed plants and pathogens were recorded with at least one biogeographically novel associate. Native plants had more alien pathogens than vice versa. Taxonomic similarity between the native and alien flora and the total number of recorded associations (a measure of sampling effort) best explained the number of novel associates among species. The frequency of novel associations and the importance of sampling effort as an explanatory variable emphasize the need for effective monitoring and risk assessment tools to mitigate the potential environmental and economic impact of novel pathogen associations.

Novel ecological interactions alter physiological responses of range-extending tropical and local temperate fishes under ocean warming.

Global warming facilitates species range-expansions, leading to novel biological interactions between local and range-expanding species. Little is still known of how such novel interactions modify the performance of interacting species or how these interactions might be altered under climate change. Here, we used an aquarium experiment to investigate the novel ecological interactions between a poleward range-extending coral reef damselfish ("tropical-vagrant") and a local temperate species ("temperate-local") collected from a climate warming hotspot in SE Australia. We measured the effect of novel interactions (isolated vs. paired fish species) on energy expenditure (activity levels, oxidative stress, and antioxidant responses), energy gain (feeding rates), and growth rates of both fish species under present-day (23 °C) and future ocean temperatures (26 °C). Short-term growth rates were faster in both species under novel interactions (paired species), regardless of elevated temperature. Compared to isolated species, activity level, feeding rate and oxidative stress level were also higher in the paired temperate fish but not in the paired tropical fish. The tropical fish showed an increased feeding rate and long-term growth under elevated temperature, irrespective of novel interactions. We conclude that novel ecological interactions under climate change can be an important driver of physiological traits in sympatric tropical and temperate fishes and can mediate critical physiological performance of fishes under ocean warming.

Identifying the generalizable controls on insect associations of native and non-native trees.

Trees growing outside their native geographic ranges often exhibit exceptional growth and survival due in part to the lack of co-evolved natural enemies that may limit their spread and suppress population growth. While most non-native trees tend to accumulate natural enemies over time, it remains uncertain which host and insect characteristics affect these novel associations and whether novel associations follow patterns of assembly similar to those of native hosts. Here, we used a dataset of insect-host tree associations in Europe to model which native insect species are paired with which native tree species, and then tested the model on its ability to predict which native insects are paired with which non-native trees. We show that native and non-native tree species closely related to known hosts are more likely to be hosts themselves, but that native host geographic range size, insect feeding guild, and sampling effort similarly affect insect associations. Our model had a strong ability to predict which insect species utilize non-native trees as hosts, but evolutionarily isolated tree species posed the greatest challenge to the model. These results demonstrate that insect-host associations can be reliably predicted, regardless of whether insect and host trees have co-evolved, and provide a framework for predicting future pest threats using a select number of easily attainable tree and insect characteristics.

Novel food resources and conservation of ecological interactions between the Andean Araucaria and the Austral parakeet.

In fragile ecosystems, the introduction of exotic species could alter some ecological processes. The Austral parakeet (Enicognathus ferrugineous) shows close ecological and evolutionary relationships with the Andean Araucaria (Araucaria araucana), so any alteration in these interactions may have negative consequences for both partners and for ecosystem functioning and structure. We conducted extensive roadside surveys to estimate the abundance of parakeets in the northern Patagonian Andes over 4 years and recorded the food plants consumed by foraging flocks. The use of native habitats and humanized areas like villages and farms was influenced by the Araucaria seed crop. In masting years, the large seed crop allowed a massive use of this resource during the non-breeding season, and even during the breeding season. The exploitation of exotic plants was minor in the masting year, but became predominant in non-masting years, especially during the non-breeding season. This feeding switch towards exotic plants primarily arose because the low Araucaria seed crop in non-masting years is entirely consumed just after production by domestic and wild exotic mammals living in Araucaria forests year-round, thus forcing the displacement of parakeets towards anthropic habitats to exploit exotic plants. Given the degradation of the remaining Andean Araucaria forests due to the impact of exotic mammals on the ecological interaction between Araucaria and Austral parakeets, ambitious programs to exclude or reduce the density of these alien mammals, including livestock, are warranted.

Do soil biota influence the outcome of novel interactions between plant competitors?

1. Species are shifting their ranges, for example to higher elevations, in response to climate change. Different plant species and soil microbiota will likely shift their ranges at different rates, giving rise to novel communities of plants and soil organisms. However, the ecological consequences of such novel plant-soil interactions are poorly understood. We experimentally simulated scenarios for novel interactions arising between high- and low elevation plants and soil biota following asynchronous climate change range shifts, asking to what extent the ability of plants to coexist depends on the origin of the soil biota. 2. In a greenhouse experiment, we grew pairs of low- (Poa trivialis and Plantago lanceolata) and high- (Poa alpina and Plantago alpina) elevation plant species alone and against a density gradient of con- or heterospecific neighbours. Plants grew on sterilized field soil that was inoculated with a soil community sampled from either low- or high elevation in the western Swiss Alps. We used the experiment to parameterize competition models, from which we predicted the population-level outcomes of competition in the presence of the different soil biota. 3. In the absence of neighbours, three of the four species produced more biomass with the low elevation soil biota. As a result of generally similar responses across plant species, soil biota tended not to affect plant interaction outcomes, with the low elevation species generally predicted to competitively exclude high elevation species irrespective of the soil biota origin. However, the low elevation grass Poa trivialis was only able to invade communities of Poa alpina in the presence of a low elevation soil biota. This suggests that, at least in some cases, the outcome of novel competitive interactions between plants following climate change will depend on whether shifts in the distribution of plant and soil organisms are asynchronous. 4. Synthesis. Our results indicate that the changing soil communities that plants encounter during range expansion can influence plant performance. However, this is only likely to alter expectations for the ability of plants to coexist following climate change if plant species respond differently to the change in the soil community.

Nematode community responses to range-expanding and native plant communities in original and new range soils.

Many plant species expand their range to higher latitudes in response to climate change. However, it is poorly understood how biotic interactions in the new range differ from interactions in the original range. Here, in a mesocosm experiment, we analyze nematode community responses in original and new range soils to plant communities with either (a) species native in both the original and new range, (b) range-expanding species related to these natives (related range expanders), or (c) range expanders without native congeneric species in the new range (unrelated range expanders). We hypothesized that nematode community shifts between ranges are strongest for unrelated range expanders and minimal for plant species that are native in both ranges. As a part of these community shifts, we hypothesized that range expanders, but not natives, would accumulate fewer root-feeding nematodes in their new range compared to their original range. Analyses of responses of nematodes from both original and new ranges and comparison between range expanders with and without close relatives have not been made before. Our study reveals that none of the plant communities experienced evident nematode community shifts between the original and new range. However, in soils from the new range, root-feeding nematode communities of natives and related range expanders were more similar than in soils from the original range, whereas the nematode community of unrelated range expanders was distinct from the communities of natives and related range expanders in soils from both ranges. The abundances of root-feeding nematodes were comparable between the original and new range for all plant communities. Unexpectedly, unrelated range expanders overall accumulated most root-feeding nematodes, whereas related range expanders accumulated fewest. We conclude that nematode communities associated with native and range-expanding plant species differ between the original and the new range, but that range-expanding plant species do not accumulate fewer root-feeding nematodes in their new than in their original range.

Reunion overseas: introduced wild boars and cultivated orange trees interact in the Brazilian Atlantic Forest

Introduction: Little is known concerning novel interactions between species that typically interact in their native range but, as a consequence of human activity, are also interacting out of their original distribution under new ecological conditions. Objective: We investigate the interaction between the orange tree and wild boar, both of which share Asian origins and have been introduced to the Americas (i.e. the overseas). Methods: Specifically, we assessed whether i) wild boars consume orange (Citrus sinensis) fruits and seeds in orchards adjacent to a remnant of the Atlantic Forest of Brazil, ii) the orange seeds are viable after passing through boar's digestive tract and iii) whether the orange tree may naturalise in the forest remnant assisted by wild boars. Results: Our camera surveys indicated that wild boar was by far the most frequent consumer of orange fruits (40.5 % of camera trap-days). A considerable proportion of sown orange seeds extracted from fresh boar feces emerged seedlings (27.8 %, N = 386) under controlled greenhouse conditions. Further, 37.6 % of sown seeds (N = 500) in the forest remnant emerged seedlings in July 2015; however, after ~4 years (March 2019) only 9 seedlings survived (i.e. 4.8 %, N = 188). Finally, 52 sweet orange seedlings were found during surveys within the forest remnant which is intensively used by wild boars. This study indicates a high potential of boars to act as effective seed dispersers of the sweet orange. However, harsh competition with native vegetation and the incidence of lethal diseases, which quickly kill sweet orange trees under non-agricultural conditions, could seriously limit orange tree establishment in the forest. Conclusions: Our results have important implications not only because the wild boar could be a vector of potential invasive species, but also because they disperse seeds of some native species (e.g. the queen palm, Syagrus romanzofiana) in defaunated forests, where large native seed dispersers are missing; thus, wild boars could exert critical ecological functions lost due to human activity.

Introducción: Se conoce relativamente poco sobre las llamadas 'interacciones noveles' entre especies que típicamente interactúan en su área de distribución nativa pero que, como consecuencia de la actividad humana, también interactúan fuera de su distribución original bajo nuevas condiciones ecológicas. Objetivo: Investigamos la interacción entre el naranjo y el jabalí, ambos con origen asiático e introducidos en las Américas (es decir, del extranjero). Métodos: Específicamente, evaluamos si i) los jabalíes consumen frutas y semillas del naranjo (Citrus sinensis) en naranjales adyacentes a un parche remanente del bosque atlántico de Brasil, ii) las semillas de naranja son viables tras pasar por el tracto digestivo del jabalí, y iii) si el naranjo puede llegar a naturalizarse en el parche de bosque gracias a los jabalíes. Resultados: Los resultados de nuestro fototrampeo indicaron que el jabalí fue, con mucho, el consumidor más frecuente de las naranjas (40.5 % cámaras trampa-días). Una proporción considerable de semillas de naranjo extraídas de heces de jabalí frescas y sembradas emergieron plántulas bajo condiciones de invernadero controladas (27.8 %, N = 386). Además, del 37.6 % de las semillas sembradas (N = 500) en el parche remanente de bosque emergieron plántulas en julio 2015; sin embargo, después de ~ 4 años (marzo 2019) solo sobrevivieron 9 plántulas (es decir, 4.8 %, N = 188). Finalmente, se encontraron 52 plántulas de naranja dulce durante varias prospecciones dentro del parche de bosque que es utilizado intensivamente por los jabalíes. Este estudio indica un alto potencial de los jabalíes para actuar como dispersores de semillas eficaces del naranjo dulce. Sin embargo, la severa competencia con la abundante vegetación nativa y la incidencia de enfermedades letales, que matan rápidamente los naranjos dulces en condiciones no agrícolas, podrían limitar seriamente el establecimiento de naranjos en el bosque. Conclusiones: Nuestros resultados tienen implicaciones importantes no solo porque el jabalí podría ser un vector de posibles especies de plantas invasoras, sino también porque dispersan semillas de algunas especies nativas (p.e., la palmera reina, Syagrus romanzofiana) en estos bosques defaunados, donde faltan dispersores nativos de semillas de gran tamaño. Por ello, los jabalíes podrían ejercer funciones ecológicas críticas que se han perdido debido a la actividad humana.