Plant networks are more connected by invasive brome and native shrub facilitation in Central California drylands.

Dominant vegetation in many ecosystems is an integral component of structure and habitat. In many drylands, native shrubs function as foundation species that benefit other plants and animals. However, invasive exotic plant species can comprise a significant proportion of the vegetation. In Central California drylands, the facilitative shrub Ephedra californica and the invasive Bromus rubens are widely dispersed and common. Using comprehensive survey data structured by shrub and open gaps for the region, we compared network structure with and without this native shrub canopy and with and without the invasive brome. The presence of the invasive brome profoundly shifted the network measure of centrality in the microsites structured by a shrub canopy (centrality scores increased from 4.3 under shrubs without brome to 6.3, i.e. a relative increase of 42%). This strongly suggests that plant species such as brome can undermine the positive and stabilizing effects of native foundation plant species provided by shrubs in drylands by changing the frequency that the remaining species connect to one another. The net proportion of positive and negative associations was consistent across all microsites (approximately 50% with a total of 14% non-random co-occurrences on average) suggesting that these plant-plant networks are rewired but not more negative. Maintaining resilience in biodiversity thus needs to capitalize on protecting native shrubs whilst also controlling invasive grass species particularly when associated with shrubs.

Energetic costs of raising brood in honey bee colonies are high, but heater bees are cheap.

Little is known about the energetic costs to insects of raising young. Honey bees collectively raise young, or brood, through a series of complex behaviors that appear to accelerate and synchronize the timing of brood maturation. These include maintaining the brood nest at warmer and consistent temperatures (33-36 °C) and the exceptional activity of heater bees. Heater bees are a part of the larger group of nurse bees that care for brood by rapidly contracting thoracic muscles to generate high body temperatures, from 42 to 47 °C. Heater bees move among brood cells and display this behavior to regulate the temperatures of individual larvae and pupae. We constructed three sets of experimental hives to explore the energy costs of raising brood in general and the cost of heater bees specifically. One set was designed to estimate the numerical allocation of individuals to the heater bee task. The second set was designed to contain only brood, which eliminated foraging and allowed us to quantify stored honey use when rearing juveniles at 10 and 30 °C. The final set was used to measure the respiration rates and energy expenditure of individual bees displaying resting, walking, heating, and agitated behavior. By integrating honey used by brood-only experimental colonies with whole-colony measurements of honey storage in the literature, we estimated that raising brood costs colonies half of their annual energy budgets stored as honey, or approximately 43.7 ± 0.9 kg·yr-1. We estimated that roughly 2 % of individuals in a colony perform as heater bees. Respiration rates of heater bees (19 mW) were more than those of resting bees (8 mW) but similar to those of walking bees (20 mW) and about half of those that were agitated (46 mW). The energetic cost of heating was more than an order of magnitude lower than the reported values for the energetic cost of flying. By integrating data from our three experimental hives, we estimate that the annual cost of raising brood is relatively high. However, heater bee behavior and physiology may require only about 7 % of the annual honey stored by a colony.

A trait-based framework linking the soil metabolome to plant-soil feedbacks.

By modifying the biotic and abiotic properties of the soil, plants create soil legacies that can affect vegetation dynamics through plant-soil feedbacks (PSF). PSF are generally attributed to reciprocal effects of plants and soil biota, but these interactions can also drive changes in the identity, diversity and abundance of soil metabolites, leading to more or less persistent soil chemical legacies whose role in mediating PSF has rarely been considered. These chemical legacies may interact with microbial or nutrient legacies to affect species coexistence. Given the ecological importance of chemical interactions between plants and other organisms, a better understanding of soil chemical legacies is needed in community ecology. In this Viewpoint, we aim to: highlight the importance of belowground chemical interactions for PSF; define and integrate soil chemical legacies into PSF research by clarifying how the soil metabolome can contribute to PSF; discuss how functional traits can help predict these plant-soil interactions; propose an experimental approach to quantify plant responses to the soil solution metabolome; and describe a testable framework relying on root economics and seed dispersal traits to predict how plant species affect the soil metabolome and how they could respond to soil chemical legacies.

Trade-offs in non-native plant herbivore defences enhance performance.

Non-native plants are typically released from specialist enemies but continue to be attacked by generalists, albeit at lower intensities. This reduced herbivory may lead to less investment in constitutive defences and greater investment in induced defences, potentially reducing defence costs. We compared herbivory on 27 non-native and 59 native species in the field and conducted bioassays and chemical analyses on 12 pairs of non-native and native congeners. Non-natives suffered less damage and had weaker constitutive defences, but stronger induced defences than natives. For non-natives, the strength of constitutive defences was correlated with the intensity of herbivory experienced, whereas induced defences showed the reverse. Investment in induced defences correlated positively with growth, suggesting a novel mechanism for the evolution of increased competitive ability. To our knowledge, these are the first linkages reported among trade-offs in plant defences related to the intensity of herbivory, allocation to constitutive versus induced defences, and growth.

Rapid evolutionary trade-offs between resistance to herbivory and tolerance to abiotic stress in an invasive plant.

Release from enemies can lead to rapid evolution in invasive plants, including reduced metabolic investment in defence. Conversely, reassociation with enemies leads to renewed evolution of defence, but the potential costs of this evolution are poorly documented. We report increased resistance of the invader Ambrosia artemisiifolia after reassociation with a coevolved specialist herbivore, and that this increase corresponds with reduced abiotic stress tolerance. Herbivore resistance was higher, but drought tolerance was lower in plants from populations with a longer reassociation history, and this corresponded with changes in phenylpropanoids involved in insect resistance and abiotic stress tolerance. These changes were corroborated by shifts in the expression of underlying biosynthetic genes and plant anti-oxidants. Together, our findings suggest rapid evolution of plant traits after reassociation with coevolved enemies, resulting in genetically based shifts in investment between abiotic and biotic stress responses, providing insights into co-evolution, plant invasion and biological control.

Age, experience, social goals, and engagement with research scientists may promote innovation in ecological restoration.

Innovation in ecological restoration is necessary to achieve the ambitious targets established in United Nations conventions and other global restoration initiatives. Innovation is also crucial for navigating uncertainties in repairing and restoring ecosystems, and thus practitioners often develop innovations at project design and implementation stages. However, innovation in ecological restoration can be hindered by many factors (e.g., time and budget constraints, and project complexity). Theory and research on innovation has been formally applied in many fields, yet explicit study of innovation in ecological restoration remains nascent. To assess the use of innovation in restoration projects, including its drivers and inhibitors, we conducted a social survey of restoration practitioners in the United States. Specifically, we assessed relationships between project-based innovation and traits of the individual practitioner (including, for example, age, gender, experience); company (including, for example, company size and company's inclusion of social goals); project (including, for example, complexity and uncertainty); and project outcomes (such as completing the project on time/on budget and personal satisfaction with the work). We found positive relationships between project-based innovation and practitioner traits (age, gender, experience, engagement with research scientists), one company trait (company's inclusion of social goals in their portfolio), and project traits (project complexity and length). In contrast, two practitioner traits, risk aversion and the use of industry-specific information, were negatively related to project-based innovation. Satisfaction with project outcomes was positively correlated with project-based innovation. Collectively, the results provide insights into the drivers and inhibitors of innovation in restoration and suggest opportunities for research and application.

Competition for water and rapid exclusion of an island endemic by a pantropical species in a tropical climate.

Water availability has major effects on community structure and dynamics globally, yet our understanding of competition for water in the tropics is limited. On the tropical Trindade Island, we explored competition for water in the context of the rapid exclusion of an endemic sedge, Cyperus atlanticus (Cyperaceae), by a pantropical, N-fixing shrub, Guilandina bonduc (Fabaceae). Guilandina patches were generally surrounded by rings of bare soil, and dead Cyperus halos commonly surrounded these bare zones. With geo-referenced measurements, we showed that Guilandina patches and bare soil zones rapidly expanded and replaced adjacent Cyperus populations. We found that soil water potentials were much lower in bare soils than soils under Guilandina or Cyperus, and that leaf water potentials of Cyperus plants were lower when co-occurring with Guilandina than when alone. When Guilandina was removed experimentally, Cyperus populations expanded and largely covered the bare soil zones. Our results indicate that when Guilandina establishes, its root systems expand beyond its canopies and these roots pull water from soils beneath Cyperus and kill it, creating bare zone halos, and then Guilandina expands and repeats the process. This scenario indicates rapid competitive exclusion and displacement of an endemic by a common pantropical species, at least in part through competition for water.

A realized facilitation cascade mediated by biological soil crusts in a sagebrush steppe community.

Biological soil crusts can have strong effects on vascular plant communities which have been inferred from short-term germination and early establishment responses. However, biocrusts are often assumed to function as an "organizing principle" in communities because their effects can "cascade" to interactions among crust-associated plant species. We conducted surveys and experiments to explore these cascades and found that biocrusts were positively associated with large patches (> 10 m diameter) of a dominant shrub Artemisia tridentata. At the smaller scale of individual shrubs and the open matrices between shrubs, biocrusts were negatively associated with Artemisia. Juveniles of Artemisia were found only in biocrusts in intershrub spaces and never under shrubs or in soil without biocrusts. In two-year field experiments, biocrusts increased the growth of Festuca and the photosynthetic rates of Artemisia. Festuca planted under Artemisia were also at least twice as large as those planted in open sites without crusts or where Artemisia were removed. Thus, biocrusts can facilitate vascular plants over long time periods and can contribute to a "realized" cascade with nested negative and positive interactions for a range of species, but unusual among documented cascades in that it includes only autotrophs.

Evidence for scale-dependent root-antation feedback and its role in halting the spread of a pantropical shrub into an endemic sedge.

Vegetation pattern formation is a widespread phenomenon in resource-limited environments, but the driving mechanisms are largely unconfirmed empirically. Combining results of field studies and mathematical modeling, empirical evidence for a generic pattern-formation mechanism is demonstrated with the clonal shrub Guilandina bonduc L. (hereafter Guilandina) on the Brazilian island of Trindade. The mechanism is associated with water conduction by laterally spread roots and root augmentation as the shoot grows-a crucial element in the positive feedback loop that drives spatial patterning. Assuming precipitation-dependent root-shoot relations, the model accounts for the major vegetation landscapes on Trindade Island, substantiating lateral root augmentation as the driving mechanism of Guilandina patterning. Guilandina expands into surrounding communities dominated by the Trindade endemic, Cyperus atlanticus Hemsl. (hereafter Cyperus). It appears to do so by decreasing the water potential in soils below Cyperus through its dense lateral roots, leaving behind a patchy Guilandina-only landscape. We use this system to highlight a novel form of invasion, likely to apply to many other systems where the invasive species is pattern-forming. Depending on the level of water stress, the invasion can take two distinct forms: (i) a complete invasion at low stress that culminates in a patchy Guilandina-only landscape through a spot-replication process, and (ii) an incomplete invasion at high stress that begins but does not spread, forming isolated Guilandina spots of fixed size, surrounded by bare-soil halos, in an otherwise uniform Cyperus grassland. Thus, drier climates may act selectively on pattern-forming invasive species, imposing incomplete invasion and reducing the negative effects on native species.

Plant litter strengthens positive biodiversity-ecosystem functioning relationships over time.

Plant biodiversity-productivity relationships become stronger over time in grasslands, forests, and agroecosystems. Plant shoot and root litter is important in mediating these positive relationships, yet the functional role of plant litter remains overlooked in long-term experiments. We propose that plant litter strengthens biodiversity-ecosystem functioning relationships over time in four ways by providing decomposing detritus that releases nitrogen (N) over time for uptake by existing and succeeding plants, enhancing overall soil fertility, changing soil community composition, and reducing the impact of residue-borne pathogens and pests. We bring new insights into how diversity-productivity relationships may change over time and suggest that the diversification of crop residue retention through increased residue diversity from plant mixtures will improve the sustainability of food production systems.

Temporal stability of productivity is associated with complementarity and competitive intensities in intercropping.

Year-to-year stability in crop production is a crucial aspect of feeding a growing global population. Evidence from natural ecosystems shows that increasing plant diversity generally increases the temporal stability of productivity; however, we have little knowledge of the mechanisms by which diversity affects stability. In fact, understanding the drivers of stability is a major knowledge gap in our understanding of biodiversity and ecosystem function in general. We varied resource inputs into crop monocultures and intercropping of maize/pea and maize/rapeseed for 3 years in field experiments to create a wide range of values for temporal stability, complementarity effects, selection effects, competition, and facilitation. We correlated whole-system temporal stability in productivity with these values and the stability of competitively subordinate species and competitively dominant species in the intercrops. We then used structural equation modeling (SEM), which combines complex path models with latent variables, to estimate how interspecific interactions for water, nitrogen, and phosphorus affected the relationships between stability and these values. Intercropping treatments did not increase stability, but the wide range of stability created by our experiments allowed us to explore the relationship of many factors with stability. Complementarity correlated positively with the temporal stability of grain yield and aboveground biomass, suggesting that either facilitative interactions or niche partitioning shifted over time in ways that promoted stability. Furthermore, the temporal stability of total productivity of intercropping relied most on the stability of more productive species. However, facilitation tested by relative interaction index independently did not correlate with stability, but the temporal stability of the whole system increased as the competitive effects of competitively dominant species (pea and rapeseed) on competitively subordinate species (maize) decreased and was highest when these competitive effects were virtually zero. SEM indicated that as competition for soil nitrogen from competitively dominant species on competitively subordinate species decreased, the overall temporal stability of whole-system aboveground biomass increased. This stability then led to greater stability in grain production. Our findings indicate that complex shifts in complementarity and competitive intensities are likely to be key mechanisms that maintain temporal stability in species-diverse agriculture and, potentially, in natural systems.

The EICA is dead? Long live the EICA!

An important hypothesis for how plants respond to introduction to new ranges is the evolution of increased competitive ability (EICA). EICA predicts that biogeographical release from natural enemies initiates a trade-off in which exotic species in non-native ranges become larger and more competitive, but invest less in consumer defences, relative to populations in native ranges. This trade-off is exceptionally complex because detecting concomitant biogeographical shifts in competitive ability and consumer defence depends upon which traits are targeted, how competition is measured, the defence chemicals quantified, whether defence chemicals do more than defend, whether 'herbivory' is artificial or natural, and where consumers fall on the generalist-specialist spectrum. Previous meta-analyses have successfully identified patterns but have yet to fully disentangle this complexity. We used meta-analysis to reevaluate traditional metrics used to test EICA theory and then expanded on these metrics by partitioning competitive effect and competitive tolerance measures and testing Leaf-Specific Mass in detail as a response trait. Unlike previous syntheses, our meta-analyses detected evidence consistent with the classic trade-off inherent to EICA. Plants from non-native ranges imposed greater competitive effects than plants from native ranges and were less quantitatively defended than plants from native ranges. Our results for defence were not based on complex leaf chemistry, but instead were estimated from tannins, toughness traits and primarily Leaf-Specific Mass. Species specificity occurred but did not influence the general patterns. As for all evidence for EICA-like trade-offs, we do not know if the biogeographical differences we found were caused by trade-offs per se, but they are consistent with predictions derived from the overarching hypothesis. Underestimating physical leaf structure may have contributed to two decades of tepid perspectives on the trade-offs fundamental to EICA.

Acquisition and evolution of enhanced mutualism-an underappreciated mechanism for invasive success?

Soil biota can determine plant invasiveness, yet biogeographical comparisons of microbial community composition and function across ranges are rare. We compared interactions between Conyza canadensis, a global plant invader, and arbuscular mycorrhizal (AM) fungi in 17 plant populations in each native and non-native range spanning similar climate and soil fertility gradients. We then grew seedlings in the greenhouse inoculated with AM fungi from the native range. In the field, Conyza plants were larger, more fecund, and associated with a richer community of more closely related AM fungal taxa in the non-native range. Fungal taxa that were more abundant in the non-native range also correlated positively with plant biomass, whereas taxa that were more abundant in the native range appeared parasitic. These patterns persisted when populations from both ranges were grown together in a greenhouse; non-native populations cultured a richer and more diverse AM fungal community and selected AM fungi that appeared to be more mutualistic. Our results provide experimental support for evolution toward enhanced mutualism in non-native ranges. Such novel relationships and the rapid evolution of mutualisms may contribute to the disproportionate abundance and impact of some non-native plant species.

Belowground feedbacks as drivers of spatial self-organization and community assembly.

Vegetation patterning in water-limited and other resource-limited ecosystems highlights spatial self-organization processes as potentially key drivers of community assembly. These processes provide insight into predictable landscape-level relationships between organisms and their abiotic environment in the form of regular and irregular patterns of biota and resources. However, two aspects have largely been overlooked; the roles played by plant - soil-biota feedbacks and allelopathy in spatial self-organization, and their potential contribution, along with plant-resource feedbacks, to community assembly through spatial self-organization. Here, we expand the drivers of spatial self-organization from a focus on plant-resource feedbacks to include plant - soil-biota feedbacks and allelopathy, and integrate concepts of nonlinear physics and community ecology to generate a new hypothesis. According to this hypothesis, below-ground processes can affect community assemblages through two types of spatial self-organization, global and local. The former occurs simultaneously across whole ecosystems, leading to self-organized patterns of biota, allelochemicals and resources, and niche partitioning. The latter occurs locally in ecotones, and determines ecotone structure and motion, invasion dynamics, and species coexistence. Studies of the two forms of spatial self-organization are important for understanding the organization of plant communities in drier climates which are likely to involve spatial patterning or re-patterning. Such studies are also important for developing new practices of ecosystem management, based on local manipulations at ecotones, to slow invasion dynamics or induce transitions from transitive to intransitive networks of interspecific interactions which increase species diversity.

Belowground facilitation and trait matching: two or three to tango?

High biodiversity increases ecosystem functions; however, belowground facilitation remains poorly understood in this context. Here, we explore mechanisms that operate via 'giving-receiving feedbacks' for belowground facilitation. These include direct effects via root exudates, signals, and root trait plasticity, and indirect biotic facilitation via the effects of root exudates on soil biota and feedback from biota to plants. We then highlight that these two- or three-way mechanisms must affect biodiversity-ecosystem function relationships via specific combinations of matching traits. To tango requires a powerful affinity and harmony between well-matched partners, and such matches link belowground facilitation to the effect of biodiversity on function. Such matching underpins applications in intercropping, forestry, and pasture systems, in which diversity contributes to greater productivity and sustainability.

Plasticity in response to plant-plant interactions and water availability.

The plastic responses of plants to abiotic and biotic environmental factors have generally been addressed separately; thus we have a poor understanding of how these factors interact. For example, little is known about the effects of plant-plant interactions on the plasticity of plants in response to water availability. Furthermore, few studies have compared the effects of intra- and interspecific interactions on plastic responses to abiotic factors. To explore the effects of intraspecific and interspecific plant-plant interactions on plant responses to water availability, we grew Leucanthemum vulgare and Potentilla recta with a conspecific or the other species, and grew pairs of each species as controls in pots with the roots, but not shoots, physically separated. We subjected these competitive arrangements to mesic and dry conditions, and then measured shoot mass, root mass, total mass and root : shoot ratio and calculated plasticity in these traits. The total biomass of both species was highly suppressed by both intra- and interspecific interactions in mesic soil conditions. However, in drier soil, intraspecific interactions for both species and the effect of P. recta on L. vulgare were facilitative. For plasticity in response to water supply, when adjusted for total biomass, drought increased shoot mass, and decreased root mass and root : shoot ratios for both species in intraspecific interactions. When grown alone, there were no plastic responses in any trait except total mass, for either species. Our results suggested that plants interacting with other plants often show improved tolerance for drought than those grown alone, perhaps because of neighbor-induced shifts in plasticity in biomass allocation. Facilitative effects might also be promoted by plasticity to drought in root : shoot ratios.

Network motifs involving both competition and facilitation predict biodiversity in alpine plant communities.

Biological diversity depends on multiple, cooccurring ecological interactions. However, most studies focus on one interaction type at a time, leaving community ecologists unsure of how positive and negative associations among species combine to influence biodiversity patterns. Using surveys of plant populations in alpine communities worldwide, we explore patterns of positive and negative associations among triads of species (modules) and their relationship to local biodiversity. Three modules, each incorporating both positive and negative associations, were overrepresented, thus acting as "network motifs." Furthermore, the overrepresentation of these network motifs is positively linked to species diversity globally. A theoretical model illustrates that these network motifs, based on competition between facilitated species or facilitation between inferior competitors, increase local persistence. Our findings suggest that the interplay of competition and facilitation is crucial for maintaining biodiversity.