w ww ww w. .f fr ro on nt ti ie er rs si in ne ec co ol lo og gy y. .o or rg g R Re eb be ec cc ca a S S E Ep pa an nc ch hi in n--N Ni ie el ll l 1 1* * , , M Ma at tt th he ew w B B H Hu uf ff fo or rd d 2 2 , , C Cl la ar re e E E A As sl la an n 3 3 , , J Ja as so on n P P S Se ex xt to on n 2 2 , , J Je ef ff fr re ey y D D P Po or rt t 4 4 , , a an nd d T Ti im mo ot th hy y M M W Wa ar ri in ng g 5 5Control of biological invasions depends on the collective decisions of resource managers across invasion zones. Regions with high land-use diversity, which we refer to as "management mosaics", may be subject to severe invasions, for two main reasons. First, as land becomes increasingly subdivided, each manager assumes responsibility for a smaller portion of the total damages imposed by invasive species; the incentive to control invasives is therefore diminished. Secondly, managers opting not to control the invasion increase control costs for neighboring land managers by allowing their lands to act as an invader propagule source. Coordination among managers can help mitigate these effects, but greater numbers -and a wider varietyof land managers occupying a region hinder collective action. Here, we discuss the challenges posed by management mosaics, using a case study of the yellow starthistle (Centaurea solstitialis) invasion in the Sierra Nevada foothills of California. We suggest that the incorporation of management mosaic dynamics into invasive species research and management is essential for successful control of invasions, and provide recommendations to address this need.

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BackgroundAs global environmental change accelerates, biodiversity losses can disrupt interspecific interactions. Extinctions of mutualist partners can create “widow” species, which may face reduced ecological fitness. Hypothetically, such mutualism disruptions could have cascading effects on biodiversity by causing additional species coextinctions. However, the scope of this problem – the magnitude of biodiversity that may lose mutualist partners and the consequences of these losses – remains unknown.Methodology/Principal FindingsWe conducted a systematic review and synthesis of data from a broad range of sources to estimate the threat posed by vertebrate extinctions to the global biodiversity of vertebrate-dispersed and -pollinated plants. Though enormous research gaps persist, our analysis identified Africa, Asia, the Caribbean, and global oceanic islands as geographic regions at particular risk of disruption of these mutualisms; within these regions, percentages of plant species likely affected range from 2.1–4.5%. Widowed plants are likely to experience reproductive declines of 40–58%, potentially threatening their persistence in the context of other global change stresses.ConclusionsOur systematic approach demonstrates that thousands of species may be impacted by disruption in one class of mutualisms, but extinctions will likely disrupt other mutualisms, as well. Although uncertainty is high, there is evidence that mutualism disruption directly threatens significant biodiversity in some geographic regions. Conservation measures with explicit focus on mutualistic functions could be necessary to bolster populations of widowed species and maintain ecosystem functions.

There is growing realization that intraspecific variation in seed dispersal can have important ecological and evolutionary consequences. However, we do not have a good understanding of the drivers or causes of intraspecific variation in dispersal, how strong an effect these drivers have, and how widespread they are across dispersal modes. As a first step to developing a better understanding, we present a broad, but not exhaustive, review of what is known about the drivers of intraspecific variation in seed dispersal, and what remains uncertain. We start by decomposing ‘drivers of intraspecific variation in seed dispersal’ into intrinsic drivers (i.e. variation in traits of individual plants) and extrinsic drivers (i.e. variation in ecological context). For intrinsic traits, we further decompose intraspecific variation into variation among individuals and variation of trait values within individuals. We then review our understanding of the major intrinsic and extrinsic drivers of intraspecific variation in seed dispersal, with an emphasis on variation among individuals. Crop size is the best-supported and best-understood intrinsic driver of variation across dispersal modes; overall, more seeds are dispersed as more seeds are produced, even in cases where per seed dispersal rates decline. Fruit/seed size is the second most widely studied intrinsic driver, and is also relevant to a broad range of seed dispersal modes. Remaining intrinsic drivers are poorly understood, and range from effects that are probably widespread, such as plant height, to drivers that are most likely sporadic, such as fruit or seed colour polymorphism. Primary extrinsic drivers of variation in seed dispersal include local environmental conditions and habitat structure. Finally, we present a selection of outstanding questions as a starting point to advance our understanding of individual variation in seed dispersal.

Summary 1. Invasion biologists use two main approaches to evaluate the effects of non‐native species (NNS) on diversity of native species (DNS), namely space‐for‐time and time approaches. These approaches have pitfalls related to lack of controls: the former lacks pre‐invasion data, while the latter often lacks data from non‐invaded sites. 2. We propose a framework that combines space‐for‐time and time approaches and which should result in more focused mechanistic hypotheses and experiments to test the causes of invasibility and the effects of NNS on DNS. We illustrate the usefulness of our framework using two case studies: one with the submersed macrophyte, Hydrilla verticillata, in reservoir and the other with the fish, Geophagus proximus, in a large river–floodplain system. 3. Hydrilla verticillata invaded sites with DNS similar to that found in non‐invaded sites, indicating that biotic and/or abiotic factors did not influence invasion success; however, DNS increased over time in invaded sites compared with non‐invaded sites, suggesting that H. verticillata facilitated natives. In contrast, G. proximus invaded sites with higher DNS than non‐invaded sites, suggesting that biotic and/or abiotic factors favouring natives were important for invasion success, but DNS increased in invaded and non‐invaded sites over time, indicating that an independent factor contributed to DNS increases. 4. Conclusions from both studies would have been inaccurate or incomplete if the space‐for‐time and time approaches had not been used in combination as proposed in our framework.

Native plant species that have lost their mutualist partners may require non-native pollinators or seed dispersers to maintain reproduction. When natives are highly specialized, however, it appears doubtful that introduced generalists will partner effectively with them. We used visitation observations and pollination treatments (experimental manipulations of pollen transfer) to examine relationships between the introduced, generalist Japanese White-eye (Zosterops japonicus) and 3 endemic Hawaiian plant species (Clermontia parviflora, C. montis-loa, and C. hawaiiensis). These plants are characterized by curved, tubular flowers, apparently adapted for pollination by curve-billed Hawaiian honeycreepers. Z. japonicus were responsible for over 80% of visits to flowers of the small-flowered C. parviflora and the midsize-flowered C. montis-loa. Z. japonicus-visited flowers set significantly more seed than did bagged flowers. Z. japonicus also demonstrated the potential to act as an occasional Clermontia seed disperser, although ground-based frugivory by non-native mammals likely dominates seed dispersal. The large-flowered C. hawaiiensis received no visitation by any birds during observations. Unmanipulated and bagged C. hawaiiensis flowers set similar numbers of seeds. Direct examination of Z. japonicus and Clermontia morphologies suggests a mismatch between Z. japonicus bill morphology and C. hawaiiensis flower morphology. In combination, our results suggest that Z. japonicus has established an effective pollination relationship with C. parviflora and C. montis-loa and that the large flowers of C. hawaiiensis preclude effective visitation by Z. japonicus.

I examined the role of bird dispersal in invasiveness of three non-native plant species in California, USA: Triadica sebifera, Ligustrum lucidum, and Olea europaea. I selected these species because their invasiveness in California is uncertain, but a survey of ornithologists highlighted them as likely bird-dispersed. I quantified bird frugivory of these plants, compared them with a native species (Heteromeles arbutifolia), and explored the management implications of dispersal mutualisms for these and other incipient invasive plants. Fruit removal by birds was sufficient to permit spread for all study species. Seed dispersers (rather than seed predators) and pulse feeders (flocking species with potential for long distance dispersal) performed most fruit removal for the non-native species, a pattern indicative of an effective dispersal regime. The number of fruiting trees per stand was a significant predictor of bird visitation. Founding population size may thus be important in management of invasive, bird-dispersed plants. Disperser-defined niches were relatively narrow because a few disperser species performed the majority of fruit removal from study trees, but each fruit species was consumed by a variety of potential dispersers. This results in strong pairwise niche overlap between some plant species. Ordinated by bird use, study site-species combinations clustered more by geographic location than by plant species, emphasizing the opportunistic nature of bird foraging. None of the non-native focal plant species appears dispersal limited, and all have formed novel mutualisms in California. It is possible that these plants are now in lag phases preceding bird-mediated invasion. Consideration of bird dispersal when evaluating invasiveness is therefore an imperative.

Methods to detect and monitor the spread of invasive grasses are critical to avoid ecosystem transformations and large economic costs. The rapid spread of non‐native buffelgrass(Pennisetum ciliare) has intensified fire risk and is replacing fire intolerant native vegetation in the Sonoran Desert of the southwestern US. Coarse‐resolution satellite imagery has had limited success in detecting small patches of buffelgrass, whereas ground‐based and aerial survey methods are often cost prohibitive. To improve detection, we trained 2 m resolution DigitalGlobe WorldView‐2 satellite imagery with 12 cm resolution unmanned aerial vehicle (UAV) imagery and classified buffelgrass on Google Earth Engine, a cloud computing platform, using Random Forest (RF) models in Saguaro National Park, Arizona, USA. Our classification models had an average overall accuracy of 93% and producer's accuracies of 94–96% for buffelgrass, although user's accuracies were low. We detected a 2.92 km2 area of buffelgrass in the eastern Rincon Mountain District (1.07% of the total area) and a 0.46 km2 area (0.46% of the total area) in the western Tucson Mountain District of Saguaro National Park. Buffelgrass cover was significantly greater in the Sonoran Paloverde‐Mixed Cacti Desert Scrub vegetation type, on poorly developed Entisols and Inceptisol soils and on south‐facing topographic aspects compared to other areas. Our results demonstrate that high‐resolution imagery improve on previous attempts to detect and classify buffelgrass and indicate potential areas where the invasive grass might spread. The methods demonstrated in this study could be employed by land managers as a low‐cost strategy to identify priority areas for control efforts and continued monitoring.