There is growing evidence that insects in high-density populations invest relatively more in pathogen resistance than those in low-density populations (i.e. density-dependent prophylaxis). Such increases in resistance are often accompanied by cuticular melanism, which is characteristic of the high-density form of many phase polyphenic insects. Both melanism and pathogen resistance involve the prophenoloxidase enzyme system. In this paper the link between resistance, melanism and phenoloxidase activity is examined in Spodoptera larvae. In S. exempta, cuticular melanism was positively correlated with phenoloxidase activity in the cuticle, haemolymph and midgut. Melanic S. exempta larvae\ud were found to melanize a greater proportion of eggs of the ectoparasitoid Euplectrus laphygmae than non-melanic larvae, and melanic S. littoralis were more resistant to the entomopathogenic fungus Beauveria bassiana (in S. exempta the association between melanism and fungal resistance was non-sign®cant). These results strengthen the link between melanism and disease resistance and implicate the involvement of phenoloxidase

Parasite resistance mechanisms can be costly to maintain. We would therefore predict that organisms should invest in resistance only when it is likely to be required. Insects that show density-dependent phase polyphenism, developing di¡erent phenotypes at high and low population densities, have the opportunity to match their levels of investment in resistance with the likelihood of exposure to pathogens. As high population densities often precipitate disease epidemics, the high-density form should be selected to invest relatively more in resistance. We tested this prediction in larvae of the noctuid Spodoptera exempta. Larvae reared at a high density were found to be considerably more resistant to a nuclear polyhedrosis virus than those reared in isolation. A conspicuous feature of the high-density phase of S. exempta and other phase-polyphenic Lepidoptera is cuticular melanization. As melanization is controlled by the phenoloxidase enzyme system, which is also involved in the immune response, this suggests a possible mechanism for increased resistance at high population densities. We demonstrated that melanized S. exempta larvae were more resistant than non-melanized forms, independent of rearing density. We also found that haemolymph phenoloxidase activity was correlated with cuticular melanization, providing further evidence for a link between melanization and immunity. These results suggest that pathogen resistance in S. exempta is phenotypically plastic, and that the melanized cuticles characteristic of the highdensity form may be indicative of a more active immune system.

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Conservation decision tools based on cost-effectiveness analysis are used to assess threat management strategies for improving species persistence. These approaches rank alternative strategies by their benefit to cost ratio but may fail to identify the optimal sets of strategies to implement under limited budgets because they do not account for redundancies. We devised a multiobjective optimization approach in which the complementarity principle is applied to identify the sets of threat management strategies that protect the most species for any budget. We used our approach to prioritize threat management strategies for 53 species of conservation concern in the Pilbara, Australia. We followed a structured elicitation approach to collect information on the benefits and costs of implementing 17 different conservation strategies during a 3-day workshop with 49 stakeholders and experts in the biodiversity, conservation, and management of the Pilbara. We compared the performance of our complementarity priority threat management approach with a current cost-effectiveness ranking approach. A complementary set of 3 strategies: domestic herbivore management, fire management and research, and sanctuaries provided all species with >50% chance of persistence for $4.7 million/year over 20 years. Achieving the same result cost almost twice as much ($9.71 million/year) when strategies were selected by their cost-effectiveness ranks alone. Our results show that complementarity of management benefits has the potential to double the impact of priority threat management approaches.

1. Threats to biodiversity and the integrity of ecological systems are escalating globally, both within and outside of protected areas. Decision makers have inadequate resources to manage all threats and typically lack information on the likely outcomes and cost-effectiveness of possible management strategies. Priority Threat Management (PTM) is an emerging approach designed to address this challenge, by defining and appraising cost-effective strategies for mitigating threats to biodiversity across regions. The scientific and practical impacts of PTM are increasing, with a growing number of case study applications across the globe.2. Here, we provide guidance and resource material for conducting the PTM process based on our experience delivering six large-scale projects across Australia and Canada. Our handbook describes the four stages of PTM: scoping and planning; defining and collecting key elements; analysing the cost-effectiveness of strategies; and communicating and integrating recommendations. We summarise critical tips, strengths, and limitations and scope for possible enhancements of the approach. Priority Threat Management harnesses scientific and expert-derived informationto prioritise management strategies based on their benefit to biodiversity, management costs and feasibility. The approach involves collaboration with key experts and stakeholders in a region to improve knowledge sharing and conservation support. The PTM approach identifies sets of regional level strategies that together provide the greatest benefits for multiple species under a limited budget, which can be used to inform existing processes for decision-making. 4. The PTM approach applies some generalisations in management strategies and resolution, in order to address complex challenges. Further developments of the approach include testing in a greater range of socioecological systems with adaptations that cater for multiobjective decisions. 482 | Journal of Applied Ecology CARWARDINE Et Al.5. Synthesis and applications. Priority Threat Management is a decision science approach that brings people together to define and prioritise strategies for managing threats to biodiversity across broad regions. It delivers a prospectus for investment in the biodiversity of a region that is transparent, repeatable, participatory, and based on the best available information. Our handbook provides the necessary guidance and resources for expanding the Priority Threat Management approach to new locations, contexts, and challenges. K E Y W O R D S biodiversity conservation, cost-effectiveness analysis, expert elicitation, extinction, Prioritising Threat Management, return on investment, structured decision-making, threatened species | 483 Journal of Applied Ecology CARWARDINE Et Al.