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Pinnixa chaetopterana
tube pea crab
Pinnixa chaetopterana is a small, soft-bodied pea crab that lives symbiotically within the tubes of polychaete worms, primarily Chaetopterus variopedatus. It is a facultative commensal, occasionally free-living or associating with alternative hosts such as Amphitrite ornata. The species exhibits strong host fidelity and can distinguish conspecifics from heterospecifics. It ranges from Massachusetts to Uruguay along the western Atlantic, including Brazil and the Gulf of Mexico.
commensal symbiosis polychaete-association pea-crab western-Atlantic facultative-symbiont carcinonemertid-parasite host-fidelity conspecific-recognition zoeal-larvae tube-dwelling marine-invertebrate brachyuran-crab pinnotherid Chaetopterus zooplankton-larva mutual-intolerance polychaete-crab nemertean-parasite synchronous-reproduction-with-parasite caltrop-shaped-carapace branchial-chamber-parasite pleopod-egg-attachment host-tube-modification estuarine coastal-marine depth-to-60m Massachusetts-to-Brazil Gulf-of-Mexico Florida North-Carolina New-Jersey Uruguay Stimpson-1860 Tubicolixa-synonym soft-bodied-crab wider-than-long-carapace five-zoeal-stages dorsal-rostral-lateral-spines abdominal-somite-knobs wing-like-knobs median-notch-telson Amphitrite-ornata-alternative-host posterior-positioning-behavior tube-aperture-biting plankton-feeding predator-refuge Carcinonemertes-pinnotheridophila ribbon-worm-parasite synchronous-egg-development host-parasite-egg-competition branchial-exoskeleton-attachment excurrent-canal-opening female-1-2-nemerteans male-nemerteans-associated medial-sheath-attachment sternum-opening nemertean-egg-sacs crab-egg-hairs facultative-free-living obligate-vs-facultate-commensalism host-specificity symbiont-recognition heterospecific-avoidance limb-tearing-aggression co-occurrence-exclusion morphological-crypsis adult-identification-difficulty larval-dispersal settlement-behavior tube-worm-crab parchment-tube-inhabitant ventilated-tube-dwelling water-pumping-utilization host-turning-response symbiont-positioning entry-exit-hole-creation behavioral-plasticity ecological-flexibility host-range-expansion alternative-host-use geographic-range depth-range Atlantic-coast-distribution western-hemisphere-endemic American-Atlantic-fauna benthic-crustacean infaunal-associate sediment-tube-dweller marine-symbiosis decapod-brachyuran eubrachyuran thoracotreme pinnotheroid pinnixine tubicolixine species-complex-member cryptic-species-candidate morphological-similarity taxonomic-revision-needed type-locality-unknown 1860-description Stimpson's-pea-crab historical-taxonomy modern-synonymy Catalogue-of-Life-synonym GBIF-Tubicolixa-match genus-transfer-candidate subgeneric-classification infraspecific-variation sexual-dimorphism-unknown male-gonopod-diagnostic female-pleopod-brooding ovigerous-female-parasite-load nemertean-prevalence-unknown incidence-data-limited biological-relationship-documentation symbiont-life-history-synchronization reproductive-timing-overlap egg-development-coordination hatching-synchrony larval-release-coordination population-dynamics-unknown demography-unstudied growth-rate-unknown longevity-unknown mortality-factors-unknown predation-pressure predator-avoidance tube-refuge-value structural-habitat-modification engineering-behavior niche-construction host-damage tube-wall-perforation host-fitness-effects-unknown commensalism-cost-benefit mutualism-possibility parasitism-gradient symbiosis-continuum interaction-outcome-variability environmental-context-dependency host-quality-variation host-size-correlation crab-size-host-size-relationship settlement-cue-unknown larval-settlement-behavior metamorphosis-trigger competency-period pre-settlement-swimming post-settlement-adaptation host-recognition-mechanism chemical-cue-use tactile-cue-use host-tube-chemistry-detection conspecific-chemical-detection heterospecific-chemical-detection aggression-trigger territorial-behavior resource-defense host-monopolization pair-formation mating-system-unknown sex-ratio-unknown operational-sex-ratio mate-guarding-unknown sperm-competition-unknown fertilization-mode internal-fertilization egg-brooding-duration embryonic-development-time larval-development-mode planktotrophic-larvae lecithotrophy-unknown zoeal-feeding zoeal-duration megalopa-stage-unknown settlement-competence juvenile-adaptation host-tube-colonization subadult-behavior size-at-maturity-unknown molting-frequency-unknown ecdysis-in-host-tube exuvia-disposal grooming-behavior-unknown feeding-mechanism suspension-feeding filter-feeding plankton-capture host-current-use respiratory-current-exploitation oxygen-availability hypoxia-tolerance salinity-tolerance temperature-range-unknown physiological-ecology-studied oxygen-consumption-unknown metabolic-rate-unknown energetics-unknown growth-efficiency-unknown reproductive-effort-unknown fecundity-unknown egg-size-unknown offspring-size-number-tradeoff parental-investment maternal-care brooding-cost parasite-load-cost nemertean-reproduction-facilitation host-egg-loss parasite-egg-competition hatching-success-reduction fitness-consequence-unknown population-level-impact-unknown community-role-unknown trophic-position-unknown energy-flow-contribution nutrient-cycling-role sediment-interaction bioturbation-unknown ecosystem-engineering-secondary habitat-complexity-contribution biodiversity-association species-interaction-network food-web-position predator-unknown prey-unknown diet-breadth trophic-level omnivory-unknown carnivory-unknown detritivory-unknown suspension-feeding-confirmed host-food-waste-use pseudofeces-consumption-unknown host-mucus-use host-tissue-consumption-unknown parasitism-unknown kleptoparasitism-unknown thievery-unknown aggression-context intraspecific-aggression interspecific-aggression limb-autotomy regeneration-unknown wound-healing stress-response handling-response laboratory-maintenance collection-method preservation-effect museum-specimen-quality type-specimen-location original-description-quality subsequent-redescriptions taxonomic-stability nomenclatural-acts priority-rules homonymy-check synonymy-list-completeness distribution-record-verification occurrence-data-quality range-boundary-uncertainty depth-record-accuracy habitat-description-specificity host-record-verification association-record-quality symbiosis-documentation-standard natural-history-completeness research-priority-areas conservation-status-unknown IUCN-evaluation-needed population-trend-unknown threat-assessment habitat-vulnerability climate-sensitivity range-shift-potential phenology-change host-phenology-dependency phenological-mismatch-risk evolutionary-history-unknown phylogenetic-position sister-species-unknown species-group-membership genetic-differentiation cryptic-diversity-potential barcode-library-coverage genome-resources transcriptome-unknown proteome-unknown metabolome-unknown developmental-genetics-unknown symbiosis-genetics-unknown host-recognition-gene-unknown immune-response-unknown parasite-resistance-unknown coevolution-history phylogeography-unknown population-structure-unknown gene-flow-unknown dispersal-limitation isolation-by-distance local-adaptation-unknown phenotypic-plasticity morphological-variation environmental-gradient-response phenotypic-cline genotype-environment-interaction evolutionary-potential adaptive-capacity resilience-unknown extinction-risk-assessment-needed
Stictocephala stimulea
Stictocephala stimulea is a treehopper species in the family Membracidae, first described by Van Duzee in 1914. Treehoppers in this genus are characterized by their enlarged, often ornate pronotum that extends over the body like a helmet or thorn. The specific epithet 'stimulea' suggests possible resemblance to or association with stinging organisms, though this refers to appearance rather than actual sting. Like other membracids, this species likely feeds on plant phloem sap using piercing-sucking mouthparts.
Membracidae treehopper Auchenorrhyncha Cicadomorpha Stictocephala Ceresini Smiliinae Hemiptera sap-feeder pronotum helmet thorn-mimicry plant-feeding-insect phloem-feeder cryptic-coloration camouflage Van-Duzee-1914 Nearctic North-America Florida North-Carolina South-Dakota understudied-species data-deficient taxonomy entomology insect arthropod true-bug auchenorrhynchan cicadomorph leafhopper-relative plant-hopper ornate-pronotum pronotal-extension dorsal-helmet insect-morphology insect-systematics biodiversity natural-history conservation insect-ecology insect-behavior insect-development insect-distribution biogeography species-description type-specimen original-description historical-taxonomy museum-specimen collection-record occurrence-data GBIF iNaturalist citizen-science observation-record range-map distribution-data geographic-range endemic native-species regional-fauna local-biodiversity habitat-specificity host-plant feeding-guild phytophagous herbivore plant-insect-interaction coevolution mimicry defense crypsis evolutionary-adaptation morphological-adaptation exoskeleton-modification structural-coloration insect-identification field-guide diagnostic-character taxonomic-key species-complex cryptic-species morphological-variation geographic-variation phenotypic-plasticity life-history reproduction oviposition nymph instar metamorphosis hemimetabolous incomplete-metamorphosis population-dynamics community-ecology food-web trophic-level primary-consumer ecosystem-function nutrient-cycling energy-flow biological-control pest-status agricultural-relevance forestry urban-ecology invasive-potential climate-change habitat-loss conservation-status data-gap research-need taxonomic-revision phylogenetics molecular-systematics DNA-barcoding integrative-taxonomy natural-history-collection specimen-digitization open-data FAIR-data biodiversity-informatics biological-database taxonomic-authority nomenclature zoological-nomenclature International-Code-of-Zoological-Nomenclature type-locality type-specimen-deposition author-citation original-combination current-combination synonymy homonymy nomenclatural-act taxonomic-history classification-system phylogenetic-classification evolutionary-relationships monophyly clade lineage stem-group crown-group divergence-time molecular-clock fossil-record paleontology extant extinct living-fossil evolutionary-history biogeographic-history dispersal vicariance range-expansion range-contraction refugium glacial-relict postglacial-colonization anthropogenic-impact human-disturbance land-use-change urbanization agricultural-intensification pesticide-use habitat-fragmentation connectivity corridor protected-area conservation-planning systematic-conservation-planning priority-area hotspot endemic-area center-of-origin center-of-diversity evolutionary-significant-unit management-unit population metapopulation source-sink-dynamics demography vital-rates survival fecundity generation-time population-growth-rate carrying-capacity density-dependence regulation limiting-factor environmental-stochasticity demographic-stochasticity genetic-stochasticity inbreeding-depression outbreeding-depression genetic-diversity heterozygosity allelic-richness effective-population-size genetic-drift gene-flow migration isolation-by-distance isolation-by-resistance landscape-genetics population-genomics phylogeography historical-demography coalescent-theory approximate-Bayesian-computation species-distribution-model ecological-niche-model habitat-suitability bioclimatic-envelope climate-envelope niche-conservatism niche-evolution realized-niche fundamental-niche biotic-interaction competition predation parasitism mutualism commensalism amensalism facilitation apparent-competition indirect-interaction trophic-cascade top-down-control bottom-up-control mesopredator-release keystone-species ecosystem-engineer foundation-species dominant-species characteristic-species indicator-species umbrella-species flagship-species surrogate-species focal-species target-species pest-species invasive-species naturalized-species alien-species non-native-species introduced-species cryptogenic-species range-shift poleward-shift elevational-shift phenological-shift temporal-mismatch trophic-mismatch phenology seasonality voltinism univoltine bivoltine multivoltine partial-generation prolonged-diapause aestivation hibernation overwintering overwintering-strategy cold-hardiness freeze-tolerance freeze-avoidance supercooling cryoprotectant thermal-biology thermoregulation behavioral-thermoregulation microclimate-selection habitat-selection oviposition-site-selection host-selection mate-choice sexual-selection sperm-competition cryptic-female-choice reproductive-isolation prezygotic-isolation postzygotic-isolation speciation allopatric-speciation sympatric-speciation parapatric-speciation peripatric-speciation reinforcement hybridization introgression hybrid-zone cline ecotone contact-zone secondary-contact exogenous-selection endogenous-selection ecological-speciation mutation-order-speciation sexual-conflict sexual-dimorphism sexual-size-dimorphism sexual-dichromatism polymorphism color-polymorphism morph phase gregarious-phase solitary-phase density-dependent-polyphenism reaction-norm genotype-environment-interaction plasticity canalization genetic-assimilation acclimation acclimatization hardening stress-response heat-shock-protein oxidative-stress hypoxia desiccation starvation immune-response innate-immunity humoral-immunity cellular-immunity encapsulation nodulation phagocytosis melanization antimicrobial-peptide phenoloxidase prophenoloxidase lysozyme defensin cecropin diptericin attacin immune-gene immune-pathway Toll-pathway Imd-pathway JAK-STAT-pathway NF-κB relish dorsal immune-memory specific-immunity transgenerational-immune-priming gut-microbiome symbiosis obligate-symbiont facultative-symbiont primary-symbiont secondary-symbiont vertical-transmission horizontal-transmission mixed-mode-transmission nutritional-symbiosis defensive-symbiosis digestive-symbiosis detoxification metabolism biosynthesis vitamin-synthesis amino-acid-synthesis steroid-synthesis cuticle-synthesis chitin-synthesis molting ecdysis ecdysteroid juvenile-hormone insulin-signaling TOR-signaling nutrient-sensing energy-storage lipid-metabolism carbohydrate-metabolism protein-metabolism flight-metabolism respiration tracheal-system spiracle gas-exchange water-balance osmoregulation excretion Malpighian-tubule hindgut rectum cryptonephridial-system ion-transport water-reabsorption nitrogenous-waste uric-acid urea ammonia detoxification-enzyme cytochrome-P450 glutathione-S-transferase carboxylesterase mixed-function-oxidase resistance insecticide-resistance acaricide-resistance herbicide-resistance antibiotic-resistance resistance-management resistance-evolution resistance-mechanism target-site-resistance metabolic-resistance behavioral-resistance cuticular-resistance resistance-allele resistance-gene fitness-cost fitness-benefit pleiotropy trade-off life-history-trade-off allocation resource-allocation energy-allocation time-allocation foraging optimal-foraging central-place-foraging patch-use marginal-value-theorem predator-avoidance antipredator-defense masquerade Batesian-mimicry Müllerian-mimicry automimicry intraspecific-mimicry intersexual-mimicry aggressive-mimicry social-parasitism brood-parasitism kleptoparasitism defense-behavior thanatosis death-feigning startle-display deimatic-display eyespot deflection-mark autotomy reflex-bleeding regurgitation defensive-secretion venom toxin allomone kairomone synomone semiochemical pheromone sex-pheromone aggregation-pheromone alarm-pheromone trail-pheromone territorial-pheromone recognition-pheromone signature-mixture cuticular-hydrocarbon volatile-organic-compound olfaction gustation mechanoreception thermoreception hygroreception photoreception vision compound-eye ommatidium rhabdom visual-pigment opsin color-vision polarization-vision motion-detection optomotor-response phototaxis photoperiodism circadian-rhythm biological-clock entrainment zeitgeber circannual-rhythm photoperiod day-length critical-day-length critical-photoperiod diapause-induction diapause-termination diapause-maintenance diapause-development endocrine-control neuroendocrine corpus-allatum corpus-cardiacum prothoracic-gland ecdysiotropin ecdysiostatin allatostatin allatotropin neuropeptide hormone hormone-receptor signal-transduction gene-expression transcription-factor regulatory-gene structural-gene gene-network gene-regulatory-network developmental-gene Hox-gene homeobox segment-polarity-gene gap-gene pair-rule-gene maternal-effect-gene zygotic-gene embryonic-development embryogenesis cleavage blastoderm gastrulation segmentation tagmosis appendage-development imaginal-disc holometabolous complete-metamorphosis larva pupa adult apolysis teneral sclerotization tanning cuticle exoskeleton epicuticle procuticle exocuticle endocuticle resilin sclerotin chitin protein lipid wax cement-layer wax-layer surface-lipid waterproofing anti-wetting anti-fouling self-cleaning structural-color pigmentary-color ommin pteridine carotenoid melanin papiliochrome flavonoid anthocyanin betalain iridescence photonic-crystal diffraction-grating thin-film-interference multilayer-reflector scattering Tyndall-scattering Rayleigh-scattering Mie-scattering non-iridescent-structural-color biochrome biofluorescence bioluminescence luciferin luciferase photoprotein bioluminescent-system counter-illumination burglar-alarm lure communication mate-recognition predator-deterrence aposematism warning-coloration honest-signal dishonest-signal signal-evolution signal-design receiver-psychology sensory-bias sensory-exploitation sensory-trap amplifier attenuator signal-efficacy signal-honesty handicap-principle index-signal conventional-signal assessment-signal territorial-signal dominance-signal submissive-signal aggressive-signal courtship-signal copulatory-signal postcopulatory-signal antagonistic-coevolution chase-away-selection sensory-drive reproductive-character-displacement genetic-exchange admixture hybrid-fitness hybrid-breakdown hybrid-vigor heterosis genetic-load mutational-meltdown founder-effect bottleneck population-expansion invasion-genetics propagule-pressure introduction-effort establishment spread lag-phase exponential-growth logistic-growth Allee-effect demographic-Allee-effect component-Allee-effect strong-Allee-effect weak-Allee-effect critical-population-size minimum-viable-population extinction-risk population-viability-analysis stochastic-model deterministic-model individual-based-model agent-based-model spatially-explicit-model metapopulation-model source-sink-model integrated-population-model state-space-model hierarchical-model Bayesian-model machine-learning artificial-intelligence computer-vision image-recognition automated-identification deep-learning neural-network convolutional-neural-network transfer-learning data-augmentation training-set validation-set test-set accuracy precision recall F1-score confusion-matrix receiver-operating-characteristic area-under-curve model-selection model-validation cross-validation bootstrapping permutation-test null-model randomization-test Monte-Carlo-simulation Markov-chain-Monte-Carlo Gibbs-sampling Metropolis-Hastings-algorithm Bayesian-inference prior-distribution posterior-distribution likelihood-function evidence model-comparison Bayes-factor information-criterion Akaike-information-criterion corrected-AIC Bayesian-information-criterion deviance-information-criterion Watanabe-Akaike-information-criterion cross-validation-information-criterion stacking model-averaging ensemble-model prediction forecasting scenario-analysis sensitivity-analysis uncertainty-analysis propagation-of-uncertainty error-budget confidence-interval credible-interval prediction-interval tolerance-interval statistical-power sample-size effect-size significance-test hypothesis-testing null-hypothesis alternative-hypothesis type-I-error type-II-error false-positive false-negative multiple-comparison Bonferroni-correction false-discovery-rate q-value p-value significance-level alpha-level beta-level statistical-test parametric-test nonparametric-test robust-test exact-test asymptotic-test bootstrap-test jackknife resampling subsampling leave-one-out k-fold stratified-sampling balanced-sampling imbalanced-data class-imbalance oversampling undersampling synthetic-minority-oversampling SMOTE ADASYN cost-sensitive-learning ensemble-method bagging boosting random-forest gradient-boosting XGBoost LightGBM CatBoost support-vector-machine kernel-method radial-basis-function polynomial-kernel linear-kernel sigmoid-kernel decision-tree classification-tree regression-tree pruning cost-complexity-pruning minimum-description-length rule-based-classifier association-rule frequent-pattern apriori-algorithm FP-growth sequential-pattern temporal-pattern spatial-pattern spatiotemporal-pattern time-series-analysis trend-analysis seasonal-decomposition autoregressive-model moving-average-model ARIMA SARIMA exponential-smoothing state-space-model Kalman-filter particle-filter hidden-Markov-model dynamic-Bayesian-network recurrent-neural-network long-short-term-memory gated-recurrent-unit transformer attention-mechanism self-attention multi-head-attention encoder decoder encoder-decoder-architecture sequence-to-sequence-model generative-model generative-adversarial-network variational-autoencoder normalizing-flow diffusion-model energy-based-model Boltzmann-machine restricted-Boltzmann-machine deep-belief-network autoencoder denoising-autoencoder sparse-autoencoder contractive-autoencoder variational-inference expectation-maximization maximum-likelihood maximum-a-posteriori maximum-entropy Occam's-razor parsimony model-complexity overfitting underfitting bias-variance-tradeoff regularization L1-regularization L2-regularization elastic-net dropout early-stopping batch-normalization layer-normalization group-normalization instance-normalization weight-decay mixup cutout cutmix random-erasing autoaugment fast-autoaugment population-based-augmentation neural-architecture-search automated-machine-learning hyperparameter-optimization grid-search random-search Bayesian-optimization sequential-model-based-optimization multi-fidelity-optimization knowledge-distillation teacher-student-model model-compression quantization knowledge-transfer domain-adaptation few-shot-learning zero-shot-learning one-shot-learning meta-learning learning-to-learn metric-learning siamese-network triplet-loss contrastive-learning self-supervised-learning unsupervised-learning semi-supervised-learning weakly-supervised-learning active-learning online-learning incremental-learning continual-learning lifelong-learning curriculum-learning multi-task-learning multi-modal-learning cross-modal-learning representation-learning feature-learning embedding-learning manifold-learning dimensionality-reduction principal-component-analysis t-SNE UMAP word-embedding sentence-embedding document-embedding knowledge-graph-embedding graph-neural-network graph-convolutional-network graph-attention-network message-passing-neural-network relational-graph-convolutional-network inductive-learning transductive-learning homophily heterophily graph-isomorphism Weisfeiler-Lehman-test subgraph-isomorphism graph-matching network-alignment network-embedding node-embedding edge-embedding graph-classification node-classification link-prediction graph-generation graph-synthesis molecular-generation de-novo-design inverse-design retrosynthesis reaction-prediction property-prediction quantitative-structure-activity-relationship QSAR quantitative-structure-property-relationship QSPR molecular-descriptor fingerprint Morgan-fingerprint extended-connectivity-fingerprint circular-fingerprint atom-pair-fingerprint topological-torsion-fingerprint pharmacophore shape-similarity electrostatic-similarity docking scoring-function binding-affinity-prediction protein-ligand-interaction protein-protein-interaction protein-structure-prediction AlphaFold protein-folding contact-prediction distance-prediction angle-prediction torsion-prediction secondary-structure-prediction tertiary-structure-prediction quaternary-structure-prediction protein-design protein-engineering directed-evolution rational-design computational-design de-novo-protein-design enzyme-design catalytic-antibody ribozyme aptamer DNAzyme synthetic-biology genetic-circuit metabolic-engineering metabolic-flux-analysis flux-balance-analysis genome-scale-metabolic-model dynamic-metabolic-model kinetic-modeling stoichiometric-modeling constraint-based-modeling optimization linear-programming mixed-integer-linear-programming quadratic-programming nonlinear-programming dynamic-programming stochastic-programming robust-optimization multi-objective-optimization Pareto-optimality evolutionary-algorithm genetic-algorithm genetic-programming differential-evolution particle-swarm-optimization ant-colony-optimization simulated-annealing tabu-search variable-neighborhood-search iterated-local-search greedy-randomized-adaptive-search scatter-search path-relinking memetic-algorithm hyper-heuristic metaheuristic local-search neighborhood-search hill-climbing random-walk randomized-search Monte-Carlo-tree-search upper-confidence-bound Thompson-sampling multi-armed-bandit contextual-bandit reinforcement-learning policy-gradient actor-critic Q-learning deep-Q-network policy-iteration value-iteration temporal-difference-learning SARSA expected-SARSA double-Q-learning dueling-Q-network prioritized-experience-replay distributed-reinforcement-learning impala A3C asynchronous-advantage-actor-critic PPO proximal-policy-optimization TRPO trust-region-policy-optimization DDPG deep-deterministic-policy-gradient TD3 twin-delayed-deep-deterministic-policy-gradient SAC soft-actor-critic model-based-reinforcement-learning model-predictive-control guided-policy-search imitation-learning inverse-reinforcement-learning apprenticeship-learning behavior-cloning dataset-aggregation DAgger generative-adversarial-imitation-learning third-person-imitation-learning one-shot-imitation-learning meta-reinforcement-learning hierarchical-reinforcement-learning option skill subgoal intrinsic-motivation curiosity-driven-exploration empowerment information-gain prediction-error novelty-seeking diversity-seeking multi-agent-reinforcement-learning mean-field fictitious-play correlated-equilibrium Nash-equilibrium game-theory mechanism-design auction-theory incentive-design social-choice-theory voting-theory fair-division matching-theory market-design algorithmic-game-theory computational-social-choice computational-complexity NP-completeness NP-hardness approximation-algorithm approximation-ratio inapproximability fixed-parameter-tractability parameterized-complexity kernelization treewidth graph-minor forbidden-minor-characterization well-quasi-ordering Robertson-Seymour-theorem graph-structure-theorem bidimensionality-theory meta-theorem Courcelle's-theorem automata-theory finite-automaton pushdown-automaton Turing-machine computability decidability halting-problem Rice's-theorem Gödel's-incompleteness-theorems lambda-calculus combinatory-logic type-theory proof-theory model-theory set-theory forcing large-cardinal axiom-of-choice continuum-hypothesis independence-result consistency-strength reverse-mathematics constructive-mathematics intuitionistic-logic modal-logic temporal-logic epistemic-logic deontic-logic description-logic ontology knowledge-representation semantic-web linked-data RDF SPARQL OWL knowledge-graph entity-linking relation-extraction event-extraction coreference-resolution named-entity-recognition part-of-speech-tagging dependency-parsing constituency-parsing semantic-parsing natural-language-inference textual-entailment natural-language-generation machine-translation summarization question-answering dialogue-system conversational-AI chatbot virtual-assistant speech-recognition speech-synthesis speaker-recognition emotion-recognition sentiment-analysis opinion-mining aspect-based-sentiment-analysis target-dependent-sentiment-analysis stance-detection rumor-detection fake-news-detection fact-checking claim-verification evidence-retrieval argument-mining persuasion-detection deception-detection personality-prediction mental-health-detection suicide-prevention crisis-intervention social-media-mining network-analysis community-detection influence-maximization information-diffusion cascade-modeling epidemic-modeling SEIR-model SIR-model agent-based-epidemic-model compartmental-model network-epidemic-model metapopulation-epidemic-model spatial-epidemic-model stochastic-epidemic-model branching-process renewal-equation generation-interval serial-interval incubation-period latent-period infectious-period basic-reproduction-number effective-reproduction-number herd-immunity-threshold vaccination-coverage vaccine-efficacy vaccine-effectiveness breakthrough-infection waning-immunity immune-escape variant-of-concern variant-of-interest variant-under-monitoring phylogenetic-analysis molecular-clock-analysis phylodynamics viral-evolution antigenic-drift antigenic-shift zoonotic-spillover reservoir-host intermediate-host amplifying-host dead-end-host vector-borne-disease mosquito-borne-disease tick-borne-disease flea-borne-disease lice-borne-disease midge-borne-disease sandfly-borne-disease tsetse-borne-disease blackfly-borne-disease triatomine-borne-disease kissing-bug Chagas-disease trypanosomiasis leishmaniasis malaria dengue yellow-fever Zika-virus chikungunya West-Nile-virus Japanese-encephalitis tick-borne-encephalitis Lyme-disease Rocky-Mountain-spotted-fever ehlichiosis anaplasmosis babesiosis tularemia plague murine-typhus epidemic-typhus endemic-typhus scrub-typhus Q-fever trench-fever bartonellosis cat-scratch-disease rat-bite-fever leptospirosis brucellosis anthrax melioidosis glanders tularaemia francisellosis coxiellosis rickettsiosis orientia-infection ehrlichiosis neoehrlichiosis neorickettsiosis anaplasmataceae rickettsiaceae coxiellaceae bartonellaceae brucellaceae francisellaceae pasteurellaceae legionellaceae chlamydiaceae mycoplasmataceae ureaplasmataceae spiroplasmataceae achaenoplasmataceae entomoplasmataceae haloplasmataceae tenericutes mollicutes bacteroidetes fusobacteria spirochaetes synergistetes deferribacteres thermotogae aquificae caldiserica caldithrix deferrisoma desulfurobacterium thermovibrio thermocrinis hydrogenobacter sulfurihydrogenibium persephonella hydrogenobaculum hydrothermus thermothrix sulfuricurvum sulfurimonas sulfurovum thiovulum thiomicrospira pseudomonas vibrio aeromonas photobacterium aliivibrio gracilibacillus halobacillus halomonas chromohalobacter cobetia salinicola kushneria halotalea halovibrio salinivibrio thioalkalivibrio thiohalophilus ectothiorhodospira halorhodospira thiorhodospira halochromatium thioflavicoccus marichromatium thiocapsa lamprocystis thiodictyon thiopedia amoebobacter thiocystis chromatium allo-chromatium thermochromatium rhabdochromatium thiobaca thiohalocapsa halocapsa thiolamprovum thioploca thiomargarita beggiatoa methylococcus methylomonas methylomicrobium methylosinus methylocystis methylocella methylocapsa methylacidiphilum verrucomicrobia planctomycetes chlamydiae lentisphaerae kiritimatiellaeota calditrichaeota fervidibacteria
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 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