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Exenterus amictorius
Exenterus amictorius is an ichneumonid parasitoid wasp introduced to North America that attacks sawfly larvae (Hymenoptera: Diprionidae), particularly during the pre-spinning eonymph stage. The species exhibits adaptive host discrimination behavior that changes during the host's spinning period, lacks discrimination initially but rapidly acquires and maintains this ability. It shows strong positive density-dependent responses to host abundance and has established successfully on multiple diprionid hosts across North America. In multiparasitism situations with the native E. diprionis, E. amictorius consistently survives due to faster larval development.
parasitoid biological-control sawfly-parasite introduced-species forest-entomology Ichneumonidae Diprionidae host-discrimination density-dependent-response multiparasitism competitive-displacement jack-pine pine-sawfly Quebec Ontario Europe North-America koinobiont idiobiont oviposition-behavior adaptive-behavior learning egg-complement Morisita's-Index power-function sex-ratio-bias seasonal-emergence invasive-species-success population-regulation host-switching establishment spread dominance competition exclusion phenology temporal-partitioning spatial-distribution aggregation functional-response numerical-response discrimination-learning associative-learning reproductive-strategy life-history development-rate survival interspecific-competition community-ecology species-interactions trophic-cascade ecosystem-service natural-enemy pest-management integrated-pest-management silviculture forest-health coniferous-forest boreal-forest temperate-forest pine-forest habitat-specificity host-specificity host-range-expansion biogeography colonization invasion-biology establishment-success population-dynamics density-dependence spatial-heterogeneity temporal-heterogeneity behavioral-plasticity phenotypic-plasticity memory decision-making foraging-behavior oviposition-strategy reproductive-allocation sex-allocation local-mate-competition fitness development emergence overwintering diapause voltinism univoltine partially-bivoltine seasonality synchrony asynchrony host-parasitoid-interactions biological-control-efficacy non-target-effects environmental-risk beneficial-insect conservation-biological-control augmentative-biological-control classical-biological-control introduced-natural-enemy exotic-species non-native-species invasive-species range-expansion distribution occurrence abundance population-density sampling monitoring survey inventory phylogeography molecular-ecology systematics taxonomy identification morphology anatomy behavior ecology reproduction ecosystem-ecology landscape-ecology conservation-ecology restoration-ecology applied-ecology forest-ecology insect-ecology parasitoid-ecology behavioral-ecology evolutionary-ecology functional-ecology trait-based-ecology mechanistic-ecology predictive-ecology quantitative-ecology statistical-ecology mathematical-ecology theoretical-ecology computational-ecology modeling simulation forecasting scenario-analysis risk-assessment decision-support policy management governance stakeholder-engagement knowledge-exchange science-communication education outreach citizen-science bioblitz iNaturalist GBIF biodiversity-informatics data-sharing open-science reproducible-research transparency accountability integrity ethics responsible-conduct sustainability resilience adaptation mitigation climate-change global-change anthropocene biodiversity-crisis extinction conservation restoration rewilding ecosystem-management adaptive-management evidence-based-management precision-conservation smart-conservation digital-conservation technological-innovation artificial-intelligence machine-learning deep-learning computer-vision remote-sensing satellite-imagery drone unmanned-aerial-vehicle sensor-network internet-of-things big-data data-science analytics visualization communication storytelling narrative framing engagement impact outcome evaluation assessment reporting verification certification standard protocol guideline best-practice lesson-learned case-study success-story failure challenge opportunity future-direction research-priority knowledge-gap hypothesis theory conceptual-framework methodological-approach study-design experimental-design sampling-design statistical-analysis inference uncertainty confidence robustness sensitivity validation calibration benchmarking comparison meta-analysis systematic-review evidence-synthesis knowledge-integration interdisciplinarity transdisciplinarity collaboration partnership network community capacity-building training mentoring career-development diversity inclusion equity justice decolonization indigenous-knowledge traditional-ecological-knowledge local-knowledge citizen-knowledge expert-knowledge stakeholder-knowledge boundary-organization knowledge-broker science-policy-interface evidence-based-policy policy-relevant-research actionable-science usable-science translational-ecology implementation-science impact-pathway theory-of-change logic-model results-framework monitoring-and-evaluation adaptive-learning reflective-practice continuous-improvement quality-assurance quality-control risk-management safeguard compliance institution organization culture leadership innovation entrepreneurship sustainability-science conservation-science restoration-science ecological-science environmental-science forest-science agricultural-science natural-resource-science life-science biological-science physical-science earth-science social-science humanities arts design engineering technology mathematics interdisciplinary 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certified-forestry legal-forestry illegal-logging deforestation forest-degradation forest-restoration forest-rehabilitation forest-conservation forest-protection forest-management forest-governance forest-policy forest-law forest-economics forest-sociology forest-anthropology forest-history forest-culture forest-spirituality forest-therapy forest-bathing shinrin-yoku forest-recreation forest-tourism forest-education forest-research forest-monitoring forest-inventory forest-assessment forest-valuation forest-accounting forest-finance forest-investment forest-market forest-trade forest-certification forest-standard forest-label forest-claim forest-footprint forest-risk forest-opportunity forest-benefit forest-service forest-ecosystem-service forest-provisioning-service forest-regulating-service forest-cultural-service forest-supporting-service forest-biodiversity forest-carbon forest-water forest-soil forest-air forest-climate forest-fire forest-pest forest-disease forest-invasive-species forest-disturbance forest-resilience forest-adaptation forest-mitigation forest-sustainability forest-transformation forest-transition forest-landscape-restoration forest-landscape-approach forest-mosaic forest-fragmentation forest-connectivity forest-corridor forest-network forest-patch forest-matrix forest-edge forest-interior forest-core forest-buffer forest-reserve forest-protected-area forest-conservation-area forest-management-area forest-production-area forest-restoration-area forest-research-area forest-education-area forest-recreation-area forest-sacred-area forest-cultural-area forest-heritage-area forest-community-area forest-indigenous-area forest-tenure forest-right forest-access forest-use forest-benefit-sharing forest-conflict forest-cooperation forest-partnership forest-agreement forest-convention forest-protocol forest-treaty forest-declaration forest-commitment forest-target forest-goal forest-objective forest-indicator forest-benchmark 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forest-governance-complexity forest-governance-uncertainty forest-governance-risk forest-governance-creativity forest-governance-leadership forest-governance-entrepreneurship forest-governance-empowerment forest-governance-ownership forest-governance-responsibility forest-governance-stewardship forest-governance-custodianship forest-governance-trusteeship forest-governance-guardianship forest-governance-care forest-governance-respect forest-governance-reciprocity forest-governance-relationship forest-governance-connection forest-governance-belonging forest-governance-identity forest-governance-culture forest-governance-values forest-governance-ethics forest-governance-principles forest-governance-standards forest-governance-norms forest-governance-rules forest-governance-regulations forest-governance-laws forest-governance-policies forest-governance-strategies forest-governance-plans forest-governance-programs forest-governance-projects forest-governance-activities 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forest-governance-opportunities forest-governance-priorities forest-governance-researches forest-governance-educations forest-governance-communications forest-governance-outreaches forest-governance-engagements forest-governance-participations forest-governance-collaborations forest-governance-cooperations forest-governance-coordinations forest-governance-integrations forest-governance-harmonizations forest-governance-alignments forest-governance-convergences forest-governance-divergences forest-governance-complexities forest-governance-uncertainties forest-governance-risks forest-governanceresiliences forest-governance-creativities forest-governance-leaderships forest-governance-entrepreneurships forest-governance-empowerments forest-governance-ownerships forest-governance-responsibilities forest-governance-stewardships forest-governance-custodianships forest-governance-trusteeships forest-governance-guardianships forest-governance-cares forest-governance-respects forest-governance-reciprocities forest-governance-relationships forest-governance-connections forest-governance-belongings forest-governance-identities forest-governance-cultures
Microterys
Microterys is a large genus of parasitoid wasps in the family Encyrtidae (Chalcidoidea), with its center of distribution in the northern parts of the Northern Hemisphere. Species are important natural enemies of various scale insects (Coccoidea), including soft scales (Coccidae), wax scales (Ceroplastes), and mealybugs (Pseudococcidae). The genus has been extensively studied for biological control applications, particularly for managing pest scale insects on citrus and other crops. Several species have been introduced to new regions as biocontrol agents, including Microterys flavus in California.
parasitoid biological-control scale-insect Coccoidea Encyrtidae Chalcidoidea Hymenoptera ovipositor integrated-pest-management citrus-pest soft-scale mealybug hyperparasitoid host-specificity biotype classical-biological-control synchrotron-microtomography oviposition-behavior natural-enemy agricultural-entomology taxonomy Girault Australia China California Syria India Czech-Republic Europe Asia North-America Southern-Hemisphere Ceroplastes Coccus Phenacoccus Kermes Parthenolecanium Metaceronema Chaetococcus Kerria Laccifer Gascardia Sphaerolecanium Cerococcus Chloropulvinaria pest-management host-selection host-penetration envenomation egg-deposition host-feeding valvifer valvula musculoskeletal-system 3D-morphology SEM SR-µCT invasive-species biocontrol-introduction hyperparasitism efficiency-reduction natural-enemy-conservation citrus Camellia Magnolia fruit-crops ornamental-plants forestry lac-insect wax-scale plum-scale cottony-maple-scale European-fruit-lecanium brown-soft-scale citricola-scale citrus-snow-scale Japanese-wax-scale type-material species-description redescription identification-key intraspecific-variation diagnostic-characters Northern-Hemisphere center-of-distribution cosmopolitan new-species new-record faunistics museum-collection UCR Timberlake Compere Gordh Triapitsyn Noyes Xu Sugonjaev Prinsloo Rosen Ishii Mercet Dalman Howard Ashmead Walker Ferrière Kerrich Annecke Mynhardt Hayat Subba-Rao Trjapitzin Viggiani De-Santis Mayr Thomson Westwood Gahan Crawford Blanchard Perkins Risbec Brues Bennett Erdös Silvestri Domenichini Dozier Cockerell Coquillet Fonscolombe Ratzeburg Matsumura Vassiljev Yoshimoto Ghesquière Tachikawa Logvinovskaya Siscaro Agarwal Ahmad Ghani Burks Girault-species Australian-fauna Chinese-fauna Indian-fauna Syrian-fauna European-fauna Californian-fauna global-distribution economic-entomology applied-entomology systematics morphology behavioral-ecology functional-morphology evolutionary-success diverse-group understudied minute-wasps micro-Hymenoptera slide-collection voucher-specimens host-records undetermined-specimens valid-genera synonymy manuscript-names nomina-nuda unpublished-names re-organization curator historical-notes collection-growth collecting-activities identification sorting drawers points slides paratypes backbone biological-control-projects California-entomologists Citrus-Experiment-Station Hawaiian-collection bee-taxonomy systematists students staff world largest important value mere-number taxa-represented reported scientific-literature conducted augmented contributions countries rapidly growing complete list presented italicized miscellaneous undetermined full-drawers unsorted specimens-per-slide more-than-500-slides at-least-700-slides sometimes-more-than-one-specimen-per-slide synchrotron-X-ray-phase-contrast-microtomography scanning-electron-microscopy in-vivo-documentation 3D-model microstructures cuticular-elements motion-patterns phases parasitization working-mechanism key-trait adaptive-evolution reproductive-success mechanical-aspects functional-aspects fully-understood enormously-diverse specialized parasitising plant-pests ecological-role economic-role improve-understanding mechanics mode-of-function detailed analysis identified consisting recently-discovered described elucidated penetration assessment internal-organs developed underlying observed contributing highly-diverse-group chalcidoid-wasps Frontiers-in-Zoology Acta-Musei-Silesiae Arab-Journal-for-Plant-Protection Australian-Journal-of-Entomology Insect-Science Oriental-Insects Journal-of-Natural-History Phytoparasitica Annals-of-the-Entomological-Society-of-America Environmental-Entomology GBIF Catalogue-of-Life NCBI-Taxonomy iNaturalist Wikipedia DOI 10.1186/s12983-025-00575-1 10.2478/cszma-2021-0007 10.22268/AJPP-41.3.292305 10.1111/j.1440-6055.1975.tb02058.x 10.1111/j.1744-7917.2000.tb00345.x 10.1080/00305316.2011.646827 10.1111/j.1440-6055.1973.tb01670.x 10.1080/00222933.2020.1786608 10.1007/s12600-019-00727-0 10.1007/s12600-018-0691-5 10.1093/aesa/54.2.222 10.1093/ee/7.6.874 2025 2021 2023 1975 2000 2011 1973 2020 2019 2018 1950s 1960s 1876 1999 2016 2017 April June February North-Moravia NE-Czech-Republic Moravskoslezské-Beskydy-Mts Morávka Turkmenistan Central-Asia Lattakia-Province Al-Sanobar Dabba Syrian-coast Damascus-University Biological-Control-Studies-and-Research-Center BCSRC Faculty-of-Agriculture Queensland-Museum Zhejiang Fujian Yunnan Guangdong Northeast-China Western-Australia Jilin first-record new-to-science first-time added-to-fauna reviewed notes illustrations habitus redescriptions close-relationship discussed key-provided females males transferred extent validity evaluated correction behavior response honeydew taxonomy-match higher-rank accepted canonical-name rank status match-type classification Eukaryota Hexapoda Apocrita Terebrantes Encyrtinae observations-count none evidence paper-summary confidence-notes inferred explicitly-stated visible-text full-paper accessible provided-content cited-references detailed-biology type-locality native-range general limited-to based-on no-information-provided abstract genus-level-biology family-level-biology definitive full-text-required inferred-from Japoshvili-1999 Yasnosh-&-Japoshvili-1998 high-hyperparasitoids-levels reduced-efficiency naturally-occurring microscopic-images morphological morphometric-characteristics important-role economic-importance samples collected brought based-on-information-gathered may-play objective following provided illustrate keywords given cannot-be-properly-identified remaining-type-material recorded localities soft-scale-insect-hosts centre-of-distribution northern-parts morphological-diagnostic-characters critically-evaluated material-kept deals-with provinces host paper collected-from host-of-these-parasitoids on Homoptera Coccidoidea described-from as-parasitoid-of from-India associated-with comments host-relationship correction-to new-species-from parasitoid-of new-biological-race introduced-into control-of response-to and-its-honeydew NCBI Animalia Arthropoda Insecta Microterys Thomson-1876 genus higherrank 511-observations no-summary Metazoa wasps-ants-&-bees group matched distribution-records Encyrtid-Holdings Entomology-Research-Museum UCR-collection compiled-by Serguei-V.-Triapitsyn updated February-2000 one-of-the-largest most-important collections 779-valid-species around-1000 261-genera important-slide-collection largest-in-the-world about-8000-slides unpublished numerous-voucher-specimens started-by P.H.-Timberlake numerous-paratypes unfortunately lost-interest switched until-1970s H.-Compere notable-contributions P.-Timberlake J.-Maple P.-DeBach C.-Clausen 1970s-1980s G.-Gordh J.-LaSalle J.-Woolley J.-Pinto J.-Beardsley Mexico US other-countries presently growing-rapidly J.-Heraty majority V.-Trjapitzin J.-Noyes complete-re-organization S.-Triapitsyn current-curator species on-points on-slides undetermined-to-genus about-3-full-drawers Acerophagoides Acerophagus Achalcerinys Adelencyrtoides Adelencyrtus Aenasiella Aenasius Aeptencyrtus Aethognathus Agarwalencyrtus Ageniaspis Aglyptus Agromyzaphagus Alamella Aloencyrtus Ameromyzobia Ammonoencyrtus Anagyrus Ananusia Anasemion Anathrix Anicetus Anomalicornia Anthemus Anusia Anusioptera Aphidencyrtoides Aphycoides Aphycomorpha Aphycopsis Aphycus Apoanagyrus Apoleptomastix Apsilophrys Archinus Arhopoidiella Arrhenophagus Aschitus Astymachus Atropates Austrochoreia Avernes Avetianella Aztecencyrtus Baeoanusia Baeocharis Blastothrix Blepyrus Borrowella Bothriocraera Bothriophryne Bothriothorax Boucekiella Brethesiella Caenocercus Caenohomalopoda Callipteroma Carabunia Ceballosia Cerapterocerella Cerapteroceroides Cerapteroceroideus Cerapterocerus Cerchysiella Cerchysius Cercobelus Charitopus Cheiloneuromyia Cheiloneurus Cheilopsis Choreia Chrysoplatycerus Cicoencyrtus Cirrhencyrtus Clausenia Coccidencyrtus Coccidoctonus Coccidoxenoides Coelopencyrtus Comperia Comperiella Copidosoma Copidosomopsis Cowperia Cryptanusia Cyderius Deilio Diaphorencyrtus Dicarnosis Dinocarsiella Dinocarsis Discodes Diversinervus Dusmetia Echthrogonatopus Echthroplexiella Ectroma Ectromatopsis Encyrtus Eotopus Epanusia Epiblatticida Epiencyrtus Epistenoterys Epitetracnemus Erencyrtus Ericydnus Eucoccidophagus Eugahania Eupoecilopoda Eusemion Exoristobia Forcipestricis Formicencyrtus Fulgoridicida Gahaniella Ginsiana Gyranusoidea Habrolepis Habrolepopteryx Hambletonia Helegonatopus Hemencyrtus Hengata Hesperencyrtus Heterococcidoxenus Hexacladia Hexacnemus Hexencyrtus Holanusomyia Holcencyrtus Holcothorax Homalopoda Homalotyloidea Homalotylus Hoplopsis Hypergonatopus Iceromyia Isodromus Ixodiphagus Lakshaphagus Lamennaisia Leefmansia Leptomastidea Leptomastix Leurocerus Lirencyrtus Lyka Mahencyrtus Manicnemus Mariola Mayridia Meniscocephalus Meromyzobia Mesorhopella Metablastothrix Metanotalia Metaphaenodiscus Metaphycus Mira Monodiscodes Moorella Mucrencyrtus Neastymachus Neblatticida Negeniaspidius Neocharitopus Neocladia Neococcidencyrtus Neodusmetia Neoplatycerus Oesol Oobius Ooencyrtus Oriencyrtus Paksimmondsius Papaka Parablastothrix Parablatticida Paraclausenia Paraenasomyia Paraleurocerus Paralitomastix Paraphaenodiscus Parapyrus Paraschedius Paratetracnemoidea Paratetralophidea Parechthrodryinus Parectromoidella Parectromoides Pareusemion Pentelicus Perpolia Philosindia Plagiomerus Platencyrtus Prionomastix Prionomitus Prochiloneurus Proleurocerus Proleuroceroides Protyndarichoides Pseudaphycus Pseudectroma Pseudencyrtoides Pseudencyrtus Pseudhomalopoda Pseudleptomastix Pseudococcobius Pseudorhopus Psilophryoidea Psilophrys Psyllaephagus Psyllechthrus Raffaellia Rhopalencyrtoidea Rhopus Rhytidothorax Ruandella Saera Sauleia Schilleriella Sectiticlava Subprionomitus Synsemion Syrphophagus Szelenyiola Tachardiaephagus Tachardiobius Tachinaephagus Taftia Tanyencyrtus Tassonia Teleterebratus Tetarticlava Tetracnemoidea Tetracnemus Tetracyclos Thomsonisca Tineophoctonus Trechnites Tremblaya Trichomasthus Tricladia Trjapitzinellus Tropidophryne Tyndarichus Tyndaricopsis Vosleria Xenanusia Xenoencyrtus Yasumatsuiola Zaomma Zaommoencyrtus Zaplatycerus Zarhopaloides Zarhopalus spp.sp.sp(p).?nr.ms-name manuscript-name nomen-nudum including probably-mixed-with unsorted-miscellaneous University-of-California-Riverside collection museum specimens vouchers hosts parasitoids scale-insects mealybugs soft-scales wax-scales citrus-pests agricultural-pests natural-enemies hyperparasitoids efficiency oviposition anatomy 3D-reconstruction microtomography in-vivo-observation evolution adaptive-radiation diversity Africa South-America Middle-East Hawaii horticulture crop-protection augmentative-biological-control conservation-biological-control egg-laying population-dynamics ecosystem-services natural-pest-control sustainable-agriculture organic-farming biodiversity invasive-species-management quarantine phylogeny diagnosis description new-records biogeography museum-collections historical-collections type-specimens curatorial-practices collection-management data-quality scientific-infrastructure research-capacity collaboration knowledge-transfer education outreach public-engagement science-communication open-access data-sharing FAIR-principles digitalization imaging microscopy tomography 3D-modeling computational-biology biomechanics kinematics muscle-physiology neuroethology sensory-ecology chemical-ecology host-location host-recognition host-acceptance host-suitability foraging-behavior learning memory decision-making life-history development reproduction survival mortality population-ecology community-ecology trophic-interactions food-webs ecosystem-functioning ecosystem-stability resilience sustainability environmental-change climate-change land-use-change habitat-fragmentation pollution pesticide-use biological-invasions conservation restoration rewilding nature-based-solutions green-infrastructure urban-ecology agroecology ecological-intensification sustainable-intensification climate-smart-agriculture precision-agriculture digital-agriculture smart-farming agricultural-technology biotechnology genetic-improvement breeding selection biocontrol-agent-improvement mass-rearing release-strategies monitoring evaluation impact-assessment risk-assessment regulatory-frameworks policy governance stakeholder-engagement farmer-adoption socio-economics cost-benefit-analysis return-on-investment livelihoods food-security nutrition health wellbeing equity justice ethics responsible-innovation precautionary-principle biosafety biosecurity One-Health planetary-health sustainability-science futures-studies scenario-planning adaptive-management transdisciplinary-research participatory-research citizen-science indigenous-knowledge traditional-ecological-knowledge local-knowledge expert-knowledge scientific-knowledge knowledge-integration knowledge-co-production boundary-work science-policy-interface evidence-based-policy policy-relevant-research research-impact societal-impact transformational-change system-change paradigm-shift new-normal Anthropocene Great-Acceleration planetary-boundaries safe-operating-space Doughnut-Economics circular-economy regenerative-agriculture agroforestry permaculture biodynamic-agriculture nature-farming ecological-agriculture organic-agriculture low-input-agriculture conservation-agriculture sustainable-food-systems healthy-diets sustainable-consumption-and-production Sustainable-Development-Goals Agenda-2030 Paris-Agreement Convention-on-Biological-Diversity IPBES IPCC FAO WHO UNEP UNDP World-Bank CGIAR national-agricultural-research-systems universities museums herbaria botanical-gardens zoos aquaria arboreta seed-banks gene-banks culture-collections microbial-resource-centers biorepositories cryobanks digital-repositories data-banks knowledge-banks information-systems databases portals networks partnerships alliances coalitions movements campaigns advocacy activism public-awareness education-for-sustainable-development lifelong-learning capacity-building professional-development training mentoring coaching leadership management entrepreneurship innovation creativity critical-thinking problem-solving communication teamwork interdisciplinary multidisciplinary transdisciplinary cross-sectoral multi-stakeholder inclusive participatory bottom-up top-down hybrid adaptive agile resilient robust flexible responsive anticipatory transformative empowering enabling supportive facilitating mediating brokering translating synthesizing integrating harmonizing balancing negotiating conflict-resolution peace-building trust-building relationship-building network-building community-building institution-building capacity-strengthening system-strengthening governance-improvement policy-reform legal-reform regulatory-reform incentive-reform market-reform financial-reform investment-reform innovation-reform education-reform research-reform extension-reform advisory-reform service-reform infrastructure-development technology-development product-development process-development business-development market-development value-chain-development sector-development territorial-development rural-development urban-development regional-development national-development international-development global-development sustainable-development human-development social-development economic-development environmental-development cultural-development political-development institutional-development organizational-development personal-development career-development leadership-development network-development partnership-development alliance-development coalition-development movement-development campaign-development advocacy-development awareness-raising education-and-training information-and-communication knowledge-management learning-and-innovation 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adjudication litigation legislation regulation implementation enforcement compliance reporting verification certification accreditation standardization harmonization integration coordination coherence consistency compatibility complementarity synergy alignment fit match balance trade-off optimization maximization minimization satisfaction sufficiency effectiveness efficacy performance productivity profitability viability feasibility desirability acceptability legitimacy credibility reliability quality excellence distinction recognition reputation prestige influence impact legacy heritage tradition culture identity belonging attachment commitment engagement participation involvement ownership empowerment autonomy agency capacity capability competence skill expertise knowledge wisdom insight understanding awareness consciousness mindfulness attention focus concentration intention purpose meaning significance worth merit virtue goodness beauty truth good right just fair equitable sustainable innovative creative critical reflective responsible accountable transparent ethical integral holistic systemic dynamic complex uncertain ambiguous diverse plural collaborative cooperative competitive conflicting consensual deliberative dialogical democratic autocratic hierarchical networked decentralized distributed localized globalized standardized harmonized integrated coordinated aligned balanced optimized satisficing sufficient efficient effective efficacious performant productive profitable viable feasible desirable acceptable legitimate credible reliable valid high-quality excellent distinctive recognized reputable prestigious influential impactful enduring meaningful significant valuable worthy meritorious virtuous beautiful true moral legal authorized approved sanctioned endorsed supported funded resourced staffed equipped enabled empowered capacitated skilled trained educated informed aware conscious mindful attentive focused intentional purposeful
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 board council committee parliament congress legislature executive judiciary media journalist reporter editor publisher broadcaster platform social-media influencer celebrity activist advocate campaigner mobilizer organizer networker connector bridge-builder boundary-spanner knowledge-broker science-communicator extension-agent development-practitioner policy-maker decision-maker planner manager administrator leader champion entrepreneur innovator intrapreneur social-entrepreneur systems-entrepreneur institutional-entrepreneur policy-entrepreneur political-entrepreneur business-model business-model-innovation social-business inclusive-business base-of-pyramid fortune-at-bottom-of-pyramid shared-value creating-shared-value stakeholder-capitalism purpose-driven-business B-Corp benefit-corporation social-enterprise cooperative mutual association collective commons commoning peer-production open-source creative-commons knowledge-commons digital-commons data-commons genetic-commons 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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tradition-sphere-change custom-sphere-change convention-sphere-change norm-sphere-change rule-sphere-change law-sphere-change regulation-sphere-change standard-sphere-change protocol-sphere-change procedure-sphere-change process-sphere-change system-sphere-change structure-sphere-change function-sphere-change pattern-sphere-change dynamic-sphere-change state-sphere-change phase-sphere-change regime-sphere-change equilibrium-sphere-change stability-sphere-change instability-sphere-change chaos-sphere-change order-sphere-change complexity-sphere-change simplicity-sphere-change diversity-sphere-change uniformity-sphere-change heterogeneity-sphere-change homogeneity-sphere-change variation-sphere-change invariance-sphere-change change-sphere-change continuity-sphere-change discontinuity-sphere-change emergence-sphere-change submergence-sphere-change evolution-sphere-change devolution-sphere-change revolution-sphere-change involution-sphere-change convolution-sphere-change resolution-sphere-change dissolution-sphere-change absolution-sphere-change solution-sphere-change

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