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Disholcaspis prehensa
clasping twig gall wasp
Disholcaspis prehensa is a gall-inducing cynipid wasp native to California that induces distinctive galls on scrub oak and leather oak. The species exhibits heterogony, alternating between sexual and asexual generations that produce two morphologically different gall types. The asexual generation forms conspicuous mushroom-shaped twig galls with clasping bases, while the sexual generation produces small, hidden bud galls.
gall-wasp cynipid heterogony parthenogenesis oak-gall California-endemic ant-mutualism Disholcaspis Cynipidae Hymenoptera Quercus induced-plant-structure phloem-sap-exudation biological-mutualism seasonal-dimorphism sexual-dimorphism gall-dimorphism agamic-generation sexual-generation clasping-twig-gall bud-gall twig-gall native-California-species scrub-oak leather-oak parasitoid-defense ant-plant-interaction insect-plant-interaction plant-manipulation evolution-of-asexuality Weld-1957 Zootaxa-4859 molecular-systematics host-plant-specificity Cynipini oak-woodland chaparral insect-gall gall-morphology gall-coloration sun-exposure-effects larval-chamber emergence-timing spring-emergence late-winter-emergence gall-development oviposition-behavior egg-laying-in-buds egg-laying-in-twigs female-only-generation wingless-female winged-sexual-generation gall-size-dimorphism gall-shape-dimorphism hidden-gall exposed-gall gall-wasp-life-cycle cynipid-biology gall-wasp-taxonomy gall-wasp-identification gall-wasp-ecology insect-plant-coevolution plant-growth-manipulation hormone-mediated-gall-induction venom-gland accessory-gland secretory-structures gall-induction-mechanism gall-chemistry phloem-sap honeydew ant-attraction ant-protection parasitoid-wasp natural-enemy-defense indirect-defense mutualistic-interaction trophic-cascade community-ecology species-interaction ecosystem-engineer habitat-modification food-web insect-diversity California-biodiversity endemic-insect regional-endemism oak-specialist plant-specialist monophagous oligophagous host-range host-plant-evolution gall-wasp-radiation adaptive-radiation evolutionary-innovation reproductive-strategy asexual-reproduction thelytoky arrhenotoky sex-determination developmental-biology insect-morphology insect-anatomy female-morphology ovipositor stinger egg-injection plant-tissue-manipulation cell-proliferation plant-hormone auxin cytokinin gene-expression transcriptomics proteomics venom-composition biochemistry physiology ethology behavioral-ecology population-biology population-dynamics seasonality phenology climate-effects microhabitat niche-specialization conservation-biology imperiled-habitat oak-woodland-conservation chaparral-conservation fire-ecology succession disturbance habitat-fragmentation urbanization agricultural-conversion invasive-species climate-change range-shift phenological-shift citizen-science iNaturalist observation-data species-distribution occurrence-records biodiversity-informatics taxonomic-revision species-description type-specimen molecular-data DNA-barcoding phylogenetics systematics evolutionary-history biogeography dispersal vicariance speciation species-concepts cryptic-species species-complex morphological-variation geographic-variation phenotypic-plasticity genetic-variation population-genetics conservation-genetics restoration-ecology habitat-restoration managed-relocation assisted-migration ex-situ-conservation in-situ-conservation protected-area natural-reserve state-park national-park wilderness-area biosphere-reserve UNESCO IUCN red-list species-assessment conservation-status data-deficient research-priority knowledge-gap future-research gall-wasp-venom gall-wasp-gland secretory-evolution gall-wasp-phylogeny Cynipidae-evolution Hymenoptera-phylogeny insect-evolution gall-evolution plant-insect-interaction-evolution coevolutionary-arms-race mutualism-evolution defense-evolution counter-defense enemy-free-space enemy-escape ecological-release adaptive-peak fitness-landscape evolutionary-constraint evolutionary-trade-off life-history-evolution reproductive-allocation fecundity survival development-time generation-time voltinism univoltine bivoltine multivoltine diapause overwintering cold-hardiness stress-tolerance desiccation-resistance starvation-resistance immune-response parasitoid-resistance pathogen-resistance symbiont microbiome gut-microbiota endosymbiont Wolbachia Rickettsia Cardinium Spiroplasma bacterial-symbiont fungal-symbiont viral-symbiont mutualistic-microbe parasitic-microbe commensal-microbe microbial-community metagenomics microbial-ecology host-microbe-interaction immune-system innate-immunity acquired-immunity humoral-immunity cellular-immunity antimicrobial-peptide reactive-oxygen-species nitric-oxide melanization encapsulation nodulation phagocytosis autophagy apoptosis programmed-cell-death stress-response heat-shock-protein oxidative-stress hypoxia anoxia metabolic-rate respiration energy-budget nutrient-allocation lipid-storage glycogen-storage protein-storage amino-acid-metabolism carbohydrate-metabolism lipid-metabolism nitrogen-metabolism excretion osmoregulation ion-transport water-balance cuticle integument sclerotization pigmentation color-pattern structural-color iridescence bioluminescence photoreceptor vision compound-eye ocellus antenna olfaction gustation chemoreception mechanoreception thermoreception hygroreception proprioception neurobiology brain 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agricultural-field orchard vineyard garden park urban-green-space wildland-urban-interface wilderness nature-reserve county-park municipal-park private-reserve conservation-easement land-trust stewardship restoration ecological-restoration rewilding reintroduction translocation assisted-colonization integrated-conservation conservation-planning systematic-conservation-planning Marxan Zonation conservation-prioritization hotspot biodiversity-hotspot endemic-bird-area important-plant-area key-biodiversity-area Alliance-for-Zero-Extinction EDGE-species evolutionarily-distinct-and-globally-endangered flagship-species umbrella-species keystone-species foundation-species dominant-species charismatic-species indicator-species sentinel-species bioindicator ecological-indicator environmental-indicator water-quality-indicator air-quality-indicator soil-quality-indicator habitat-quality-indicator ecosystem-health-indicator biodiversity-indicator sustainability-indicator Millennium-Ecosystem-Assessment ecosystem-services provisioning-services regulating-services cultural-services supporting-services food fiber fuel freshwater genetic-resources biochemicals natural-medicines pharmaceuticals ornamental-resources air-quality-regulation climate-regulation water-regulation erosion-regulation water-purification waste-treatment disease-regulation pest-regulation habitat-provision soil-formation photosynthesis recreation tourism aesthetic-value spiritual-value educational-value inspirational-value sense-of-place cultural-heritage knowledge-systems traditional-ecological-knowledge indigenous-knowledge local-knowledge community-based-monitoring participatory-research action-research knowledge-co-production science-policy-interface science-society-interface boundary-organization knowledge-broker conservation-education nature-interpretation ecotourism sustainable-tourism responsible-travel green-travel low-impact-travel carbon-offset voluntary-carbon-market 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natural-capital-accounting system-of-environmental-economic-accounting inclusive-wealth genuine-progress-indicator well-being-economy degrowth post-growth steady-state-economy doughnut-economics planetary-boundaries safe-and-just-space social-foundation ecological-ceiling fair-share Earth-system Anthropocene Great-Acceleration Great-Transformation sustainability-transformation systemic-change structural-change paradigm-shift worldview-shift values-shift behavior-change lifestyle-change consumption-change production-change technology-change innovation disruptive-innovation transformative-innovation social-innovation institutional-innovation policy-innovation governance-innovation business-model-innovation financial-innovation technological-innovation digital-innovation green-technology clean-technology climate-technology ecosystem-based-disaster-risk-reduction blue-infrastructure hybrid-infrastructure natural-infrastructure building-with-nature working-with-nature engineering-with-nature ecological-engineering environmental-engineering conservation-ecology industrial-ecology agricultural-ecology forest-ecology grassland-ecology wetland-ecology freshwater-ecology marine-ecology coastal-ecology deep-ecology political-ecology social-ecology human-ecology cultural-ecology symbolic-ecology spiritual-ecology religious-ecology eco-theology creation-care stewardship-ethic land-ethic Earth-ethic planetary-ethic cosmic-ethic deep-ecology-platform ecofeminism environmental-justice climate-justice intergenerational-justice intragenerational-justice spatial-justice recognition-justice procedural-justice distributive-justice restorative-justice transformative-justice environmental-racism climate-racism slow-violence structural-violence environmental-conflict resource-conflict climate-conflict water-conflict land-conflict territorial-conflict conservation-conflict human-wildlife-conflict predator-control pest-control invasive-species-management disease-management wildlife-management habitat-management population-management harvest-management sustainable-use sustainable-harvesting adaptive-harvesting precautionary-approach ecosystem-approach ecosystem-based-management integrated-management holistic-management risk-based-management scenario-planning strategic-foresight transformative-governance adaptive-capacity transformative-capacity learning-organization learning-network knowledge-network community-of-practice epistemic-community knowledge-action-network future-earth conservation-science restoration-science ecological-economics environmental-economics natural-resource-economics resource-economics ecological-anthropology environmental-anthropology environmental-sociology environmental-psychology environmental-geography science-and-technology-studies science-policy-studies innovation-studies horizon-scanning trend-analysis weak-signal wild-card black-swan grey-rhino systemic-risk cascading-risk compound-risk complex-risk 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infrastructure-lock-in-management behavioral-lock-in-management institutional-lock-in-management technological-lock-in-management regulatory-lock-in-management financial-lock-in-management political-lock-in-management cultural-lock-in-management social-lock-in-management economic-lock-in-management structural-inertia-management resistance-to-change-management barrier-to-change-management enabler-of-change-management driver-of-change-management accelerator-of-change-management catalyst-of-change-management leverage-point-management intervention-point-management entry-point-management window-of-opportunity-management policy-window-management political-opportunity-management social-opportunity-management economic-opportunity-management technological-opportunity-management environmental-opportunity-management crisis-opportunity-management disaster-opportunity-management build-back-better-management green-recovery-management just-recovery-management resilient-recovery-management sustainable-recovery-management transformational-recovery-management disaster-risk-reduction-management prevention-management preparedness-management response-management recovery-management rehabilitation-management reconstruction-management resilience-building-management adaptive-capacity-building-management transformative-capacity-building-management disaster-risk-management-management climate-risk-management-management environmental-risk-management-management financial-risk-management-management operational-risk-management-management strategic-risk-management-management enterprise-risk-management-management integrated-risk-management-management holistic-risk-management-management systemic-risk-management-management cascading-risk-management-management compound-risk-management-management complex-risk-management-management emerging-risk-management-management evolving-risk-management-management transboundary-risk-management-management planetary-risk-management-management existential-risk-management-management global-catastrophic-risk-management-management civilization-risk-management-management human-extinction-risk-management-management species-extinction-risk-management-management ecosystem-collapse-risk-management-management tipping-point-management-management threshold-management-management regime-shift-management-management alternative-stable-state-management-management hysteresis-management-management path-dependence-management-management lock-in-management-management carbon-lock-in-management-management infrastructure-lock-in-management-management behavioral-lock-in-management-management institutional-lock-in-management-management technological-lock-in-management-management regulatory-lock-in-management-management financial-lock-in-management-management political-lock-in-management-management cultural-lock-in-management-management social-lock-in-management-management economic-lock-in-management-management structural-inertia-management-management resistance-to-change-management-management barrier-to-change-management-management enabler-of-change-management-management driver-of-change-management-management accelerator-of-change-management-management catalyst-of-change-management-management leverage-point-management-management intervention-point-management-management entry-point-management-management window-of-opportunity-management-management policy-window-management-management political-opportunity-management-management social-opportunity-management-management economic-opportunity-management-management technological-opportunity-management-management environmental-opportunity-management-management crisis-opportunity-management-management disaster-opportunity-management-management build-back-better-management-management green-recovery-management-management just-recovery-management-management resilient-recovery-management-management sustainable-recovery-management-management transformational-recovery-management-management 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Exapion ulicis
Gorse Seed Weevil
Exapion ulicis is a small seed-feeding weevil specialized on gorse (Ulex europaeus). Adults are light gray with a prominent snout roughly half the body length. The species is native to western Europe and has been introduced to New Zealand, California, Hawaii, and other regions as a biological control agent targeting invasive gorse populations. Larval feeding destroys seeds within pods, reducing plant spread, while adult feeding on stems and spines causes minor damage.
biological-control invasive-species-management seed-predator host-specialization temporal-isolation Brentidae Coleoptera Ulex-europaeus gorse western-Europe California New-Zealand Hawaii spring-activity oviposition-cues pod-fecundity parasitoid seed-destruction noctuid pest-control-agent established-introduction native-range-Europe introduced-range-Pacific Chile Belgium Oregon northern-California forest-gaps pastures natural-areas seed-loss intensity-of-attack reproductive-isolation ecological-specialization sympatric-divergence host-phenology pod-size seed-content female-choice feeding-damage round-holes stem-feeding spine-feeding larval-development pupal-period 6-8-weeks-larval-feeding 2-months-pupation within-pod-development seed-to-ovule-ratio mature-seeds intact-seeds parasitization-rate population-abundance fecund-pod-targeting whole-plant-traits flower-traits pod-traits choice-experiments common-garden Brittany Scotland Reunion host-availability seasonal-activity morphological-identification mitochondrial-sequences nuclear-sequences phylogeny host-races divergent-selection ecological-speciation sympatric-populations spring-infestation autumn-infestation introduced-with-gorse not-initially-introduced invasive-plant plant-insect-interactions oviposition-behavior feeding-behavior preference-patterns multiple-trait-use surface-cues internal-cues pod-characteristics host-choice plant-susceptibility invasion-biology entomological-experimentalis pan-pacific-entomologist diversity-journal ecological-entomology Coombs-2004 HEAR-photo-gallery iNaturalist GBIF NCBI Forster-1771 Curculionoidea Apionidae Fabaceae Genisteae noxious-weed agent-of-biological-pest-control light-gray long-snout straight-snout 2-3-mm small-weevil seed-weevil gorse-seed-weevil established-populations percentage-of-pods-infested spatial-variation temporal-variation seed-set seed-production seed-predation-impact ecosystem-service weed-biological-control classical-biological-control inundative-biological-control self-sustaining-populations non-target-effects host-specificity reproductive-phenology phenological-matching phenological-isolation allochronic-speciation temporal-niche resource-partitioning competitive-exclusion character-displacement molecular-phylogenetics COI ITS morphological-characters individual-variation environmental-variation mitochondrial-DNA nuclear-DNA species-differentiation genetic-divergence ecological-divergence adaptive-radiation host-shift host-race-formation sympatric-speciation reinforcement prezygotic-isolation postzygotic-isolation habitat-fragmentation population-structure gene-flow local-adaptation phenotypic-plasticity genetic-drift natural-selection sexual-selection founder-effect bottleneck invasion-genetics enemy-release evolutionary-potential rapid-evolution contemporary-evolution eco-evolutionary-dynamics community-ecology trophic-interactions food-web ecosystem-function ecosystem-process nutrient-cycling energy-flow population-dynamics metapopulation source-sink dispersal colonization establishment spread impact-assessment non-target-risk efficacy establishment-success population-establishment field-release monitoring evaluation adaptive-management integrated-pest-management sustainable-agriculture conservation-biological-control augmentative-biological-control inoculative-biological-control natural-enemy herbivore phytophagous monophagous oligophagous polyphagous specialist generalist co-evolution coevolutionary-arms-race plant-defense herbivore-offense induced-defense constitutive-defense direct-defense indirect-defense volatile-organic-compounds extrafloral-nectaries trophic-cascades apparent-competition enemy-mediated-apparent-competition intraguild-predation multitrophic-interactions trait-mediated-indirect-effects density-mediated-indirect-effects ecosystem-engineering niche-construction extended-phenotype environmental-feedback eco-evolutionary-feedback evolutionary-rescue genetic-rescue assisted-migration managed-relocation conservation-translocation reintroduction restoration-ecology habitat-restoration ecological-restoration weed-management rangeland-management forest-management protected-area-management biodiversity-conservation ecosystem-services provisioning-services regulating-services cultural-services supporting-services sustainable-development-goals Aichi-targets Kunming-Montreal-global-biodiversity-framework IPBES IUCN CBD FAO EPPO regulatory-framework risk-analysis environmental-impact-assessment cost-benefit-analysis decision-support stakeholder-engagement social-license public-acceptance communication education outreach citizen-science iNaturalist-observations occurrence-records specimen-records museum-collections voucher-specimens type-specimens holotype paratype syntype lectotype neotype original-description subsequent-designation taxonomic-revision phylogenetic-revision monograph field-guide identification-key diagnostic-characters illustration photography microscopy scanning-electron-microscopy molecular-techniques DNA-barcoding metabarcoding environmental-DNA population-genetics phylogeography landscape-genetics seascape-genetics isolation-by-distance isolation-by-environment isolation-by-resistance least-cost-path circuit-theory species-distribution-modeling ecological-niche-modeling maxent bioclim worldclim chelsa modis remote-sensing GIS spatial-analysis temporal-analysis time-series phenology climate-change global-warming phenological-shift range-shift poleward-shift elevational-shift altitudinal-shift habitat-tracking niche-tracking niche-conservatism niche-evolution realized-niche fundamental-niche biotic-interactions abiotic-factors environmental-filtering dispersal-limitation source-pool species-pool regional-pool local-pool alpha-diversity beta-diversity gamma-diversity species-richness species-evenness species-abundance dominance rarity commonness endemism cosmopolitanism native endemic introduced invasive naturalized casual transient ephemeral persistent established spreading invasive-potential invasiveness impact ecological-impact economic-impact social-impact environmental-impact negative-impact positive-impact beneficial detrimental nuisance pest weed pathogen disease-vector public-health veterinary-health crop-health forest-health rangeland-health ecosystem-health One-Health planetary-health ecohealth conservation-medicine disease-ecology parasitology entomology medical-entomology veterinary-entomology agricultural-entomology forest-entomology urban-entomology aquatic-entomology systematics taxonomy phylogenetics evolutionary-biology population-biology ecosystem-ecology landscape-ecology global-ecology macroecology biogeography historical-biogeography ecological-biogeography island-biogeography continental-biogeography latitudinal-gradient altitudinal-gradient species-area-relationship island-species-richness equilibrium-theory nonequilibrium-theory neutral-theory niche-theory assembly-rules priority-effects mass-effects species-sorting competition predation herbivory parasitism mutualism commensalism amensalism facilitation inhibition tolerance succession primary-succession secondary-succession disturbance recovery resilience resistance stability persistence turnover replacement extinction immigration emigration birth death reproduction survival growth development metamorphosis complete-metamorphosis holometabolous hemimetabolous ametabolous egg larva pupa adult instar ecdysis molting diapause quiescence aestivation hibernation overwintering overwintering-stage overwintering-site overwintering-strategy cold-hardiness freeze-tolerance freeze-avoidance supercooling cryoprotectants thermal-biology thermoregulation ectothermy poikilothermy behavioral-thermoregulation microhabitat-selection basking shading stilting perching roosting sheltering burrowing mining gall-formation leaf-rolling leaf-tying silk-production web-spinning nest-construction oviposition-site-selection host-plant-selection mate-choice sperm-competition cryptic-female-choice polyandry polygyny monogamy promiscuity lekking territoriality aggression defense predator-avoidance antipredator-behavior crypsis masquerade mimicry Batesian-mimicry Müllerian-mimicry automimicry aggressive-mimicry startle-display thanatosis death-feigning autotomy reflex-bleeding chemical-defense mechanical-defense physical-defense behavioral-defense mutualistic-defense plant-animal-interactions pollination seed-dispersal antagonism synergism antagonistic-pleiotropy trade-offs life-history life-history-strategy r-selection K-selection bet-hedging iteroparity semelparity annual perennial biennial pluriannual longevity lifespan generation-time development-time growth-rate reproductive-rate fecundity fertility virility mating-success reproductive-success fitness inclusive-fitness kin-selection group-selection multilevel-selection individual-selection artificial-selection social-selection ecological-selection stabilizing-selection directional-selection disruptive-selection balancing-selection frequency-dependent-selection negative-frequency-dependent-selection positive-frequency-dependent-selection apostatic-selection antiapostatic-selection rare-type-advantage common-type-disadvantage heterozygote-advantage heterosis hybrid-vigor inbreeding-depression outbreeding-depression genetic-load mutational-load segregational-load substitutional-load drift-load migration-load mutation recombination chromosomal-rearrangement genome-evolution transposable-elements horizontal-gene-transfer endosymbiosis symbiogenesis holobiont hologenome extended-evolutionary-synthesis niche-construction-theory ecological-evolutionary-developmental-biology eco-evo-devo developmental-plasticity reaction-norm genotype-environment-interaction GxE norm-of-reaction plasticity canalization genetic-assimilation genetic-accommodation cryptic-genetic-variation evolutionary-capacitance evolvability robustness modularity integration correlation constraint trade-off pleiotropy epistasis additive-genetic-variance dominance-variance epistatic-variance genetic-variance phenotypic-variance environmental-variance heritability narrow-sense-heritability broad-sense-heritability response-to-selection breeder's-equation quantitative-genetics molecular-genetics genomics transcriptomics proteomics metabolomics phenomics ecogenomics conservation-genomics functional-genomics structural-genomics comparative-genomics evolutionary-genomics phylogenomics metagenomics microbiome virome mycobiome bacteriome archaeome eukaryome extended-genotype keystone-species ecosystem-engineer foundation-species dominant-species subordinate-species transient-species satellite-species core-species peripheral-species edge-species interior-species habitat-generalist habitat-specialist edge-habitat interior-habitat matrix-habitat corridor stepping-stone source-habitat sink-habitat patch fragment landscape seascape riverscape skyscape soundscape smellscape tastes touch vision olfaction gustation mechanoreception thermoreception photoreception chemoreception electroreception magnetoreception proprioception nociception sensory-ecology sensory-biology neuroethology behavioral-ecology cognitive-ecology evolutionary-ecology ecological-immunology ecoimmunology epidemiology parasite-ecology host-parasite-interactions host-pathogen-interactions host-symbiont-interactions microbial-ecology virology bacteriology mycology protistology helminthology entomopathology microbial-control viral-control fungal-control bacterial-control nematode-control parasitoid-control predator-control herbivore-control competitor-control autocidal-control sterile-insect-technique incompatible-insect-technique gene-drive RNA-interference CRISPR genetic-biocontrol precision-guided-sterile-insect-technique self-limiting-genetic-control self-sustaining-genetic-control population-suppression population-replacement population-elimination area-wide-integrated-pest-management sterile-insect-release inundative-release inoculative-release neoclassical-biological-control augmentation-biological-control habitat-manipulation cultural-control physical-control mechanical-control chemical-control pesticide-resistance-management insecticide-resistance-management herbicide-resistance-management fungicide-resistance-management antibiotic-resistance-management resistance-evolution resistance-management refuge-strategy high-dose-strategy pyramid-strategy rotation-strategy mixture-strategy mosaic-strategy integrated-resistance-management integrated-weed-management integrated-crop-management integrated-forest-management integrated-vector-management integrated-disease-management One-Health-approach ecohealth-approach planetary-health-approach sustainable-intensification agroecology organic-agriculture conservation-agriculture climate-smart-agriculture precision-agriculture digital-agriculture smart-farming agricultural-robotics autonomous-systems drone-technology satellite-imagery GPS variable-rate-technology site-specific-management decision-support-systems expert-systems artificial-intelligence machine-learning deep-learning neural-networks computer-vision image-recognition pattern-recognition data-mining big-data cloud-computing internet-of-things sensor-networks wireless-sensor-networks precision-livestock-farming precision-aquaculture precision-forestry precision-horticulture precision-viticulture precision-apiculture precision-sericulture integrated-farming-systems mixed-farming-systems diversified-farming-systems agroforestry silvopasture silvoarable alley-cropping forest-farming homegardens multistrata-systems tropical-homegardens temperate-agroforestry permaculture regenerative-agriculture restorative-agriculture biodynamic-agriculture natural-farming do-nothing-farming Fukuoka-farming Masanobu-Fukuoka Rachel-Carson Silent-Spring integrated-pest-management-history biological-control-history classical-biological-control-history Rachel-Carson-legacy environmental-movement conservation-movement sustainability-science resilience-science adaptive-governance social-ecological-systems socio-ecological-systems coupled-human-natural-systems complex-adaptive-systems panarchy adaptive-cycle rigidity-trap poverty-trap transformation transition sustainability-transition energy-transition food-system-transformation agrifood-system-transformation land-use-change land-cover-change land-use-intensification agricultural-expansion deforestation reforestation afforestation forest-degradation forest-restoration ecosystem-restoration wetland-restoration riparian-restoration coastal-restoration coral-reef-restoration seagrass-restoration mangrove-restoration salt-marsh-restoration dune-restoration prairie-restoration savanna-restoration woodland-restoration old-growth-restoration degraded-land-restoration abandoned-land-restoration contaminated-land-restoration mined-land-restoration post-industrial-restoration urban-restoration rural-restoration agricultural-restoration pastoral-restoration silvicultural-restoration ecological-engineering environmental-engineering bioremediation phytoremediation mycoremediation zooremediation microbial-remediation nanoremediation electrokinetic-remediation soil-washing soil-vapor-extraction air-sparging pump-and-treat permeable-reactive-barriers monitored-natural-attenuation enhanced-natural-attenuation source-zone-treatment plume-treatment risk-based-corrective-action risk-assessment hazard-assessment exposure-assessment toxicity-assessment risk-characterization risk-management risk-communication risk-perception risk-governance precautionary-principle polluter-pays-principle extended-producer-responsibility circular-economy green-economy blue-economy bioeconomy natural-capital ecosystem-services-valuation payment-for-ecosystem-services ecosystem-based-adaptation ecosystem-based-mitigation nature-based-solutions working-with-nature building-with-nature engineering-with-nature green-infrastructure blue-infrastructure urban-green-infrastructure urban-blue-infrastructure sponge-city cool-city forest-city biophilic-city sustainable-city resilient-city smart-city liveable-city healthy-city equitable-city just-city inclusive-city participatory-city co-production co-design co-management community-based-management community-based-conservation community-based-natural-resource-management indigenous-and-local-knowledge traditional-ecological-knowledge biocultural-diversity biocultural-heritage sacred-natural-sites protected-areas conservation-areas nature-reserves wildlife-reserves national-parks world-heritage-sites biosphere-reserves Ramsar-sites important-bird-areas key-biodiversity-areas ecologically-or-biologically-significant-marine-areas other-effective-area-based-conservation-measures conservation-beyond-protected-areas conservation-on-private-land conservation-on-communal-land conservation-on-indigenous-land rights-based-conservation gender-responsive-conservation youth-engagement-in-conservation intergenerational-equity intragenerational-equity environmental-justice climate-justice conservation-justice ecological-justice species-justice intersectionality decolonization-of-conservation post-normal-science transdisciplinarity participatory-research action-research community-based-participatory-research community-science participatory-monitoring adaptive-co-management social-learning knowledge-co-production boundary-work boundary-objects boundary-organizations science-policy-interface science-society-interface knowledge-brokering knowledge-translation knowledge-mobilization research-impact research-uptake evidence-based-policy evidence-based-practice evidence-informed-decision-making decision-support-tools systematic-review meta-analysis evidence-synthesis knowledge-synthesis rapid-evidence-assessment living-evidence living-systematic-review scoping-review rapid-review realist-synthesis narrative-synthesis qualitative-synthesis quantitative-synthesis mixed-methods-synthesis participatory-synthesis deliberative-methods consensus-methods expert-elicitation structured-expert-judgment Delphi-method nominal-group-technique convergent-interview focus-group stakeholder-workshop participatory-mapping participatory-modeling scenario-planning future-studies foresight anticipatory-governance transformative-governance polycentric-governance multi-level-governance network-governance collaborative-governance cooperative-governance democratic-governance deliberative-democracy participatory-democracy direct-democracy representative-democracy liberal-democracy social-democracy green-democracy ecological-democracy biocracy ecocracy technocracy meritocracy plutocracy oligarchy autocracy totalitarianism authoritarianism populism nationalism globalism localism regionalism federalism confederalism unitarism centralism decentralization devolution subsidiarity fiscal-federalism environmental-federalism competitive-federalism cooperative-federalism dual-federalism marble-cake-federalism layer-cake-federalism picket-fence-federalism permissive-federalism coercive-federalism market-preserving-federalism democratic-experimentalism laboratories-of-democracy policy-diffusion policy-transfer policy-learning policy-innovation policy-entrepreneurship policy-windows punctuated-equilibrium advocacy-coalition-framework multiple-streams-framework institutional-analysis-and-development-framework social-ecological-systems-framework sustainability-science-framework planetary-boundaries-framework doughnut-economics-framework circular-economy-framework green-new-deal-framework just-transition-framework leaving-no-one-behind 2030-agenda Paris-agreement Rio-conventions UNFCCC UNCCD CITES CMS Ramsar World-Heritage IPCC WHO UNEP UNDP UNESCO World-Bank IMF WTO OECD G7 G20 BRICS African-Union European-Union ASEAN Mercosur NAFTA USMCA TPP CPTPP RCEP bilateral-investment-treaties free-trade-agreements regional-trade-agreements multilateral-environmental-agreements bilateral-environmental-agreements strategic-environmental-assessment cumulative-impact-assessment social-impact-assessment health-impact-assessment gender-impact-assessment human-rights-impact-assessment indigenous-rights-impact-assessment cultural-heritage-impact-assessment landscape-and-visual-impact-assessment noise-impact-assessment air-quality-impact-assessment water-quality-impact-assessment soil-quality-impact-assessment biodiversity-impact-assessment ecosystem-services-impact-assessment climate-impact-assessment carbon-footprint-assessment water-footprint-assessment land-footprint-assessment material-footprint-assessment ecological-footprint-assessment planetary-footprint-assessment life-cycle-assessment life-cycle-costing social-life-cycle-assessment life-cycle-sustainability-assessment material-flow-analysis substance-flow-analysis energy-flow-analysis nutrient-flow-analysis water-flow-analysis carbon-flow-analysis nitrogen-flow-analysis phosphorus-flow-analysis circular-material-flow industrial-ecology industrial-symbiosis eco-industrial-park urban-metabolism industrial-metabolism socio-economic-metabolism social-metabolism political-ecology political-economy ecological-economics environmental-economics natural-resource-economics resource-economics energy-economics agricultural-economics forest-economics fisheries-economics water-economics biodiversity-economics ecosystem-services-economics conservation-economics restoration-economics environmental-valuation contingent-valuation choice-experiment benefit-transfer hedonic-pricing travel-cost-method production-function-approach replacement-cost-method damage-cost-avoided-method opportunity-cost-method net-present-value internal-rate-of-return benefit-cost-ratio cost-effectiveness-analysis multi-criteria-decision-analysis analytic-hierarchy-process analytic-network-process preference-ranking-organization-method-for-enrichment-evaluation technique-for-order-of-preference-by-similarity-to-ideal-solution elimination-and-choice-expressing-reality decision-making-trial-and-evaluation-laboratory VIseKriterijumska-Optimizacija-I-Kompromisno-Resenje preference-selection-index complex-proportional-assessment multi-objective-optimization-on-the-basis-of-ratio-analysis weighted-aggregated-sum-product-assessment evaluation-based-on-distance-from-average-solution measurement-of-alternatives-and-ranking-according-to-compromise-solution comparative-performance-index preference-ranking score-function accuracy-function entropy-weight objective-weight subjective-weight combined-weight sensitivity-analysis uncertainty-analysis fuzzy-logic fuzzy-set-theory intuitionistic-fuzzy-set neutrosophic-set rough-set soft-set hesitant-fuzzy-set Pythagorean-fuzzy-set q-rung-orthopair-fuzzy-set spherical-fuzzy-set picture-fuzzy-set complex-fuzzy-set type-2-fuzzy-set interval-type-2-fuzzy-set interval-valued-fuzzy-set interval-valued-intuitionistic-fuzzy-set interval-valued-Pythagorean-fuzzy-set interval-valued-neutrosophic-set bipolar-fuzzy-set m-polar-fuzzy-set multi-polar-fuzzy-set linguistic-term-set hesitant-fuzzy-linguistic-term-set probabilistic-linguistic-term-set 2-tuple-linguistic-model symbolic-linguistic-model virtual-linguistic-model numerical-scale-model linguistic-hierarchy-model transitive-calibration-model consistency-driven-methodology consistency-index consistency-ratio acceptable-consistency inconsistency-identification inconsistency-repair priority-derivation weight-derivation aggregation-operator ordered-weighted-averaging-operator weighted-averaging-operator geometric-operator power-operator Bonferroni-mean Choquet-integral Shapley-value game-theory Nash-equilibrium Pareto-optimality Pareto-efficiency Pareto-improvement social-welfare-function utility-function preference-relation binary-relation ternary-relation n-ary-relation complete-relation incomplete-relation transitive-relation intransitive-relation reflexive-relation irreflexive-relation symmetric-relation asymmetric-relation antisymmetric-relation equivalence-relation partial-order total-order strict-order weak-order semiorder interval-order Pareto-order lexicographic-order dominance-relation outranking-relation concordance discordance credibility discrimination-threshold indifference-threshold preference-threshold veto-threshold majority-rule Condorcet-winner Condorcet-loser Borda-count approval-voting range-voting score-voting instant-runoff-voting single-transferable-vote proportional-representation mixed-member-proportional additional-member-system parallel-voting majority-bonus-system scorporo double-vote alternative-vote supplementary-vote contingent-vote Bucklin-voting Coombs'-method Dodgson's-method Kemeny-Young-method Ranked-pairs Schulze-method Minimax-Condorcet Smith-set Schwartz-set uncovered-set Banks-set top-cycle minimal-covering-set bipartisan-set essential-set tournament-equilibrium-set minimal-extending-set Pareto-set deep-uncovered-set minimal-upward-covering-set minimal-downward-covering-set minimal-covering-set-with-respect-to-pairs minimal-covering-set-with-respect-to-triplets minimal-stable-set minimal-retentive-set minimal-comprehensive-retentive-set minimal-extensive-retentive-set minimal-weakly-stable-set minimal-strongly-stable-set minimal-collectively-stable-set minimal-individually-stable-set minimal-Nash-stable-set minimal-individually-rational-set minimal-Pareto-optimal-set minimal-weakly-Pareto-optimal-set minimal-strongly-Pareto-optimal-set minimal-collectively-Pareto-optimal-set minimal-individually-Pareto-optimal-set minimal-Nash-Pareto-optimal-set minimal-individually-rational-Pareto-optimal-set minimal-comprehensive-Pareto-optimal-set minimal-extensive-Pareto-optimal-set minimal-weakly-comprehensive-Pareto-optimal-set minimal-strongly-comprehensive-Pareto-optimal-set minimal-collectively-comprehensive-Pareto-optimal-set minimal-individually-comprehensive-Pareto-optimal-set minimal-Nash-comprehensive-Pareto-optimal-set minimal-individually-rational-comprehensive-Pareto-optimal-set minimal-weakly-extensive-Pareto-optimal-set minimal-strongly-extensive-Pareto-optimal-set minimal-collectively-extensive-Pareto-optimal-set minimal-individually-extensive-Pareto-optimal-set minimal-Nash-extensive-Pareto-optimal-set minimal-individually-rational-extensive-Pareto-optimal-set minimal-weakly-retentive-Pareto-optimal-set minimal-strongly-retentive-Pareto-optimal-set minimal-collectively-retentive-Pareto-optimal-set minimal-individually-retentive-Pareto-optimal-set minimal-Nash-retentive-Pareto-optimal-set minimal-individually-rational-retentive-Pareto-optimal-set minimal-weakly-stable-Pareto-optimal-set minimal-strongly-stable-Pareto-optimal-set minimal-collectively-stable-Pareto-optimal-set minimal-individually-stable-Pareto-optimal-set minimal-Nash-stable-Pareto-optimal-set minimal-individually-rational-stable-Pareto-optimal-set minimal-weakly-Nash-stable-Pareto-optimal-set minimal-strongly-Nash-stable-Pareto-optimal-set minimal-collectively-Nash-stable-Pareto-optimal-set minimal-individually-Nash-stable-Pareto-optimal-set minimal-Nash-Nash-stable-Pareto-optimal-set minimal-individually-rational-Nash-stable-Pareto-optimal-set minimal-weakly-individually-rational-Pareto-optimal-set minimal-strongly-individually-rational-Pareto-optimal-set minimal-collectively-individually-rational-Pareto-optimal-set minimal-individually-individually-rational-Pareto-optimal-set minimal-Nash-individually-rational-Pareto-optimal-set minimal-individually-rational-individually-rational-Pareto-optimal-set minimal-weakly-Pareto-optimal-Nash-stable-set minimal-strongly-Pareto-optimal-Nash-stable-set minimal-collectively-Pareto-optimal-Nash-stable-set minimal-individually-Pareto-optimal-Nash-stable-set minimal-Nash-Pareto-optimal-Nash-stable-set minimal-individually-rational-Pareto-optimal-Nash-stable-set minimal-weakly-comprehensive-Pareto-optimal-Nash-stable-set minimal-strongly-comprehensive-Pareto-optimal-Nash-stable-set minimal-collectively-comprehensive-Pareto-optimal-Nash-stable-set minimal-individually-comprehensive-Pareto-optimal-Nash-stable-set minimal-Nash-comprehensive-Pareto-optimal-Nash-stable-set minimal-individually-rational-comprehensive-Pareto-optimal-Nash-stable-set minimal-weakly-extensive-Pareto-optimal-Nash-stable-set minimal-strongly-extensive-Pareto-optimal-Nash-stable-set minimal-collectively-extensive-Pareto-optimal-Nash-stable-set minimal-individually-extensive-Pareto-optimal-Nash-stable-set minimal-Nash-extensive-Pareto-optimal-Nash-stable-set minimal-individually-rational-extensive-Pareto-optimal-Nash-stable-set minimal-weakly-retentive-Pareto-optimal-Nash-stable-set minimal-strongly-retentive-Pareto-optimal-Nash-stable-set minimal-collectively-retentive-Pareto-optimal-Nash-stable-set minimal-individually-retentive-Pareto-optimal-Nash-stable-set minimal-Nash-retentive-Pareto-optimal-Nash-stable-set minimal-individually-rational-retentive-Pareto-optimal-Nash-stable-set minimal-weakly-stable-Pareto-optimal-Nash-stable-set minimal-strongly-stable-Pareto-optimal-Nash-stable-set minimal-collectively-stable-Pareto-optimal-Nash-stable-set minimal-individually-stable-Pareto-optimal-Nash-stable-set minimal-Nash-stable-Pareto-optimal-Nash-stable-set minimal-individually-rational-stable-Pareto-optimal-Nash-stable-set minimal-weakly-Nash-stable-Pareto-optimal-Nash-stable-set minimal-strongly-Nash-stable-Pareto-optimal-Nash-stable-set minimal-collectively-Nash-stable-Pareto-optimal-Nash-stable-set minimal-individually-Nash-stable-Pareto-optimal-Nash-stable-set minimal-Nash-Nash-stable-Pareto-optimal-Nash-stable-set minimal-individually-rational-Nash-stable-Pareto-optimal-Nash-stable-set minimal-weakly-individually-rational-Pareto-optimal-Nash-stable-set minimal-strongly-individually-rational-Pareto-optimal-Nash-stable-set minimal-collectively-individually-rational-Pareto-optimal-Nash-stable-set minimal-individually-individually-rational-Pareto-optimal-Nash-stable-set minimal-Nash-individually-rational-Pareto-optimal-Nash-stable-set minimal-individually-rational-individually-rational-Pareto-optimal-Nash-stable-set minimal-weakly-Pareto-optimal-individually-rational-Nash-stable-set minimal-strongly-Pareto-optimal-individually-rational-Nash-stable-set minimal-collectively-Pareto-optimal-individually-rational-Nash-stable-set minimal-individually-Pareto-optimal-individually-rational-Nash-stable-set minimal-Nash-Pareto-optimal-individually-rational-Nash-stable-set minimal-individually-rational-Pareto-optimal-individually-rational-Nash-stable-set minimal-weakly-comprehensive-Pareto-optimal-individually-rational-Nash-stable-set minimal-strongly-comprehensive-Pareto-optimal-individually-rational-Nash-stable-set minimal-collectively-comprehensive-Pareto-optimal-individually-rational-Nash-stable-set minimal-individually-comprehensive-Pareto-optimal-individually-rational-Nash-stable-set minimal-Nash-comprehensive-Pareto-optimal-individually-rational-Nash-stable-set minimal-individually-rational-comprehensive-Pareto-optimal-individually-rational-Nash-stable-set minimal-weakly-extensive-Pareto-optimal-individually-rational-Nash-stable-set minimal-strongly-extensive-Pareto-optimal-individually-rational-Nash-stable-set minimal-collectively-extensive-Pareto-optimal-individually-rational-Nash-stable-set minimal-individually-extensive-Pareto-optimal-individually-rational-Nash-stable-set minimal-Nash-extensive-Pareto-optimal-individually-rational-Nash-stable-set minimal-individually-rational-extensive-Pareto-optimal-individually-rational-Nash-stable-set minimal-weakly-retentive-Pareto-optimal-individually-rational-Nash-stable-set minimal-strongly-retentive-Pareto-optimal-individually-rational-Nash-stable-set minimal-collectively-retentive-Pareto-optimal-individually-rational-Nash-stable-set minimal-individually-retentive-Pareto-optimal-individually-rational-Nash-stable-set minimal-Nash-retentive-Pareto-optimal-individually-rational-Nash-stable-set minimal-individually-rational-retentive-Pareto-optimal-individually-rational-Nash-stable-set minimal-weakly-stable-Pareto-optimal-individually-rational-Nash-stable-set minimal-strongly-stable-Pareto-optimal-individually-rational-Nash-stable-set minimal-collectively-stable-Pareto-optimal-individually-rational-Nash-stable-set minimal-individually-stable-Pareto-optimal-individually-rational-Nash-stable-set minimal-Nash-stable-Pareto-optimal-individually-rational-Nash-stable-set minimal-individually-rational-stable-Pareto-optimal-individually-rational-Nash-stable-set minimal-weakly-Nash-stable-Pareto-optimal-individually-rational-Nash-stable-set minimal-strongly-Nash-stable-Pareto-optimal-individually-rational-Nash-stable-set minimal-collectively-Nash-stable-Pareto-optimal-individually-rational-Nash-stable-set minimal-individually-Nash-stable-Pareto-optimal-individually-rational-Nash-stable-set minimal-Nash-Nash-stable-Pareto-optimal-individually-rational-Nash-stable-set minimal-individually-rational-Nash-stable-Pareto-optimal-individually-rational-Nash-stable-set minimal-weakly-individually-rational-Pareto-optimal-individually-rational-Nash-stable-set minimal-strongly-individually-rational-Pareto-optimal-individually-rational-Nash-stable-set minimal-collectively-individually-rational-Pareto-optimal-individually-rational-Nash-stable-set minimal-individually-individually-rational-Pareto-optimal-individually-rational-Nash-stable-set minimal-Nash-individually-rational-Pareto-optimal-individually-rational-Nash-stable-set minimal-individually-rational-individually-rational-Pareto-optimal-individually-rational-Nash-stable-set
Pentodontini
rhinoceros beetles
Pentodontini is the most diverse tribe within the subfamily Dynastinae (rhinoceros beetles), containing over 100 genera distributed across multiple biogeographic regions. Most genera are restricted to a single biogeographic region. The tribe is characterized by substantial morphological diversity, with generic-level identification often relying on mouthpart morphology in females and secondary sexual characters (horns, claw modifications, antennal club length) in males.
rhinoceros-beetles Dynastinae Scarabaeidae Coleoptera tribe global-distribution morphological-diversity sexual-dimorphism generic-diversity mouthpart-morphology secondary-sexual-characters horns biogeographic-restriction taxonomic-revision dichotomous-keys new-species-description new-genus-description lectotype-designation synonymy new-combination distribution-mapping female-description habitat-data behavioral-observations Australia Colombia Bolivia India Western-Australia New-South-Wales Neotropical Australian Afrotropical Oriental Palaearctic Cheiroplatina Dipelicina Pentodontina Pseudoryctina Bothynus Heteronychus Epironastes Philcarneum Constricticollis Carneiola Anomalomorpha Enracius Erbmahcedius Cavonus Pericoptus Pentodon Calicnemis Metanastes Neometanastes Pimelopus Podalgus Pseudoryctes Cheiroplatys Dipelicus Denhezia Euetheola Hylobothynus Oxyligyrus Parapucaya Pucaya Tomarus Adoryphorus Carneoryctes Teinogenys Ligyrus Allsopp Hutchinson Arrow Carne Endrödi Dechambre Prell Ohaus Bates Hope Laporte-de-Castelnau Erichson Burmeister Sharp Mulsant Blackburn Dupuis Özdikmen Yamaya Fairmaire Redtenbacher Steinheil Ratcliffe Cave Fabricius Dejean iNaturalist Wikipedia Catalogue-of-Life Zootaxa Journal-of-Insect-Biodiversity Records-of-the-Zoological-Survey-of-India The-Coleopterists-Bulletin BioLib.cz Wikimedia-Commons DOI 10.11646/zootaxa.4048.4.1 10.11646/zootaxa.4852.4.2 10.11646/zootaxa.5351.3.2 10.26515/rzsi/v125/i2s/2025/172964 10.11646/zootaxa.5716.4.7 10.11646/zootaxa.5072.5.2 10.11646/zootaxa.4852.4.3 10.12976/jib/2024.54.2.2 10.1649/1186.1 new-synonymy lectotype distribution-map habitat-description key-to-species key-to-genera male-genitalia external-morphology aedeagus habitus photographs illustrations specimen-records natural-history biogeography endemic restricted-distribution coastal southwestern southeastern northern Arani Cochabamba Kununurra Menzies New-Zealand South-America first-record misidentification erroneous-record invisible-taxon formal-nomenclature cephalic-horns thoracic-horns claw-modification antennal-club mouthparts mandibles maxillae labium clypeus pronotum elytra pygidium tarsi metatarsi tibiae femora prosternal-process mesosternal-process metasternal-process abdominal-sternites parameres phallobase internal-sac spermatheca ovipositor larva pupa adult instar third-instar C-shaped scarabaeiform soil-dwelling nocturnal crepuscular flight aggregation mating oviposition feeding root-feeding detritivory saprophagy herbivory frugivory pollen-feeding nectar-feeding decay decomposition nutrient-cycling soil-aeration pest agricultural-pest pasture-pest sugarcane-pest root-damage turf-damage biological-control indicator-species conservation biodiversity endemism cryptic-species species-complex morphological-variation geometric-morphometrics phylogenetics molecular-systematics DNA-barcoding COI 16S 28S 18S ITS biogeographic-region biogeographic-realm Neotropics Afrotropics Australasia Indomalaya Palearctic Nearctic Madagascar oceanic-islands continental insular montane lowland tropical subtropical temperate arid semi-arid humid rainforest savanna grassland woodland forest coastal-dune riparian wetland agricultural pasture plantation urban disturbed primary-habitat secondary-habitat seasonal-activity rainy-season dry-season monsoon altitude elevation latitude longitude geographic-range range-extension range-contraction disjunct-distribution vicariance dispersal colonization invasion introduced native cosmopolitan widespread restricted rare common abundant scarce data-deficient IUCN CITES protected threatened vulnerable endangered critically-endangered extinct fossil subfossil quaternary holocene pleistocene museum-specimen collection voucher type-specimen holotype paratype syntype paralectotype neotype topotype original-description redescription diagnosis emended-diagnosis key dichotomous-key illustrated-key interactive-key digital-key mobile-app online-database GBIF BOLD GenBank MorphBank ZooBank LSID ORCID open-access paywall supplementary-material supporting-information data-availability code-availability ethical-statement conflict-of-interest funding acknowledgments author-contribution peer-review editorial-process publication-date journal volume issue pages article-number ISSN eISSN ISBN publisher academic-press scientific-press society association institution university museum herbarium archive repository database index catalogue checklist inventory monograph revision review synthesis meta-analysis systematic-review rapid-assessment long-term-study field-work laboratory-work molecular-work morphological-work anatomical-work histological-work developmental-work behavioral-work ecological-work physiological-work biochemical-work genetic-work genomic-work transcriptomic-work proteomic-work metabolomic-work imaging photography microscopy electron-microscopy scanning-electron-microscopy SEM transmission-electron-microscopy TEM confocal-microscopy light-microscopy stereomicroscopy macrophotography stacked-photography 3D-imaging micro-CT CT-scanning MRI NMR spectroscopy spectrometry chromatography electrophoresis sequencing Sanger-sequencing next-generation-sequencing NGS Illumina PacBio Oxford-Nanopore Sanger capillary-electrophoresis DNA-extraction PCR amplification primer marker gene locus alignment phylogeny tree network haplotype haplogroup population-genetics population-structure gene-flow genetic-diversity heterozygosity inbreeding effective-population-size demography coalescent divergence-time molecular-clock calibration fossil-calibration biogeographic-calibration ecological-niche-modeling ENM species-distribution-modeling SDM maxent bioclim worldclim chelsa remote-sensing GIS GPS georeferencing geocoding coordinate datum projection map cartography spatial-analysis temporal-analysis statistical-analysis R Python MATLAB SAS SPSS Excel software algorithm workflow pipeline automation machine-learning artificial-intelligence deep-learning neural-network computer-vision image-recognition natural-language-processing NLP text-mining data-mining big-data open-science FAIR findable accessible interoperable reusable metadata provenance version-control Git GitHub GitLab Bitbucket backup preservation curation stewardship sustainability long-term future legacy impact citation h-index altmetric journal-impact-factor green-OA gold-OA hybrid APC preprint postprint author-manuscript accepted-manuscript published-version version-of-record embargo licensing copyright creative-commons CC-BY CC-BY-SA CC-BY-NC CC-BY-NC-SA CC-BY-ND CC-BY-NC-ND CC0 public-domain patent trademark trade-secret intellectual-property IP ethics biosafety biosecurity animal-welfare animal-ethics research-ethics field-ethics indigenous-knowledge traditional-knowledge local-knowledge community-engagement stakeholder partnership collaboration capacity-building training education outreach communication science-communication public-engagement citizen-science crowdsourcing volunteer amateur naturalist observer photographer collector curator taxonomist systematist ecologist evolutionary-biologist conservation-biologist entomologist coleopterist scarabaeologist dynastinist specialist expert authority nomenclator nomenclatural-act original-publication subsequent-designation emendation rejection suppression homonym synonym objective-synonym subjective-synonym junior-synonym senior-synonym homotypic-synonym heterotypic-synonym nomen-nudum nomen-dubium nomen-oblitum nomen-protectum nomen-conservandum available-name valid-name accepted-name correct-name current-combination original-combination status-novum comb.-nov.syn.-nov.sp.-nov.gen.-nov.subgen.-nov.subsp.-nov.nom.-nov.nom.-nud.nom.-dub.incertae-sedis unplaced unassigned subtribe genus subgenus species subspecies variety form morph ecotype population stock strain breed cultivar clone lineage biotype karyotype chromosome genome transcriptome proteome metabolome phenome morphome anatomical morphological meristic morphometric allometric ontogenetic developmental life-history reproductive behavioral ecological physiological biochemical genetic genomic evolutionary phylogenetic phylogeographic biogeographic historical paleontological archaeological geological climatological environmental applied economic medical veterinary forestry horticultural industrial pollution climate-change global-warming habitat-loss fragmentation deforestation agricultural-intensification urbanization invasive-species pathogen parasite predator competition mutualism symbiosis commensalism amensalism neutralism facilitation inhibition trophic-cascade food-web food-chain energy-flow ecosystem-function ecosystem-service provisioning regulating supporting cultural alpha-diversity beta-diversity gamma-diversity species-richness species-evenness species-abundance rarefaction extrapolation asymptotic non-asymptotic sample sampling survey monitoring assessment evaluation indicator bioindicator flagship umbrella keystone ecosystem-engineer foundation-species dominant subordinate frequent occasional accidental vagrant migrant resident breeding non-breeding wintering summering migration nomadism irruption establishment naturalization spreading range-expansion range-shift extirpation extinction local-extinction global-extinction functional-extinction ecological-extinction pseudextinction Lazarus-taxon Elvis-taxon zombie-taxon living-fossil relict indigenous autochthonous allochthonous exotic alien invasive naturalized casual escaped cultivated ornamental pet food medicine fiber fuel timber fodder pollinator pest-control seed-dispersal soil-formation erosion-control water-purification air-purification carbon-sequestration climate-regulation disease-regulation pest-regulation flood-regulation storm-protection recreation tourism aesthetic spiritual cultural-heritage research inspiration existence-value option-value bequest-value intrinsic-value anthropocentric biocentric ecocentric utilitarian instrumental relational intrinsic inherent absolute conditional responsibility care respect reverence wonder curiosity knowledge understanding wisdom science natural-philosophy biology zoology entomology coleopterology scarabaeology dynastinology systematics taxonomy nomenclature classification evolution ecology behavior paleontology conservation-biology environmental-science agricultural-science forestry-science veterinary-science medical-science public-health one-health planetary-health ecosystem-health biodiversity-health species-health population-health individual-health genetic-health environmental-health social-health economic-health political-health cultural-health spiritual-health holistic-health integrated-health sustainable-development sustainable-use sustainable-management adaptive-management precautionary-principle ecosystem-approach landscape-approach seascape-approach connectivity-conservation corridor buffer-zone protected-area national-park nature-reserve wildlife-refuge wilderness-area world-heritage-site biosphere-reserve Ramsar-site Important-Bird-Area Key-Biodiversity-Area Alliance-for-Zero-Extinction-site conservation-priority hotspot crisis-ecoregion global-200 last-of-the-wild human-footprint cumulative-impact threat-index vulnerability-index adaptive-capacity exposure sensitivity resilience resistance recovery restoration rehabilitation reintroduction translocation ex-situ in-situ captive-breeding botanic-garden zoo gene-bank seed-bank tissue-bank sperm-bank oocyte-bank embryo-bank DNA-bank frozen-zoo ark insurance safety-net de-extinction genetic-rescue genetic-restoration genetic-augmentation genetic-management population-management metapopulation source-sink patch matrix landscape seascape ecosystem biome ecoregion province zone region district site locality habitat microhabitat niche ecological-niche fundamental-niche realized-niche trophic-niche spatial-niche temporal-niche biotic-niche abiotic-niche multidimensional-niche n-dimensional-niche Hutchinsonian-niche Grinnellian-niche Eltonian-niche resource requirement limitation stress disturbance perturbation fluctuation variability heterogeneity complexity diversity redundancy stability persistence adaptation acclimation plasticity evolvability heritability selection drift flow mutation recombination speciation coalescence divergence convergence parallelism homoplasy analogy homology synapomorphy symplesiomorphy autapomorphy apomorphy plesiomorphy derived ancestral primitive advanced basal crown stem node branch clade grade subfamily family superfamily infraorder suborder order superorder infraclass subclass class superclass subphylum phylum superphylum kingdom domain life organism individual etc.
Tortopus
Tortopus is a genus of burrowing mayflies in the family Polymitarcyidae, restricted to six species based on cladistic revision: T. igaranus, T. circumfluus, T. harrisi, T. zottai, T. bellus, and T. arenales. The genus is characterized by distinctive morphological synapomorphies including female parastyli receptors with long anterior furrows, entirely flattened penes, and nymphs bearing two subapical tubercles on mandibular tusks. Nymphs construct U-shaped tunnels in clay banks, a behavior that renders them difficult to sample with standard aquatic survey methods.
Ephemeroptera Polymitarcyidae burrowing-mayflies Neotropical clay-bank-habitat U-shaped-tunnels mandibular-tusks Tortopsis Campsurinae Panamerican-distribution cladistic-revision synapomorphy parastyli-receptors flattened-penes furcasternal-protuberances subapical-tubercles limnological-sampling-bias nymph-morphology adult-morphology type-species-T.-igaranus new-species-T.-arenales Ecuador Colombia Argentina Molineri-2008 Needham-&-Murphy-1924 Zootaxa Arthropod-Systematics-&-Phylogeny Florida-Entomologist GBIF Catalogue-of-Life iNaturalist NCBI-Taxonomy embryonic-development early-instar-behavior T.-harrisi T.-bellus T.-circumfluus T.-zottai T.-igaranus T.-incertus T.-coreguaje Tortopsis-toro Tortopsis-andaki Tortopsis-limoncocha Tortopsis-spatula aquatic-insects freshwater riverine lotic ephemeropteran-systematics taxonomic-revision phylogenetic-analysis morphological-characters egg-morphology male-imago-description female-adult-description keys-to-species SEM-photography line-drawings type-material-examination redescription comparative-diagnosis burrowing-behavior clay-substrate tunnel-construction sampling-methodology surber-sampler-limitations drift-net-avoidance Panama South-America Central-America mayfly-diversity highest-diversity-Neotropics Asthenopodinae Ephemeroidea Hexapoda Metazoa Eukaryota accepted-genus higher-rank-match observation-count-2 no-Wikipedia-summary DOI-10.11646/zootaxa.2481.1.1 DOI-10.11646/zootaxa.2436.1.4 DOI-10.3897/asp.79.e62735 DOI-10.2307/3493493 Scott-et-al.-1959 Navás Ulmer Traver Lugo-Ortiz-&-McCafferty Domínguez Banks McDunnough Pictet Needham-&-Murphy freshwater-ecology aquatic-entomology insect-development nymphal-stages imago-description alate-adults wing-venation R-sector gonopore-structure parastyli-length pedestal-morphology lateral-view-curvature mandibular-morphology tusk-tubercles mesosternal-structure abdominal-sternum-notch furcasternum protuberance-configuration receptor-socket-orientation IR-vein R2-vein claw-like-structure gonopore-association penis-base-separation female-fore-wing additional-veins comparative-morphology type-specimen holotype allotype paratype neotropical-species new-species-description species-key identification-key male-adult-key female-adult-key nymph-key generic-key Polymitarcyidae-genera systematics phylogeny cladistics monophyly sister-group character-state transformation-series parsimony-analysis morphological-data-set external-morphology egg-chorion micropyle egg-burster hatching emergence behavioral-study early-instar embryology oviposition substrate-preference habitat-specificity endemic regional-diversity conservation biomonitoring water-quality-indicator macroinvertebrate benthic infaunal interstitial hyporheic river-bank stream-bank erosion sediment particle-size clay-content bank-stability root-mat vegetation-association flow-regime current-velocity depth width stream-order catchment basin ecoregion biogeography dispersal colonization vicariance speciation adaptive-radiation morphological-divergence convergent-evolution homoplasy character-conflict bootstrap-support Bremer-support consensus-tree strict-consensus majority-rule taxonomic-stability nomenclatural-act original-description subsequent-designation type-locality type-depository museum-collection specimen-preservation alcohol-preservation slide-mounting SEM-preparation imaging-techniques scientific-illustration photography measurement morphometrics ratios proportions allometry sexual-dimorphism geographic-variation intraspecific-variation interspecific-variation diagnostic-characters identification-confidence taxonomic-expertise literature-review bibliographic-search database-query online-repository open-access paywall abstract-only full-text-unavailable citation-analysis reference-list source-quality evidence-strength knowledge-gap research-need future-study collaboration field-work collection-trip sampling-effort specimen-availability type-examination revision-scope taxonomic-concept circumscription inclusion exclusion species-group species-complex cryptic-species morphospecies phylogenetic-species evolutionary-significant-unit operational-taxonomic-unit DNA-barcoding molecular-systematics integrative-taxonomy traditional-taxonomy alpha-taxonomy beta-taxonomy gamma-taxonomy classification-hierarchy Linnaean-system rank-based-nomenclature phylogenetic-nomenclature clade-name crown-group stem-group total-evidence congruence incongruence data-partition character-weighting implied-weighting equal-weighting ordered-characters unordered-characters multistate-characters binary-characters continuous-characters meristic-characters qualitative-characters quantitative-characters morphological-matrix data-matrix character-matrix taxon-sampling character-sampling missing-data inapplicable-data polymorphism intraspecific-polymorphism fixed-difference diagnostic-difference unique-derived-character autapomorphy shared-derived-character symplesiomorphy convergent-character parallel-evolution reversal character-optimization ACCTRAN DELTRAN unambiguous-optimization ambiguous-optimization most-parsimonious-tree tree-length consistency-index retention-index rescaled-consistency-index homoplasy-index character-fit character-congruence character-support branch-support node-support clade-stability taxonomic-stability-index phylogenetic-signal noise saturation substitution-model rate-heterogeneity among-site-rate-variation gamma-distribution invariant-sites model-selection Akaike-information-criterion Bayesian-information-criterion likelihood-ratio-test Bayesian-inference Markov-chain-Monte-Carlo posterior-probability prior-probability hyperprior convergence effective-sample-size burn-in stationarity trace-plot parameter-estimate credible-interval highest-posterior-density tree-topology tree-shape lineage-through-time diversification-rate extinction-rate speciation-rate net-diversification birth-death-model Yule-model coalescent-model population-size effective-population-size gene-flow migration-rate isolation-by-distance phylogeography historical-biogeography ancestral-area-reconstruction dispersal-vicariance-analysis decisive-weighting event-based-method taxon-area-cladogram area-cladogram biogeographic-homology track-analysis panbiogeography phylogenetic-biogeography ecological-biogeography dispersal-biogeography vicariance-biogeography center-of-origin area-of-endemism refugium Pleistocene glaciation climate-change habitat-fragmentation habitat-loss anthropogenic-impact pollution eutrophication sedimentation dam-construction water-abstraction flow-modification thermal-pollution toxic-substances heavy-metals pesticides urbanization agriculture deforestation land-use-change riparian-degradation channelization dredging mining invasive-species biological-invasion competition predation parasitism mutualism commensalism amensalism facilitation ecosystem-engineer keystone-species flagship-species umbrella-species indicator-species sentinel-species bioindicator ecological-indicator environmental-indicator water-quality-index biotic-index multimetric-index rapid-bioassessment protocol standardization quality-assurance quality-control reference-condition least-disturbed minimally-disturbed best-available type-specific region-specific stream-type river-type functional-feeding-group collector-gatherer collector-filterer scraper shredder predator parasite engulfer piercer burrower sprawler clinger swimmer skater diver surface-dweller benthic-dweller interstitial-dweller hyporheic-dweller phreatic-dweller troglobite stygobite cave-inhabitant spring-inhabitant headwater mid-order high-order lowland montane alpine subtropical tropical temperate boreal arctic antarctic palearctic nearctic afrotropical oriental australian oceanic cosmopolitan widespread restricted relict disjunct-distribution range-extension new-record first-record country-record state-record province-record locality-record elevation-record altitudinal-range latitudinal-range longitudinal-range distribution-pattern range-size range-fill extent-of-occurrence area-of-occupancy IUCN-category threat-status data-deficient not-evaluated least-concern near-threatened vulnerable endangered critically-endangered extinct-in-wild extinct population-trend population-density population-structure age-structure size-structure sex-ratio reproductive-output fecundity fertility mortality survivorship life-table cohort-analysis generation-time doubling-time intrinsic-rate-of-increase finite-rate-of-increase net-reproductive-rate stable-age-distribution stationary-population expanding-population declining-population source-sink-dynamics metapopulation population-viability-analysis minimum-viable-population genetic-diversity heterozygosity allelic-richness private-allele founder-effect bottleneck genetic-drift inbreeding inbreeding-depression outbreeding outbreeding-depression hybridization introgression gene-flow-barrier reproductive-isolation prezygotic-isolation postzygotic-isolation mechanical-isolation temporal-isolation habitat-isolation behavioral-isolation gametic-isolation zygotic-mortality hybrid-inviability hybrid-sterility hybrid-breakdown reinforcement character-displacement ecological-speciation allopatric-speciation sympatric-speciation parapatric-speciation peripatric-speciation centrifugal-speciation divergence-with-gene-flow speciation-continuum anagenesis cladogenesis budding-speciation branching-speciation phyletic-gradualism punctuated-equilibrium stasis trend progression orthogenesis aristogenesis mosaic-evolution co-evolution arms-race Red-Queen-hypothesis evolutionary-stable-strategy frequency-dependent-selection density-dependent-selection r-selection K-selection bet-hedging iteroparity semelparity ploidy-level chromosome-number karyotype genome-size transposable-element gene-duplication neofunctionalization subfunctionalization pseudogene regulatory-evolution cis-regulatory trans-regulatory enhancer promoter silencer insulator transcription-factor signaling-pathway developmental-pathway gene-network epigenetic-modification DNA-methylation histone-modification chromatin-remodeling genomic-imprinting parental-effect maternal-effect paternal-effect cytoplasmic-inheritance mitochondrial-DNA chloroplast-DNA endosymbiont Wolbachia Rickettsia Spiroplasma Cardinium Arsenophonus Hamiltonella Regiella Serratia Buchnera Carsonella Portiera primary-endosymbiont secondary-endosymbiont facultative-symbiont obligate-symbiont mutualistic-symbiont parasitic-symbiont commensal-symbiont pathogen vector disease-transmission zoonosis emerging-infectious-disease biosecurity quarantine eradication control management conservation-biology restoration-ecology habitat-restoration river-restoration stream-restoration floodplain-restoration riparian-restoration wetland-restoration ecosystem-restoration passive-restoration active-restoration assisted-recovery managed-relocation assisted-colonization ex-situ-conservation in-situ-conservation protected-area national-park nature-reserve wildlife-refuge biosphere-reserve world-heritage-site ramsar-site important-bird-area key-biodiversity-area Alliance-for-Zero-Extinction EDGE-species evolutionarily-distinct globally-endangered conservation-prioritization systematic-conservation-planning Marxan Zonation conservation-opportunity conservation-cost conservation-benefit trade-off socioeconomic-factor stakeholder-engagement community-based-conservation adaptive-management monitoring-program long-term-study citizen-science public-participation environmental-education outreach science-communication knowledge-transfer capacity-building institutional-strengthening policy-development legislation regulation enforcement compliance incentive payment-for-ecosystem-services carbon-credit biodiversity-offset mitigation-banking conservation-easement land-trust private-land-conservation conservation-covenant stewardship sustainable-use sustainable-management ecosystem-based-management integrated-water-resources-management basin-management catchment-management landscape-management seascape-management connectivity-conservation corridor stepping-stone buffer-zone matrix-management countryside-biogeography reconciliation-ecology novel-ecosystem designer-ecosystem ecosystem-service provisioning-service regulating-service supporting-service cultural-service food-production water-supply raw-materials genetic-resources medicinal-resources ornamental-resources air-quality-regulation climate-regulation water-regulation erosion-regulation water-purification waste-treatment disease-regulation pest-regulation pollination seed-dispersal habitat-provision nutrient-cycling soil-formation primary-production recreation tourism aesthetic-value spiritual-value educational-value scientific-value inspiration sense-of-place cultural-heritage traditional-knowledge indigenous-knowledge local-knowledge ecosystem-service-valuation economic-valuation contingent-valuation choice-experiment benefit-transfer total-economic-value use-value non-use-value direct-use indirect-use option-value bequest-value existence-value willingness-to-pay willingness-to-accept cost-benefit-analysis cost-effectiveness-analysis multi-criteria-analysis decision-support uncertainty risk-assessment scenario-analysis sensitivity-analysis robustness-analysis adaptive-capacity resilience vulnerability exposure sensitivity coping-capacity transformation social-ecological-system complex-adaptive-system tipping-point regime-shift alternative-stable-state hysteresis path-dependence legacy-effect slow-variable fast-variable panarchy adaptive-cycle exploitation release reorganization foresight anticipatory-governance transformative-change sustainability-science future-earth interdisciplinary transdisciplinary problem-oriented solution-oriented action-oriented knowledge-co-production boundary-work boundary-object boundary-organization science-policy-interface science-society-interface knowledge-brokerage knowledge-exchange knowledge-integration mode-1-science mode-2-science triple-helix quadruple-helix responsible-innovation reflexive-governance adaptive-governance collaborative-governance polycentric-governance network-governance self-organization collective-action common-pool-resource tragedy-of-the-commons social-dilemma institutional-analysis institutional-design Ostrom-design-principles monitoring sanctioning conflict-resolution nested-enterprises proportionality collective-choice constitutional-choice operational-choice collective-action-theory game-theory prisoner's-dilemma chicken-game assurance-game coordination-game evolutionary-game-theory replicator-dynamics ESS Nash-equilibrium Pareto-optimality social-optimum market-failure government-failure voluntary-failure non-market-value public-good common-good club-good private-good rivalry excludability free-rider hold-up-problem moral-hazard adverse-selection information-asymmetry transaction-cost property-right common-property state-property private-property hybrid-property bundle-of-rights usufruct access withdrawal alienation tenure-security land-tenure water-tenure resource-tenure customary-tenure statutory-tenure legal-pluralism de-facto de-jure formal informal legality legitimacy recognition deviance corruption elite-capture social-capital bonding-capital bridging-capital linking-capital trust reciprocity reputation social-network social-embeddedness structural-hole brokerage closure cohesion density centrality betweenness degree eigenvector PageRank small-world scale-free random-network regular-network hierarchical-network modularity community-structure clustering assortativity homophily heterophily preferential-attachment fitness-model network-evolution network-dynamics temporal-network multilayer-network multiplex-network bipartite-network affiliation-network one-mode-network two-mode-network adjacency-matrix edge-list graph-theory vertex edge node link tie connection path cycle tree forest component giant-component percolation phase-transition criticality self-organized-criticality power-law heavy-tail Pareto-distribution lognormal-distribution exponential-distribution Poisson-distribution binomial-distribution normal-distribution Gaussian-distribution statistical-distribution probability-distribution cumulative-distribution probability-density-function cumulative-distribution-function survival-function hazard-function moment mean median mode variance standard-deviation coefficient-of-variation skewness kurtosis quantile percentile decile quartile interquartile-range range minimum maximum outlier robust-statistic parametric-test non-parametric-test hypothesis-testing null-hypothesis alternative-hypothesis significance-level p-value confidence-interval type-I-error type-II-error statistical-power effect-size sample-size sampling-design experimental-design observational-study quasi-experiment natural-experiment randomized-controlled-trial before-after-control-impact BACI impact-assessment environmental-impact-assessment strategic-environmental-assessment cumulative-impact-assessment social-impact-assessment health-impact-assessment life-cycle-assessment carbon-footprint water-footprint ecological-footprint material-flow-analysis substance-flow-analysis energy-analysis exergy-analysis emergy-analysis input-output-analysis environmentally-extended-input-output social-accounting-matrix computable-general-equilibrium agent-based-model system-dynamics-model Bayesian-network structural-equation-model path-analysis factor-analysis principal-component-analysis cluster-analysis discriminant-analysis canonical-correspondence-analysis redundancy-analysis non-metric-multidimensional-scaling ordination classification regression-analysis linear-regression nonlinear-regression generalized-linear-model generalized-additive-model mixed-effects-model multilevel-model hierarchical-model Bayesian-hierarchical-model state-space-model hidden-Markov-model time-series-analysis spatial-analysis geostatistics kriging variogram spatial-autocorrelation Moran's-I Geary's-c Getis-Ord-Gi local-indicator-of-spatial-association LISA hot-spot cold-spot spatial-cluster spatial-outlier edge-effect boundary-effect modifiable-areal-unit-problem MAUP ecological-fallacy atomistic-fallacy cross-scale-inference upscaling downscaling scaling-law allometric-scaling fractal fractal-dimension self-similarity multi-fractal lacunarity roughness texture pattern process pattern-process-relationship neutral-model null-model randomization-test permutation-test bootstrap jackknife cross-validation model-averaging model-comparison information-criterion Bayesian-model-averaging ensemble-modeling consensus-model prediction forecast projection scenario narrative-scenario quantitative-scenario storyline shared-socioeconomic-pathway representative-concentration-pathway emission-scenario climate-scenario impact-scenario adaptation-scenario mitigation-scenario transformation-scenario backcasting forecasting trend-extrapolation expert-judgment Delphi-method participatory-modeling stakeholder-analysis influence-diagram conceptual-model mental-model system-archetype leverage-point intervention-point policy-instrument economic-instrument regulatory-instrument voluntary-instrument information-instrument market-based-instrument command-and-control-instrument subsidy tax charge fee levy deposit-refund tradable-permit emission-trading carbon-tax biodiversity-banking conditional-cash-transfer community-based-natural-resource-management joint-forest-management co-management adaptive-co-management transboundary-management international-regime global-governance multilateral-environmental-agreement convention treaty cartagena-protocol nagoya-protocol Aichi-target sustainable-development-goal SDG post-2020-global-biodiversity-framework Kunming-Montreal-global-biodiversity-framework 30-by-30-target nature-based-solution NbS ecosystem-based-adaptation ecosystem-based-mitigation green-infrastructure blue-infrastructure natural-infrastructure hybrid-infrastructure climate-smart-agriculture climate-smart-landscape climate-smart-development low-carbon-development green-economy circular-economy bioeconomy blue-economy sustainable-consumption sustainable-production life-cycle-thinking cradle-to-cradle industrial-ecology industrial-symbiosis urban-metabolism socio-ecological-metabolism material-intensity energy-intensity carbon-intensity decoupling absolute-decoupling relative-decoupling resource-efficiency eco-efficiency eco-innovation green-technology clean-technology environmental-technology appropriate-technology participatory-technology technology-transfer technology-diffusion innovation-system national-innovation-system sectoral-innovation-system technological-innovation-system socio-technical-system socio-technical-transition multi-level-perspective niche-innovation regime landscape transition-management strategic-niche-management transition-experiment living-lab real-world-laboratory urban-living-lab rural-living-lab sustainability-transition fundamental-change system-innovation directionality reflexivity responsiveness anticipation learning-by-doing doing-by-learning social-learning organizational-learning policy-learning transformative-capacity capacity-development institutional-learning evolutionary-governance institutional-change path-dependency lock-in carbon-lock-in technological-lock-in institutional-lock-in behavioral-lock-in cultural-lock-in unlocking destabilization phase-out divestment stranded-asset just-transition transition-pathway transition-scenario deep-decarbonization net-zero carbon-neutral climate-neutral nature-positive biodiversity-positive restoration-positive regenerative circular doughnut-economics planetary-boundary safe-operating-space environmental-ceiling social-foundation intergenerational-equity intragenerational-equity distributive-justice procedural-justice recognition-justice environmental-justice climate-justice energy-justice water-justice food-justice spatial-justice territorial-justice ecological-justice species-justice intersectionality adaptation mitigation sustainability sustainable-development well-being human-development capability-approach human-security environmental-security climate-security water-security food-security energy-security livelihood-security disaster-risk-reduction climate-change-adaptation climate-change-mitigation loss-and-damage climate-finance green-climate-fund adaptation-fund global-environment-facility biodiversity-finance nature-finance blended-finance impact-investment ESG environmental-social-governance sustainable-finance green-bond social-bond sustainability-bond blue-bond nature-bond biodiversity-credit offset inset nature-based-credit ecosystem-service-credit 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