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Weeds pose severe threats to agricultural and natural landscapes worldwide. One major reason for the failure to effectively manage weeds at landscape scales is that current Best Management Practice guidelines, and research on how to improve such guidelines, focus too narrowly on property-level management decisions. Insufficiently considered are the aggregate effects of individual actions to determine landscape-scale outcomes, or whether there are collective practices that would improve weed management outcomes. Here, we frame landscape-scale weed management as a social dilemma, where trade-offs occur between individual and collective interests. We apply a transdisciplinary system approach-integrating the perspectives of ecologists, evolutionary biologists and agronomists into a social science theory of social dilemmas-to four landscape-scale weed management challenges: achieving plant biosecurity, preventing weed seed contamination, maintaining herbicide susceptibility, and sustainably using biological control. We describe how these four challenges exhibit characteristics of "public good problems," wherein effective weed management requires the active contributions of multiple actors, while benefits are not restricted to these contributors. Adequate solutions to address these public good challenges often involve a subset of the eight design principles developed by Elinor Ostrom for "common pool social dilemmas," together with design principles that reflect the public good nature of the problems. This paper is a call to action for scholars and practitioners to broaden our conceptualization and approaches to weed management problems. Such progress begins by evaluating the public good characteristics of specific weed management challenges and applying context-specific design principles to realize successful and sustainable weed management.

Cytokinins (CKs) are a class of compounds that have long been thought to be exclusively plant growth regulators. Interestingly, some species of phytopathogenic bacteria and fungi have been shown to, and gall-inducing insects have been hypothesized to, produce CKs and use them to manipulate their host plants. We used high performance liquid chromatography-electrospray ionization tandem mass spectrometry (HPLC-MS/MS) to examine concentrations of a wide range of CKs in 17 species of phytophagous insects, including gall- and non-gall-inducing species from all six orders of Insecta that contain species known to induce galls: Thysanoptera, Hemiptera, Lepidoptera, Coleoptera, Diptera, and Hymenoptera. We found CKs in all six orders of insects, and they were not associated exclusively with gall-inducing species. We detected 24 different CK analytes, varying in their chemical structure and biological activity. Isoprenoid precursor nucleotide and riboside forms of trans-zeatin (tZ) and isopentenyladenine (iP) were most abundant and widespread across the surveyed insect species. Notably, the observed concentrations of CKs often markedly exceeded those reported in plants suggesting that insects are synthesizing CKs rather than obtaining them from the host plant via tissue consumption, compound sequestration, and bioaccumulation. These findings support insect-derived CKs as means for gall-inducing insects to manipulate their host plant to facilitate cell proliferation, and for both gall- and non-gall-inducing insects to modify nutrient flux and plant defenses during herbivory. Furthermore, wide distribution of CKs across phytophagous insects, including non-gall-inducing species, suggests that insect-borne CKs could be involved in manipulation of source-sink mechanisms of nutrient allocation to sustain the feeding site and altering plant defensive responses, rather than solely gall induction. Given the absence of any evidence for genes in the de novo CK biosynthesis pathway in insects, we postulate that the tRNA-ipt pathway is responsible for CK production. However, the unusually high concentrations of CKs in insects, and the tendency toward dominance of their CK profiles by tZ and iP suggest that the tRNA-ipt pathway functions differently and substantially more efficiently in insects than in plants.

The biocontrol weevil Mogulones crucifer, released in Canada since 1997 but not approved in the USA, is known to sporadically use native nontarget relatives of its target weed for feeding, oviposition, and larval development. Catton et al. released M. crucifer in outbreak densities in patches of mixed target and nontarget plants and nontarget plants in isolation from the target weed to determine if this nontarget herbivory is a spillover effect, or if the weevil establishes and sustains populations on the native plant.

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