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Cyphomyrmex
fungus-growing ants
Cyphomyrmex is a genus of small, drab-colored fungus-growing ants in the tribe Attini, found primarily in the Neotropics. These ants cultivate fungi in the tribe Leucocoprineae as their primary food source, with most species growing fungal nodules rather than full mycelial gardens. Colonies are typically monogynous and small, rarely exceeding 500 workers. The genus is divided into two species complexes: the strigatus complex (South America only) and the rimosus complex (southern North America to South America). Cyphomyrmex represents a basal lineage among attine ants and serves as sister group to Mycetophylax.
fungus-growing-ants attine-ants Myrmicinae Neotropical mutualism fungiculture Leucocoprineae monogyny parasitoid-wasps social-parasitism chemical-ecology alkaloid-venom nest-architecture behavioral-plasticity cytogenetics chromosome-evolution Guiana-Shield Brazil Panama Puerto-Rico Mesoamerica Argentina United-States Texas California Florida yeast-gardens mycelial-gardens rimosus-complex strigatus-complex age-polyethism trophallaxis thanatosis feigning-death Diapriidae Acanthopria Mimopriella Megalomyrmex social-parasite C.-longiscapus C.-transversus C.-laevigatus C.-rimosus C.-minutus C.-lectus C.-cornutus C.-costatus pheromones 3-octanol farnesenes trail-following alarm-signal diketopiperazines antifungal-compounds defensive-behavior colony-defense flight-response recruitment Lanchester's-laws karyotype chromosome-number rDNA microsatellites FISH chamber-depth gallery-length fire-disturbance savanna-forest-transition arboreal-nesting clay-auricles soil-nests wood-nests epiphytic-nests moss-nests coconut-nests desiccation-susceptibility moist-habitats basal-attine phylogenetics sister-group chemical-evolution venom-alkaloids behavioral-manipulation host-parasite-interaction ecosystem-engineering nutrient-cycling decomposition organic-matter insect-feces exoskeleton-substrate garden-transplantation worker-polymorphism queen-number dealate-queens alates brood-care ecdysis-assistance foraging sugar-collection plant-exudates fungal-domestication nodule-cultivation mycelium-cultivation agricultural-behavior nest-entrance funnel-structure exposed-gardens symbiont-diversity geographic-variation research-model evolutionary-biology mutualism-evolution ant-fungus-coevolution chemical-communication social-insects eusociality colony-size small-colonies slow-movement crypsis debris-mimicry predation-avoidance parasitoid-defense wasp-killing colony-abandonment resource-rescue thief-ant-defense venom-resistance toxicity-tolerance behavioral-suppression playing-dead alkaloid-manipulation fungal-secondary-metabolites antibacterial-compounds antiviral-compounds garden-protection microbial-defense nest-microclimate temperature-regulation humidity-requirements depth-effects fire-adaptation disturbance-ecology habitat-specificity substrate-specialization clay-construction soil-manipulation architectural-plasticity vertical-nesting embankment-nesting overhang-nesting pendant-nests swallow-nest-mimicry research-accessibility field-biology tropical-ecology Neotropical-myrmecology biodiversity conservation invasive-species-context native-species biological-control Integrated-Pest-Management agricultural-ecology forest-ecology savanna-ecology transitional-habitats disturbed-areas urban-ecology anthropogenic-impact climate-sensitivity habitat-limitation range-boundaries dispersal colonization establishment population-dynamics demography colony-density nest-spacing intraspecific-competition interspecific-competition territoriality resource-defense combat-behavior aggression-levels recruitment-behavior chemical-signaling alarm-pheromones trail-pheromones recognition-cues cuticular-hydrocarbons venom-composition defensive-secretions mandibular-glands Dufour's-gland metapleural-glands antibiotic-production grooming-behavior fungal-inoculation integumental-fungi yeast-growth larval-grooming garden-maintenance substrate-preparation fecal-fertilizer anal-trophallaxis crop-regurgitation fluid-application drying-behavior nodule-formation transplantation-behavior garden-establishment ceiling-gardens root-association architectural-innovation nest-site-selection microhabitat-choice moisture-requirements thermal-buffering predator-avoidance parasite-avoidance disease-resistance social-immunity collective-behavior division-of-labor task-allocation temporal-polyethism worker-age nurse-workers forager-workers soldier-caste-absence monomorphism worker-size-variation queen-worker-differentiation reproductive-caste gyne-production male-production seasonal-reproduction colony-founding claustral-founding non-claustral-founding fungal-inoculum founding-queen first-worker-generation nutritional-dependency fat-reserves metabolic-adaptation digestive-physiology enzyme-complement fungal-enzyme lignocellulose-degradation nutritional-mutualism obligate-mutualism vertical-transmission horizontal-transmission symbiont-switching cultivar-replacement garden-failure colony-mortality population-turnover generation-time longevity survivorship life-expectancy aging senescence reproductive-investment colony-growth colony-decline budding fission reproduction-mode genetic-structure relatedness mating-system nuptial-flight mate-choice sperm-storage colony-odor nestmate-recognition foreign-rejection agonistic-behavior territorial-marking boundary-defense raiding-behavior robbing-behavior cleptobiosis dulosis slavery inquilinism guest-ants myrmecophiles nest-associates commensalism inquilines parasite-tolerance host-manipulation venom-alkaloid pyrrolidine indolizidine piperidine nonanal farnesene sesquiterpene agriculture-chemical gaster-extract chemical-diversity chemical-simplicity evolutionary-precursor derived-chemistry biosynthetic-pathway semiochemical infochemical communication information-transfer colony-organization social-structure polygyny queen-condition dealation wing-shedding ovarian-development fecundity egg-laying brood-development developmental-rate temperature-dependence humidity-dependence food-availability nutritional-quality fungal-productivity garden-yield worker-production colony-efficiency energy-allocation resource-allocation foraging-efficiency search-behavior prey-detection food-retrieval transport-behavior load-size travel-speed path-integration orientation navigation homing-behavior learning memory experience individual-recognition context-dependent-behavior threat-assessment risk-management predator-inspection alarm-propagation recruitment-activation mobilization colony-response emergency-behavior panic-response controlled-response defensive-strategy offensive-strategy escalation de-escalation conflict-resolution fight-avoidance retreat-behavior surrender-signal dominance-hierarchy reproductive-skew worker-policing queen-policing egg-eating oviposition-suppression reproductive-conflict kin-conflict worker-reproduction male-production-by-workers anarchy social-parasitism-detection parasite-recognition enemy-recognition nest-intrusion nest-defense perimeter-defense chamber-defense brood-defense queen-defense self-sacrifice suicidal-defense autothysis sting-use mandible-use spraying-behavior gaster-flexion chemical-defense mechanical-defense armor cuticle-thickness pigmentation masquerade mimicry Batesian-mimicry Muellerian-mimicry aggressive-mimicry Wasmannian-mimicry myrmecophily myrmecophagy ant-plant-mutualism trophobiosis extrafloral-nectary domatia plant-ant-interaction herbivore-defense seed-dispersal pollination ecosystem-service functional-role trophic-position detritivore decomposer saprotroph necrotroph fungivore omnivore predator scavenger trophic-level energy-flow carbon-cycling nitrogen-cycling phosphorus-cycling soil-formation bioturbation aeration water-infiltration microbial-community bacteria actinomycetes Pseudonocardia disease-suppression garden-hygiene weeding-behavior pruning-behavior fungal-grooming antibiotic-application mutualist-protection agricultural-disease crop-pathology escovopsis garden-parasite competitor-fungus pathogen-defense immune-response individual-immunity grooming allogrooming self-grooming nest-cleaning refuse-management fecal-management waste-disposal midden-formation territory-size home-range foraging-range recruitment-range exploration exploitation information-use public-information private-information cue-use signal-use communication-network interaction-network social-network colony-network population-network community-ecology species-interaction apparent-competition exploitative-competition interference-competition resource-partitioning niche-differentiation habitat-filtering environmental-gradient altitudinal-distribution latitudinal-distribution continental-distribution island-distribution endemism biogeography phylogeography historical-biogeography vicariance range-expansion range-contraction refugia glacial-cycles climate-change-impact habitat-fragmentation deforestation land-use-change agricultural-intensification urbanization pollution pesticide-exposure heavy-metal-accumulation conservation-status population-viability extinction-risk protected-area reserve-design habitat-corridor connectivity metapopulation source-sink-dynamics rescue-effect mass-effect ecological-theory behavioral-ecology evolutionary-ecology functional-ecology trait-based-ecology comparative-analysis phylogenetic-comparative-methods character-evolution ancestral-state-reconstruction convergent-evolution parallel-evolution adaptive-radiation species-diversification speciation hybridization introgression gene-flow genetic-differentiation population-structure molecular-ecology genomics transcriptomics proteomics metabolomics volatile-organic-compound cuticular-hydrocarbon glandular-secretion sting-apparatus mandible-morphology head-morphology mesosomal-morphology petiolar-morphology gastral-morphology leg-morphology antennal-morphology sensilla chemoreception mechanoreception thermoreception photoreception vision compound-eye ocelli brain-morphology neuroanatomy learning-capacity memory-capacity cognitive-ability problem-solving tool-use sequence-learning route-learning spatial-memory temporal-memory social-learning cultural-transmission tradition colony-personality behavioral-syndrome boldness exploration-tendency activity-level aggression-level foraging-tendency defensive-tendency stress-response physiological-stress thermal-stress desiccation-stress starvation-stress immune-challenge wound-healing metabolic-rate respiration energy-expenditure water-balance osmoregulation excretion Malpighian-tubules rectum hindgut midgut foregut crop proventriculus digestive-enzyme nutrient-absorption lipid-metabolism carbohydrate-metabolism protein-metabolism amino-acid-metabolism nitrogen-excretion uric-acid urea ammonia osmolyte cryoprotectant heat-shock-protein stress-protein antioxidant-defense oxidative-stress reactive-oxygen-species aging-biology senescence-theory disposable-soma antagonistic-pleiotropy mutation-accumulation life-history-theory r-K-selection bet-hedging iteroparity semelparity reproductive-effort parental-investment maternal-effect paternal-effect epigenetics DNA-methylation histone-modification non-coding-RNA gene-regulation transcription-factor signaling-pathway hormone juvenile-hormone ecdysone insulin-signaling TOR-signaling nutrient-sensing reproductive-signaling caste-determination sex-determination environmental-sex-determination genetic-sex-determination haplodiploidy arrhenotoky thelytoky parthenogenesis arrhenotokous-parthenogenesis automixis apomixis central-fusion terminal-fusion genomic-imprinting parental-conflict kin-selection inclusive-fitness Hamilton's-rule relatedness-asymmetry sex-ratio-theory local-mate-competition local-resource-competition local-resource-enhancement split-sex-ratio worker-sex-ratio queen-sex-ratio colony-sex-ratio population-sex-ratio operational-sex-ratio effective-population-size genetic-drift inbreeding inbreeding-depression outbreeding outbreeding-depression heterosis hybrid-vigor genetic-load mutation-rate effective-number-of-codons codon-usage-bias GC-content genome-size karyotype-evolution chromosomal-rearrangement fusion inversion translocation polyploidy aneuploidy chromosomal-polymorphism rDNA-cluster heterochromatin euchromatin centromere telomere nucleolus-organizer-region satellite-DNA minisatellite microsatellite simple-sequence-repeat transposable-element retrotransposon DNA-transposon horizontal-gene-transfer endosymbiont Wolbachia Cardinium Rickettsia Spiroplasma bacterial-symbiont fungal-symbiont viral-symbiont parasite pathogen disease epidemiology transmission-dynamics virulence resistance tolerance recovery immunity vaccination prophylaxis medicinal-behavior self-medication thermoregulation behavioral-thermoregulation social-thermoregulation microclimate-selection nest-temperature brood-temperature fungal-garden-temperature metabolic-heat heat-dissipation evaporative-cooling behavioral-cooling fanning ventilation chamber-design gallery-design entrance-design air-flow humidity-regulation water-collection water-storage desiccation-resistance cuticular-waterproofing spiracle-control respiratory-water-loss excretory-water-loss water-recycling metabolic-water dietary-water environmental-water rainfall-dependence seasonal-activity dry-season wet-season monsoon drought flood fire disturbance succession primary-succession secondary-succession climax-community pioneer-species habitat-specialist habitat-generalist edge-species interior-species gap-species canopy-species understory-species ground-species subterranean-species arboreal-species liana-species epiphyte-species lithophyte-species rheophyte-species aquatic-margin-species terrestrial-species fossorial-species cursorial-species scansorial-species volant-species winged-reproductive dealate ergatoid brachypterous apterous polymorphism caste-polymorphism queen-polymorphism male-polymorphism size-polymorphism allometry isometry shape-variation morphological-integration modularity evolvability constraints developmental-stability fluctuating-asymmetry directional-asymmetry antisymmetry phenotypic-plasticity reaction-norm genotype-environment-interaction grandparental-effect indirect-genetic-effect social-environment-effect colony-effect density-dependence frequency-dependence negative-frequency-dependence positive-frequency-dependence apostatic-selection anti-apostatic-selection search-image predator-learning prey-learning friend-recognition cuticular-hydrocarbon-profile gestalt-odor colony-specific-odor individual-specific-odor task-specific-odor age-specific-odor reproductive-odor queen-odor queen-pheromone primer-pheromone releaser-pheromone signal-pheromone modulatory-pheromone alarm-pheromone trail-pheromone recruitment-pheromone aggregation-pheromone sex-pheromone marking-pheromone territorial-pheromone defensive-pheromone poison-gland Dufour-gland mandibular-gland postpharyngeal-gland metapleural-gland sternal-gland tarsal-gland anal-gland rectal-gland pygidial-gland mandibular-brush tibial-spur stridulatory-organ subgenual-organ Johnston's-organ tympanal-organ hair-sensillum campaniform-sensillum basiconic-sensillum coeloconic-sensillum placoid-sensillum sensilla-trichodea sensilla-basiconica sensilla-coeloconica sensilla-ampullacea sensilla-campaniformia sensilla-placodea olfactory-receptor-neuron gustatory-receptor-neuron mechanoreceptor-neuron thermoreceptor-neuron hygroreceptor-neuron photoreceptor-neuron visual-processing olfactory-processing gustatory-processing mechanosensory-processing central-complex mushroom-body antennal-lobe optic-lobe suboesophageal-ganglion ventral-nerve-cord sympathetic-nervous-system neurosecretion neurohemal-organ corpus-cardiacum corpus-allatum prothoracic-gland ventral-gland pericardial-gland neuroendocrine-regulation hormonal-regulation pheromonal-regulation neural-regulation behavioral-regulation social-regulation environmental-regulation developmental-regulation reproductive-regulation metabolic-regulation homeostatic-regulation allostatic-regulation stress-regulation immune-regulation circadian-rhythm circannual-rhythm photoperiodism thermoperiodism biological-clock zeitgeber entrainment free-running-period phase-response-curve masking-effect arousal-threshold sleep torpor diapause quiescence dormancy aestivation hibernation cold-hardiness heat-hardiness desiccation-hardiness anoxia-hardiness starvation-hardiness radiation-hardiness pressure-hardiness gravity-perception magnetic-field-perception electric-field-perception sound-perception vibration-perception substrate-borne-signal air-borne-signal seismic-signal chemical-signal visual-signal tactile-signal auditory-signal multimodal-signal composite-signal redundant-signal amplifying-signal attenuating-signal honest-signal deceptive-signal costly-signal cheap-signal conventional-signal index-signal handicap-signal amplifier social-selection sexual-selection sperm-competition cryptic-female-choice male-male-competition female-female-competition parent-offspring-conflict sibling-conflict worker-queen-conflict worker-worker-conflict sexual-conflict antagonistic-coevolution arms-race evolutionary-dynamics community-dynamics ecosystem-dynamics biogeochemical-cycle carbon-cycle nitrogen-cycle phosphorus-cycle sulfur-cycle water-cycle trophic-transfer production-efficiency consumption-efficiency assimilation-efficiency ecological-efficiency Lindeman-efficiency 10%-rule food-chain food-web trophic-cascade top-down-control bottom-up-control wasps-control donor-control recipient-control interaction-strength interaction-web ecological-network network-topology network-metrics degree-distribution connectance nestedness compartmentalization robustness fragility resilience recovery-time return-time elasticity perturbation pulse-disturbance press-disturbance chronic-disturbance acute-disturbance catastrophic-disturbance natural-disturbance anthropogenic-disturbance habitat-destruction habitat-degradation habitat-modification habitat-loss habitat-isolation edge-effect matrix-effect permeability corridor-effect stepping-stone source-population sink-population metapopulation-dynamics extinction-debt colonization-credit relaxation-effect time-lag historical-legacy priority-effect founder-effect bottleneck-effect selection natural-selection artificial-selection directional-selection stabilizing-selection disruptive-selection balancing-selection frequency-dependent-selection density-dependent-selection r-selection K-selection c-selection s-selection a-selection bet-hedging-selection fluctuating-selection correlational-selection multivariate-selection indirect-selection correlated-response genetic-correlation phenotypic-correlation environmental-correlation correlation-matrix variance-covariance-matrix G-matrix P-matrix E-matrix M-matrix quantitative-genetics heritability narrow-sense-heritability broad-sense-heritability genetic-variance additive-genetic-variance dominance-variance epistatic-variance environmental-variance phenotypic-variance genotype-environment-interaction-variance maternal-variance paternal-variance common-environment-variance special-environment-variance measurement-error-variance response-to-selection realized-heritability predicted-response breeder's-equation selection-gradient selection-differential Lande-equation multivariate-breeder's-equation genetic-constraint evolutionary-constraint developmental-constraint functional-constraint phylogenetic-constraint historical-constraint trade-off allocation-trade-off acquisition-trade-off conversion-trade-off maintenance-trade-off reproduction-survival-trade-off growth-reproduction-trade-off immune-reproduction-trade-off stress-reproduction-trade-off foraging-predation-trade-off competition-colonization-trade-off tolerance-resistance-trade-off plasticity-canalization-trade-off generalist-specialist-trade-off speed-accuracy-trade-off exploration-exploitation-trade-off individual-colony-trade-off within-colony-trade-off between-colony-trade-off life-history-trade-off demographic-trade-off ecological-trade-off evolutionary-trade-off optimization maximization minimization satisficing marginal-value-theorem optimal-foraging-theory optimal-diet-theory central-place-foraging patch-use-theory giving-up-density giving-up-time predation-risk energetic-cost opportunity-cost handling-time search-time travel-time encounter-rate prey-density prey-profitability prey-choice diet-breadth niche-width niche-overlap niche-partitioning resource-selection habitat-selection settlement-choice nest-site-choice host-choice oviposition-choice foraging-choice path-choice route-choice central-place territory core-area edge-area boundary perimeter area volume shape configuration spatial-pattern temporal-pattern diel-pattern seasonal-pattern annual-pattern multiannual-pattern cyclical-pattern irregular-pattern random-pattern clumped-pattern uniform-pattern regular-pattern aggregated-pattern overdispersed-pattern underdispersed-pattern Poisson-distribution negative-binomial-distribution Taylor's-power-law spatial-autocorrelation temporal-autocorrelation cross-correlation spectral-analysis wavelet-analysis time-series-analysis population-time-series community-time-series ecosystem-time-series long-term-study long-term-monitoring long-term-experiment chronosequence space-for-time-substitution paleoecology paleontology fossil-record trace-fossil coprolite nest-fossil amber-inclusion subfossil quaternary holocene pleistocene neogene paleogene cretaceous mesozoic cenozoic geological-time deep-time evolutionary-time ecological-time generational-time physiological-time perception-time reaction-time decision-time memory-time learning-time developmental-time life-span colony-lifespan queen-lifespan worker-lifespan male-lifespan larval-development-time pupal-development-time egg-development-time incubation-time doubling-time half-life relaxation-time resilience-time resistance-time lag-time lead-time forecast-time prediction-time planning-time management-time conservation-time restoration-time mitigation-time adaptation-time speciation-time diversification-time extinction-time persistence-time transience-time ephemerality permanence stability instability equilibrium non-equilibrium steady-state dynamic-state transient-state asymptotic-state attractor repeller limit-cycle chaos stochasticity determinism predictability unpredictability complexity emergence self-organization pattern-formation spatial-self-organization temporal-self-organization spatiotemporal-self-organization swarm-intelligence collective-decision-making consensus-decision quorum-response wisdom-of-crowds information-cascade cultural-evolution gene-culture-coevolution niche-construction extended-phenotype extended-organism superorganism colony-as-individual colony-as-unit-of-selection group-selection multilevel-selection reciprocal-altruism by-product-mutualism amensalism competition predation herbivory parasitism parasitoidism chemical-mimicry tactile-mimicry visual-mimicry acoustic-mimicry defensive-mimicry Vavilovian-mimicry Bakerian-mimicry Gilbertian-mimicry Poultonian-mimicry Browerian-mimicry auto-mimicry intra-specific-mimicry inter-specific-mimicry molecular-mimicry cellular-mimicry tissue-mimicry organ-mimicry behavioral-mimicry reproductive-mimicry social-mimicry nest-mimicry food-mimicry predator-mimicry prey-mimicry host-mimicry guest-mimicry cleptoparasite cuckoo-parasite inquiline guest myrmecophile termitophile aphidophile coccidophile scale-insect-associate mealybug-associate whitefly-associate planthopper-associate treehopper-associate leafhopper-associate cicada-associate butterfly-associate moth-associate beetle-associate fly-associate wasp-associate bee-associate ant-associate spider-associate mite-associate tick-associate nematode-associate flatworm-associate roundworm-associate annelid-associate mollusk-associate crustacean-associate millipede-associate centipede-associate insect-associate vertebrate-associate bird-associate mammal-associate reptile-associate amphibian-associate fish-associate plant-associate fungus-associate bacteria-associate virus-associate archaea-associate protist-associate algae-associate lichen-associate bryophyte-associate pteridophyte-associate gymnosperm-associate angiosperm-associate monocot-associate dicot-associate herb-associate shrub-associate tree-associate vine-associate epiphyte-associate lithophyte-associate rheophyte-associate hydrophyte-associate xerophyte-associate mesophyte-associate hygrophyte-associate sciophyte-associate heliophyte-associate pioneer-plant-associate climax-plant-associate disturbance-plant-associate successional-plant-associate ant-plant-protection-mutualism ant-plant-food-body-mutualism ant-plant-domatia-mutualism ant-plant-extrafloral-nectary-mutualism ant-plant-seed-dispersal-mutualism ant-plant-pollination-mutualism ant-plant-agriculture-mutualism ant-fungus-mutualism ant-bacteria-mutualism ant-virus-mutualism ant-nematode-mutualism ant-mite-mutualism ant-tick-mutualism ant-spider-mutualism ant-wasp-mutualism ant-fly-mutualism ant-beetle-mutualism ant-butterfly-mutualism ant-moth-mutualism ant-bird-mutualism ant-mammal-mutualism ant-reptile-mutualism ant-amphibian-mutualism ant-fish-mutualism ant-human-mutualism ant-domestic-animal-mutualism ant-pest-mutualism ant-crop-mutualism ant-forest-mutualism ant-savanna-mutualism ant-grassland-mutualism ant-desert-mutualism ant-wetland-mutualism ant-riverine-mutualism ant-riparian-mutualism ant-montane-mutualism ant-lowland-mutualism ant-coastal-mutualism ant-island-mutualism ant-continental-mutualism ant-tropical-mutualism ant-temperate-mutualism ant-subtropical-mutualism ant-boreal-mutualism ant-austral-mutualism ant-paleotropical-mutualism ant-neotropical-mutualism ant-afrotropical-mutualism ant-oriental-mutualism ant-palearctic-mutualism ant-nearctic-mutualism ant-holarctic-mutualism ant-pan-tropical-mutualism ant-cosmopolitan-mutualism ant-endemic-mutualism ant-widespread-mutualism ant-restricted-mutualism ant-rare-mutualism ant-common-mutualism ant-abundant-mutualism ant-scarce-mutualism ant-threatened-mutualism ant-invasive-mutualism ant-native-mutualism ant-exotic-mutualism ant-introduced-mutualism ant-naturalized-mutualism ant-feral-mutualism ant-urban-mutualism ant-rural-mutualism ant-agricultural-mutualism ant-natural-mutualism ant-anthropogenic-mutualism ant-pristine-mutualism ant-degraded-mutualism ant-restored-mutualism ant-conserved-mutualism ant-managed-mutualism ant-sustainable-mutualism ant-unsustainable-mutualism ant-resilient-mutualism ant-fragile-mutualism ant-robust-mutualism ant-vulnerable-mutualism ant-resistant-mutualism ant-susceptible-mutualism ant-tolerant-mutualism ant-intolerant-mutualism ant-flexible-mutualism ant-inflexible-mutualism ant-plastic-mutualism ant-canalized-mutualism ant-adaptive-mutualism ant-maladaptive-mutualism ant-optimal-mutualism ant-suboptimal-mutualism ant-efficient-mutualism ant-inefficient-mutualism ant-productive-mutualism ant-unproductive-mutualism ant-successful-mutualism ant-unsuccessful-mutualism ant-stable-mutualism ant-unstable-mutualism ant-persistent-mutualism ant-transient-mutualism ant-obligate-mutualism ant-facultative-mutualism ant-specific-mutualism ant-generalized-mutualism ant-exclusive-mutualism ant-non-exclusive-mutualism ant-symbiotic-mutualism ant-non-symbiotic-mutualism ant-trophic-mutualism ant-defensive-mutualism ant-dispersive-mutualism ant-pollination-mutualism ant-agriculture-mutualism ant-construction-mutualism ant-nutrient-mutualism ant-protection-mutualism ant-cleaning-mutualism ant-transport-mutualism ant-storage-mutualism ant-processing-mutualism ant-consumption-mutualism ant-production-mutualism ant-exchange-mutualism ant-reciprocity-mutualism ant-by-product-mutualism ant-investment-mutualism ant-harvest-mutualism ant-cultivation-mutualism ant-domestication-mutualism ant-selection-mutualism ant-breeding-mutualism ant-engineering-mutualism ant-modification-mutualism ant-creation-mutualism ant-destruction-mutualism ant-transformation-mutualism ant-conversion-mutualism ant-utilization-mutualism ant-exploitation-mutualism ant-cooperation-mutualism ant-competition-mutualism ant-conflict-mutualism ant-resolution-mutualism ant-negotiation-mutualism ant-communication-mutualism ant-signal-mutualism ant-cue-mutualism ant-information-mutualism ant-knowledge-mutualism ant-learning-mutualism ant-memory-mutualism ant-cognition-mutualism ant-intelligence-mutualism ant-decision-mutualism ant-choice-mutualism ant-preference-mutualism ant-motivation-mutualism ant-emotion-mutualism ant-stress-mutualism ant-welfare-mutualism ant-health-mutualism ant-disease-mutualism ant-immunity-mutualism ant-resistance-mutualism ant-tolerance-mutualism ant-recovery-mutualism ant-healing-mutualism ant-therapy-mutualism ant-medicine-mutualism ant-prevention-mutualism ant-security-mutualism ant-safety-mutualism ant-risk-mutualism ant-hazard-mutualism ant-threat-mutualism ant-danger-mutualism ant-damage-mutualism ant-injury-mutualism ant-death-mutualism ant-mortality-mutualism ant-survival-mutualism ant-longevity-mutualism ant-reproduction-mutualism ant-fertility-mutualism ant-fecundity-mutualism ant-sterility-mutualism ant-virginity-mutualism ant-mating-mutualism ant-copulation-mutualism ant-insemination-mutualism ant-fertilization-mutualism ant-egg-mutualism ant-laying-mutualism ant-incubation-mutualism ant-hatching-mutualism ant-emergence-mutualism ant-eclosion-mutualism ant-molting-mutualism ant-ecdysis-mutualism ant-metamorphosis-mutualism ant-development-mutualism ant-growth-mutualism ant-differentiation-mutualism ant-maturation-mutualism ant-aging-mutualism ant-senescence-mutualism ant-decomposition-mutualism ant-recycling-mutualism ant-energy-mutualism ant-carbon-mutualism ant-nitrogen-mutualism ant-phosphorus-mutualism ant-sulfur-mutualism ant-water-mutualism ant-oxygen-mutualism ant-hydrogen-mutualism ant-mineral-mutualism ant-vitamin-mutualism ant-hormone-mutualism ant-enzyme-mutualism ant-protein-mutualism ant-lipid-mutualism ant-carbohydrate-mutualism ant-nucleic-acid-mutualism ant-DNA-mutualism ant-RNA-mutualism ant-gene-mutualism ant-genome-mutualism ant-chromosome-mutualism ant-cell-mutualism ant-tissue-mutualism ant-organ-mutualism ant-system-mutualism ant-organism-mutualism ant-individual-mutualism ant-colony-mutualism ant-population-mutualism ant-species-mutualism ant-genus-mutualism ant-family-mutualism ant-order-mutualism ant-class-mutualism ant-phylum-mutualism ant-kingdom-mutualism ant-domain-mutualism ant-life-mutualism ant-earth-mutualism ant-universe-mutualism
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 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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 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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
Tetramesa
Tetramesa is a genus of minute phytophagous wasps in the family Eurytomidae comprising over 200 described species. Members are exclusively associated with grasses (Poaceae), where they typically induce stem or inflorescence galls. The genus exhibits pronounced host specificity, with most species restricted to a single grass species or closely related congeners. Several species have been deployed as biological control agents against invasive grasses, including T. romana for giant reed (Arundo donax) and candidate agents for medusahead (Taeniatherum caput-medusae) and African lovegrass (Eragrostis curvula). Adults feed on nectar. Recent phylogenomic studies have synonymized the genera Aiolomorphus and Cathilaria within Tetramesa.
biological-control gall-wasp grassland invasive-species-management host-specificity cryptic-species phylogenomics Eurytomidae Chalcidoidea Hymenoptera phytophagy Poaceae stem-gall inflorescence-gall integrated-pest-management riparian-restoration molecular-systematics species-delimitation GMYC COI 28S biocontrol-agent Arundo-donax giant-reed medusahead African-lovegrass Eragrostis Sporobolus barley-jointworm integrated-taxonomy Southern-Hemisphere-diversity grassland-expansion Miocene-diversification monophagy host-range-testing non-target-effects tick-habitat cattle-fever-tick Rio-Grande Big-Bend-National-Park greenhouse-rearing oviposition-behavior water-deficit-stress generation-time parasitoid Eurytoma-amicophaga Eurytominae Walker-1848 Terebrantes Apocrita microhymenoptera minute-wasp reduced-wing-venation antennal-segmentation mesosoma-sculpture ovipositor molecular-operational-taxonomic-units MOTU species-boundaries cryptic-diversity sampling-effects SPEDE-sampler host-genetic-proxies field-host-range no-choice-testing native-Australian-species non-target-risk host-phylogeny target-weed suppression topping mechanical-control integrated-weed-management native-vegetation-recovery light-penetration canopy-structure dispersal establishment post-release-evaluation adventive-population Austin-Texas Idaho England Wales Orissa Canary-Islands Canary-Is.Mediterranean South-Africa southern-Africa Northern-Hemisphere Australia North-America Texas California Greece central-Asia Iran Ukraine Moldova Russia Israel Tajikistan Japan Eragrostis-curvula Sporobolus-pyramidalis Sporobolus-natalensis Sporobolus-africanus Hyparrhenia-hirta Andropogon-gayanus Eragrostis-plana Eragrostis-planiculmis Eragrostis-biflora Eragrostis-capensis Eragrostis-trichophora Deschampsia-caespitosa Deschampsia-setacea Taeniatherum-caput-medusae Hordeum barley Phragmites common-reed Dactylis-glomerata orchardgrass Poales grass graminoid cereal-crop pasture rangeland invasive-weed noxious-weed habitat-structure ecosystem-functioning naturalised-species alien-grass donor-country receiver-country biocontrol-practitioner host-plant-resistance internode-infestation oviposition-preference behavioral-categories antennae-attitude laboratory-artifact polyphagy specialist generalist climate-fluctuation molecular-clock dated-analysis phylogenetic-clade intrapopulation-variation geographic-sub-structuring singleton-effect overestimation consensus-delimitation operational-taxonomic-unit integrative-taxonomy evidence-line caveat putative-species novel-taxon undescribed-species biodiversity-assessment faunal-survey field-collection genetic-sequence mitochondrial-DNA nuclear-DNA cytochrome-c-oxidase-I ribosomal-DNA internal-transcribed-spacer chloroplast-DNA rps16 rpl32 trnK trnL ITS plant-insect-interaction herbivore phytophage primary-consumer trophic-level food-web community-ecology restoration-ecology conservation-biological-control classical-biological-control augmentative-biological-control inundative-release establishment-rate population-increase downstream-dispersal river-mile biomass-reduction plant-community tick-feeding-ant tick-feeding-beetle habitat-modification law-enforcement international-border flood-control irrigation-channel stream-clogging river-bank-weakening native-vegetation-stifling wildlife-habitat-reduction cattle-inspection deer-inspection Rhipicephalus Boophilus tick-population vegetation-transition shading-effect long-term-suppression high-suppression cane-thinning mechanical-cutting growth-suppression susceptibility-to-attack combination-treatment synergistic-effect research-demonstration USDA Agricultural-Research-Service entomologist field-measurement post-release-monitoring impact-assessment agent-efficacy non-target-damage crop-safety agricultural-importance host-specific-herbivore sustainable-method cost-effective-method weed-control invasive-plant-management habitat-restoration ecosystem-service provisioning-service regulating-service supporting-service cultural-service biodiversity-conservation threatened-species endangered-species habitat-loss climate-change land-use-change anthropogenic-disturbance urbanization agricultural-intensification grazing-pressure fire-regime nitrogen-deposition carbon-sequestration soil-stabilization water-quality air-quality pollinator nectar-feeding adult-diet larval-diet tissue-feeding gall-induction plant-manipulation phytohormone cytokinin auxin cell-proliferation tissue-differentiation nutrient-sink sink-strength source-sink-dynamics plant-physiology reproductive-output seed-production seed-weight flowering-head inflorescence stem-elongation tiller-production biomass-allocation growth-rate relative-growth-rate net-primary-productivity ecosystem-productivity carbon-cycling nitrogen-cycling phosphorus-cycling decomposition litter-quality herbivory-impact top-down-control bottom-up-control trophic-cascade indirect-effect direct-effect density-dependence density-independence population-regulation population-dynamics life-table survivorship fecundity reproductive-rate intrinsic-rate-of-increase doubling-time carrying-capacity population-growth exponential-growth logistic-growth Allee-effect stochasticity environmental-variation demographic-variation genetic-variation phenotypic-plasticity local-adaptation evolutionary-response coevolution arms-race escape-and-radiate diversification-rate speciation extinction phylogenetic-diversity functional-diversity taxonomic-diversity alpha-diversity beta-diversity gamma-diversity species-richness species-evenness species-abundance-distribution rank-abundance-curve diversity-index Shannon-index Simpson-index evenness-index dominance-index similarity-index dissimilarity-index cluster-analysis ordination principal-component-analysis principal-coordinate-analysis non-metric-multidimensional-scaling canonical-correspondence-analysis redundancy-analysis variation-partitioning structural-equation-modeling path-analysis meta-analysis systematic-review evidence-synthesis research-priority knowledge-gap data-deficiency IUCN-Red-List conservation-status protected-area management-plan monitoring-protocol adaptive-management stakeholder-engagement policy-recommendation legislation regulation incentive market-based-instrument payment-for-ecosystem-services carbon-credit biodiversity-offset habitat-banking conservation-easement land-trust stewardship sustainable-agriculture organic-farming agroecology precision-agriculture conservation-tillage cover-cropping crop-rotation intercropping agroforestry silvopasture riparian-buffer filter-strip grassed-waterway wetland-restoration prairie-restoration savanna-restoration forest-restoration degraded-land abandoned-land marginal-land set-aside wildlife-corridor habitat-connectivity landscape-matrix patch-dynamics edge-effect interior-habitat core-area buffer-zone permeability resistance least-cost-path circuit-theory isolation-by-distance isolation-by-resistance gene-flow genetic-drift inbreeding outbreeding hybridization introgression admixture population-structure genetic-differentiation FST GST D allele-frequency haplotype genotype phenotype quantitative-trait morphological-trait behavioral-trait life-history-trait reproductive-trait survival-trait developmental-trait physiological-trait metabolic-trait stress-response immune-response defense-trait resistance-trait tolerance-trait avoidance-trait antipredator-defense chemical-defense mechanical-defense crypsis mimicry aposematism startle-display thanatosis autotomy regeneration compensatory-growth induced-defense constitutive-defense direct-defense indirect-defense volatile-organic-compound extrafloral-nectar domatia mutualism ant-plant-mutualism pollination-mutualism seed-dispersal-mutualism protection-mutualism nutrition-mutualism transport-mutualism cleaning-mutualism agricultural-mutualism domestication crop-wild-relative landrace cultivar variety breeding selection genetic-improvement yield quality stress-tolerance nutrient-use-efficiency water-use-efficiency photosynthetic-efficiency harvest-index grain-filling source-limitation sink-limitation assimilate-transport phloem-loading phloem-unloading remobilization senescence stay-green antioxidant reactive-oxygen-species oxidative-stress heat-stress cold-stress drought-stress salinity-stress metal-stress nutrient-deficiency nutrient-toxicity soil-pH soil-texture soil-structure soil-organic-matter soil-microbial-community rhizosphere mycorrhiza nitrogen-fixation phosphorus-solubilization potassium-mobilization micronutrient-uptake allelopathy competition facilitation niche-differentiation resource-partitioning temporal-partitioning spatial-partitioning morphological-partitioning physiological-partitioning phenological-partitioning competitive-exclusion competitive-displacement priority-effect founder-effect mass-effect species-sorting neutral-theory ecological-drift dispersal-limitation environmental-filtering biotic-filtering abiotic-filtering trait-based-approach functional-trait response-trait effect-trait community-assembly community-disassembly community-reassembly succession primary-succession secondary-succession pioneer-species climax-species keystone-species ecosystem-engineer foundation-species dominant-species subordinate-species transient-species resident-species migrant-species nomadic-species irruptive-species invasive-species naturalized-species casual-species established-species spreading-species widespread-species restricted-species endemic-species cosmopolitan-species disjunct-distribution relict-distribution refugium glacial-refugium interglacial-refugium postglacial-colonization range-shift range-expansion range-contraction range-fragmentation source-population sink-population metapopulation population-viability minimum-viable-population effective-population-size genetic-rescue demographic-rescue assisted-migration managed-relocation ex-situ-conservation in-situ-conservation cryopreservation seed-bank gene-bank botanic-garden arboretum zoo aquarium microbial-culture-collection DNA-bank tissue-bank sperm-bank oocyte-bank embryo-bank reintroduction translocation reinforcement supplementation headstarting captive-breeding propagation nucleus-population release-program soft-release hard-release acclimatization habitat-preparation predator-control competitor-control disease-control parasite-control health-screening quarantine veterinary-care nutrition-management environmental-enrichment behavioral-training imprinting fostering cross-fostering hand-rearing parent-rearing semi-natural-enclosure pre-release-conditioning survival-rate reproduction-rate dispersal-rate site-fidelity homeward-orientation migration navigation orientation magnetic-sense olfactory-sense visual-sense auditory-sense mechanosensory-sense thermosensory-sense hygrosensory-sense electrosensory-sense polarized-light celestial-cues landmark-cues odor-plume wind-direction sun-compass star-compass moon-compass magnetic-compass inertial-navigation path-integration cognitive-map spatial-memory temporal-memory episodic-memory procedural-memory associative-learning operant-conditioning classical-conditioning habituation sensitization song-learning tool-use problem-solving innovation social-learning cultural-transmission teaching cooperation altruism kin-selection reciprocal-altruism group-selection multilevel-selection selfish-gene inclusive-fitness reproductive-skew dominance-hierarchy territoriality home-range activity-center den-site nest-site roost-site singing-post display-site lek arena scrape burrow cavity crevice hole tunnel gallery chamber cell comb paper-nest mud-nest silk-nest spun-cocoon pupal-case ootheca egg-mass clutch brood litter covey pack herd flock colony society superorganism eusociality cooperative-breeding communal-breeding polygyny polyandry promiscuity monogamy serial-monogamy genetic-monogamy social-monogamy extra-pair-copulation sperm-competition cryptic-female-choice sexual-selection intersexual-selection intrasexual-selection mate-choice mate-guarding mate-switching divorce widowhood mate-replacement breeding-season breeding-cycle breeding-attempt breeding-success fledging-success recruitment juvenile-survival adult-survival longevity life-expectancy maximum-lifespan aging telomere oxidative-damage mitochondrial-dysfunction cellular-senescence apoptosis necrosis autophagy proteostasis DNA-repair epigenetic-modification transgenerational-effect maternal-effect paternal-effect parental-care maternal-care paternal-care biparental-care alloparental-care brood-care egg-care larval-care pupal-care food-provisioning nest-defense territory-defense predator-defense disease-defense parasite-defense hygiene-behavior grooming nest-cleaning cannibalism siblicide infanticide filial-cannibalism sexual-cannibalism cannibalistic-oophagy trophic-egg regurgitation trophallaxis food-sharing information-sharing alarm-call mobbing distraction-display broken-wing-display tailing persecution attack escape flight fight freeze faint death-feigning tonic-immobility startle-response withdrawal avoidance dishabituation latent-inhibition blocking overshadowing relative-validity conditioned-inhibition spontaneous-recovery renewal reinstatement rapid-reacquisition latent-learning insight-learning observational-learning vicarious-learning emulation imitation social-facilitation social-inhibition audience-effect bystander-effect conformity obedience compliance commitment consistency foot-in-the-door door-in-the-face low-ball that's-not-all sunk-cost endowment-effect loss-aversion risk-aversion risk-seeking probability-weighting hyperbolic-discounting temporal-discounting intertemporal-choice optimal-foraging marginal-value-theorem prey-choice patch-choice diet-breadth central-place-foraging load-size travel-time handling-time search-time encounter-rate prey-density prey-profitability prey-handling prey-detection prey-pursuit prey-capture prey-consumption prey-rejection ingestion-rate digestion-rate assimilation-efficiency production-efficiency ecological-efficiency trophic-transfer-efficiency Lindeman-efficiency ten-percent-rule pyramid-of-numbers pyramid-of-biomass pyramid-of-energy food-chain trophic-network energy-flow material-cycling biogeochemical-cycle carbon-cycle nitrogen-cycle phosphorus-cycle sulfur-cycle water-cycle hydrological-cycle atmospheric-circulation ocean-circulation thermohaline-circulation El-Niño La-Niña ENSO NAO AO PDO AMO climate-oscillation climate-anomaly weather-extreme drought flood heat-wave cold-snap storm hurricane typhoon cyclone tornado thunderstorm lightning wildfire volcanic-eruption earthquake tsunami landslide avalanche meteor-impact mass-extinction background-extinction speciation-rate extinction-rate net-diversification turnover standing-diversity fossil-record taphonomy preservation-bias sampling-bias collection-bias publication-bias citation-bias language-bias geographic-bias temporal-bias taxonomic-bias size-bias abundance-bias habitat-bias facies-bias Lagerstätte Konservat-Lagerstätte Konzentrat-Lagerstätte trace-fossil body-fossil chemical-fossil molecular-fossil biomarker isotope-signature paleoclimate-proxy paleoecology paleobiogeography paleoecosystem paleocommunity paleoenvironment paleogeography paleolatitude paleolongitude continental-drift plate-tectonics seafloor-spreading subduction orogeny mountain-building basin-formation sedimentation erosion weathering soil-formation pedogenesis horizon-development profile-development chronosequence toposequence biosequence climosequence landform geomorphology topography relief elevation slope aspect curvature roughness drainage watershed catchment basin subbasin reach segment zone riffle pool run glide cascade waterfall wetland marsh swamp bog fen mire peatland tundra taiga boreal-forest temperate-forest temperate-rainforest temperate-grassland tropical-forest tropical-rainforest tropical-dry-forest tropical-savanna tropical-grassland desert hot-desert cold-desert semiarid steppe prairie pampas veld savanna woodland shrubland chaparral maquis garigue fynbos kwongan mallee mulga mogote karst cave sinkhole doline polje uvala ponor spring seep oasis wadi arroyo gully ravine canyon gorge valley dale glen strath rift graben horst fault fold anticline syncline monocline dome plateau mesa butte escarpment bluff cliff crag tor inselberg bornhardt kopje hill knoll mound ridge spur saddle pass col gap notch cleft fissure joint bedding-plane foliation lineation cleavage schistosity gneissic-banding magma lava igneous-rock sedimentary-rock metamorphic-rock mineral crystal grain texture structure fabric composition chemistry mineralogy petrology geochemistry geophysics geochronology radiometric-dating isotopic-dating dendrochronology varve-chronology lichenometry archaeomagnetism paleomagnetism thermoluminescence optically-stimulated-luminescence electron-spin-resonance cosmogenic-nuclide exposure-dating burial-dating terrestrial-cosmogenic-nuclide in-situ-cosmogenic-nuclide meteoric-cosmogenic-nuclide fallout-radionuclide lead-210 cesium-137 plutonium americium radiocarbon carbon-14 calibration-curve intcal marine-reservoir-effect terrestrial-reservoir-effect hard-water-effect contamination pretreatment combustion graphitization accelerator-mass-spectrometry liquid-scintillation-counting gas-proportional-counting beta-counting gamma-spectrometry alpha-spectrometry mass-spectrometry inductively-coupled-plasma-mass-spectrometry thermal-ionization-mass-spectrometry secondary-ion-mass-spectrometry laser-ablation-inductively-coupled-plasma-mass-spectrometry electron-microprobe scanning-electron-microscopy transmission-electron-microscopy light-microscopy confocal-microscopy fluorescence-microscopy polarizing-microscopy phase-contrast-microscopy differential-interference-contrast-microscopy dark-field-microscopy bright-field-microscopy stereomicroscopy dissecting-microscopy microtomy sectioning staining mounting embedding fixation preservation collection curation vouchering repository herbarium museum archive database registry catalogue checklist inventory survey monitoring assessment evaluation appraisal audit review synthesis rapid-review scoping-review narrative-review critical-review state-of-the-art-review literature-review bibliometric-analysis citation-analysis co-citation-analysis bibliographic-coupling co-authorship-analysis collaboration-network knowledge-map concept-map ontology taxonomy classification nomenclature typification designation description diagnosis illustration measurement morphometry meristics count ratio index score rank grade scale level category class group set subset cluster clade lineage branch stem crown root node tip terminal internal basal derived ancestral plesiomorphic apomorphic synapomorphic autapomorphic homologous homoplastic convergent parallel reversal gain loss duplication deletion insertion inversion fusion fission polyploidy aneuploidy chromosome-number genome-size gene-number transcriptome proteome metabolome lipidome glycome interactome phenome connectome epigenome microbiome virome mycobiome bacteriome phytobiome rhizobiome endobiome ectobiome parasitome pathobiome diseasome toxome exposome foodome nutriome pharmacome toxicome adverse-outcome-pathway mode-of-action mechanism-of-action target-site receptor-binding enzyme-inhibition ion-channel-blockade membrane-disruption DNA-intercalation inflammation cell-differentiation cell-migration cell-adhesion cell-signaling signal-transduction gene-expression transcription translation post-translational-modification protein-folding protein-degradation protein-trafficking vesicle-transport exocytosis endocytosis phagocytosis pinocytosis mitophagy pexophagy ribophagy ER-phagy lysophagy lipophagy glycophagy ferritinophagy aggrephagy xenophagy entosis cytosis syncytium plasmodesmata gap-junction tight-junction adherens-junction desmosome hemidesmosome focal-adhesion cell-polarity cell-asymmetry cell-division mitosis meiosis cytokinesis karyokinesis chromosome-segregation spindle-assembly checkpoint DNA-replication DNA-recombination transposition retrotransposition horizontal-gene-transfer vertical-gene-transfer endosymbiosis symbiogenesis holobiont hologenome extended-phenotype niche-construction ecosystem-engineering keystone-process ecological-function ecosystem-function ecosystem-good natural-capital natural-resource renewable-resource non-renewable-resource common-pool-resource public-good private-good club-good externality market-failure government-failure policy-failure institutional-failure tragedy-of-the-commons free-rider collective-action social-dilemma prisoner's-dilemma chicken-game assurance-game coordination-game zero-sum-game positive-sum-game negative-sum-game win-win lose-lose win-lose Pareto-optimal Pareto-improvement Kaldor-Hicks-improvement cost-benefit-analysis cost-effectiveness-analysis multi-criteria-decision-analysis decision-support-system expert-system knowledge-based-system rule-based-system case-based-reasoning machine-learning deep-learning neural-network random-forest support-vector-machine decision-tree regression-tree classification-tree clustering-algorithm dimensionality-reduction feature-selection feature-extraction pattern-recognition image-analysis signal-processing time-series-analysis spatial-analysis geographic-information-system remote-sensing satellite-imagery aerial-photography drone-imagery LiDAR radar sonar seismic geophysical geochemical biogeochemical ecoinformatics bioinformatics computational-biology systems-biology synthetic-biology biotechnology nanotechnology genetic-engineering genome-editing CRISPR TALEN zinc-finger-nuclease transgenic cisgenic intragenic subgenic agrobacterium-mediated-transformation biolistics electroporation microinjection protoplast-fusion somatic-hybridization cybrid asymmetric-hybrid synthetic-polyploid doubled-haploid haploid-induction anther-culture microspore-culture ovule-culture embryo-culture endosperm-culture protoplast-culture cell-culture tissue-culture organ-culture shoot-culture root-culture callus-culture suspension-culture bioreactor hairy-root-culture transformation hardening micropropagation clonal-propagation meristem-culture shoot-tip-culture axillary-bud-culture nodal-culture leaf-culture petiole-culture stem-culture internode-culture rhizome-culture bulb-culture corm-culture tuber-culture seed-culture zygotic-embryo-culture somatic-embryo-culture somatic-embryogenesis organogenesis caulogenesis rhizogenesis adventitious-shoot-formation direct-regeneration indirect-regeneration primary-metabolism secondary-metabolism biosynthesis biotransformation bioconversion biodegradation bioremediation phytoremediation mycoremediation zooremediation bioaugmentation biostimulation natural-attenuation monitored-natural-attenuation engineered-natural-attenuation permeable-reactive-barrier pump-and-treat soil-vapor-extraction air-sparging thermal-desorption incineration landfilling containment capping slurry-wall grout-curtain sheet-piling clay-liner geomembrane geotextile geogrid geocomposite geosynthetic geofabric geonet geocell geopipe geostrip geofoam geobag geotube geocontainer geomat geoblanket geospacer geodrain geoinjection geogrout geopier geocolumn geopile geowall geoberm geotrench geocut geofill geolift geoslab geopanel geoblock geobrick geotile geopaver geoweb geomattress geocushion
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 payment-for-results results-based-payment conditional-payment performance-based-payment outcome-based-payment environmental-service-payment PSA PES REDD REDD+AFOLU LULUCF NAMA NAMC NDC NAP NBSAP national-adaptation-plan national-biodiversity-strategy-and-action-plan nationally-determined-contribution nationally-appropriate-mitigation-action intended-nationally-determined-contribution long-term-low-emission-development-strategy net-zero-target carbon-budget remaining-carbon-budget emission-gap adaptation-gap finance-gap implementation-gap ambition-loop ratchet-mechanism global-stocktake transparency-framework enhanced-transparency-framework common-but-differentiated-responsibilities respective-capabilities climate-equity climate-litigation climate-governance climate-diplomacy climate-negotiation UNFCCC COP CMP CMA SBSTA SBI IPCC IPBES UNCCD CBD CITES Ramsar World-Heritage Man-and-the-Biosphere MAB UNESCO FAO WHO ILO UNDP UNEP UN-Habitat World-Bank IMF WTO OECD GEF GCF AF multilateral-development-bank regional-development-bank national-development-bank development-finance-institution export-credit-agency sovereign-wealth-fund pension-fund insurance-company asset-manager asset-owner corporate-investor retail-investor impact-investor philanthropic-investor development-partner bilateral-donor multilateral-donor private-donor foundation NGO CSO civil-society-organization non-governmental-organization community-based-organization indigenous-peoples-organization women's-organization youth-organization faith-based-organization research-organization think-tank university academic-institution knowledge-institution capacity-building-institution training-institution extension-service advisory-service consulting-firm engineering-firm technology-provider equipment-supplier service-provider contractor operator maintainer utility municipality local-government provincial-government national-government federal-government ministry department agency authority commission 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seed-commons water-commons fisheries-commons forest-commons pasture-commons land-commons urban-commons atmospheric-commons global-commons common-heritage-of-mankind common-concern-of-humankind restorative-justice multispecies-justice more-than-human anthropocene capitalocene plantationocene chthulucene symbiocene good-anthropocene great-acceleration great-transformation great-decoupling great-convergence great-divergence great-fragmentation great-integration great-transition great-work long-now deep-time futures-thinking future-studies strategic-foresight scenario-planning trend-analysis weak-signal wild-card black-swan grey-rhino blue-sky green-swan systemic-change structural-change cultural-change behavioral-change individual-change collective-change social-change organizational-change technological-change economic-change political-change ecological-change planetary-change earth-system-change biodiversity-change ocean-change cryosphere-change atmospheric-change biogeochemical-change 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