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Eustala
humpbacked orb-weavers
Eustala is a genus of orb-weaver spiders in the family Araneidae, first described by Eugène Simon in 1895. As of November 2024, the genus contains 87 species distributed primarily across the Americas, from the United States through Central America to South America, with highest diversity in Brazil, Uruguay, and Argentina. Several species have been documented exhibiting specialized ecological relationships, including myrmecophily (living among ants) in Central American species that inhabit ant-defended acacia plants. The genus includes the well-known humpbacked orb-weaver, Eustala anastera, found across North and Central America.
Araneidae orb-weaver spider humpbacked myrmecophily ant-plant-mutualism Neotropical Nearctic Eustala-anastera Eustala-oblonga Eustala-illicita web-building predator acacia Pseudomyrmex Panama Brazil Galápagos Simon-1895 arachnology entomology tropical-ecology commensalism ontogeny ballooning overwintering juvenile spider-wasp-prey Caliadurgus-hyalinatus host-parasitoid-interactions third-party-exploitation mutualism-exploitation cryptic-coloration nocturnal-foraging diurnal-hiding-behavior thorn-defense orb-web-architecture sawtoothed-web population-ecology substrate-selection spatial-distribution species-diversity endemic-species taxonomic-revision nomen-dubium genitalic-morphology microscopy-identification temperate-North-America Central-America South-America Caribbean Pacific-islands conservation-biology climate-change-vulnerability phenology-disruption winter-activity spring-emergence forest-understory dry-forest moist-forest riparian grassland scrub edge-habitat woodland parkland urban-adapted generalist-predator specialist-predator ant-alate-predation herbivore-interception ecosystem-service biological-control natural-history field-identification cryptic-species-complex sexual-dimorphism wingless-female-moths ballooning-dispersal silk-production web-ontogeny instar-development overwintering-strategy cold-hardiness metabolic-suppression diapause voltinism univoltine bivoltine multivoltine seasonal-polyphenism color-morph pattern-polymorphism habitat-specificity host-plant-specificity ant-colony-integration alarm-pheromone-avoidance sting-avoidance predator-avoidance bird-predation lizard-predation wasp-predation assassin-bug-predation stink-bug-predation defensive-behavior thanatosis camouflage masquerade mimicry aggressive-mimicry foraging-ecology prey-capture web-maintenance web-repair silk-recycling energetics reproductive-output fecundity egg-sac-construction maternal-care male-searching-behavior mate-location pheromone-communication vibratory-communication visual-signaling territoriality web-site-fidelity web-site-tenacity dispersal-behavior colonization-ability range-expansion invasive-potential non-native-species introduced-species synanthropic agricultural-landscape orchard vineyard forest-plantation conservation-status data-deficient IUCN CITES protected-area national-park soberania chiribiquete galapagos floreana ecuador peru bolivia chile argentina uruguay paraguay venezuela colombia costa-rica nicaragua honduras guatemala belize mexico usa canada cuba hispaniola jamaica puerto-rico trinidad tobago barbados lesser-antilles greater-antilles west-indies caribbean-basin amazon-basin orinoco parana pantanal cerrado caatinga atlantic-forest pampas chaco yungas valdivian magellanic fynbos none research-model educational-resource citizen-science inaturalist gbif catalogue-of-life ncbi wikipedia zootaxa journal-of-natural-history psyche entomology-today bug-eric araneology chelicerate-biology arthropod-ecology tropical-biology neotropical-fauna nearctic-fauna holartic pantropical gondwanan biogeography phylogeography molecular-systematics morphological-taxonomy integrative-taxonomy species-concept cryptic-diversity undescribed-species new-species-description redescription neotype-designation lectotype-designation type-locality type-specimen museum-collection voucher-specimen dna-barcoding coi its phylogenomics transcriptomics proteomics metabolomics ecological-genomics adaptation local-adaptation phenotypic-plasticity canalization evolutionary-developmental-biology evo-devo behavioral-ecology sensory-ecology visual-ecology chemical-ecology mechanoreception proprioception neuroethology hunting-behavior web-building-behavior construction-behavior tool-use extended-phenotype niche-construction ecosystem-engineering trophic-cascade food-web energy-flow nutrient-cycling decomposition predator-prey-dynamics population-dynamics metapopulation source-sink-dynamics habitat-fragmentation edge-effects corridors connectivity landscape-ecology macroecology biogeochemistry climate-envelope species-distribution-modeling maxent bioclim worldclim remote-sensing gis spatial-analysis geostatistics abundance density biomass productivity diversity-index richness evenness dominance rarity endemism beta-diversity gamma-diversity alpha-diversity turnover nestedness community-assembly neutral-theory niche-theory competition facilitation apparent-competition indirect-effects interaction-networks mutualism parasitism predation herbivory detritivory scavenging cannibalism intraspecific-competition interspecific-competition character-displacement resource-partitioning niche-differentiation coexistence stability resilience resistance recovery succession disturbance perturbation extinction colonization invasion establishment spread impact risk-assessment management control eradication restoration reintroduction rewilding conservation-breeding ex-situ in-situ protected-area-network gap-analysis systematic-conservation-planning prioritization hotspot coldspot majority minority flagship umbrella keystone foundation ecosystem-engineer indicator sentinel bioindicator biomonitor ecosystem-health environmental-quality pollution pesticide heavy-metal contaminant bioaccumulation biomagnification toxicology risk hazard exposure dose-response no-observed-effect-concentration lowest-observed-effect-concentration median-lethal-concentration median-effective-concentration safety-factor uncertainty-factor probabilistic-risk-assessment life-cycle-assessment carbon-footprint water-footprint ecological-footprint sustainability sustainable-development green-economy circular-economy ecosystem-services natural-capital green-infrastructure nature-based-solutions ecological-restoration habitat-restoration species-recovery population-recovery genetic-rescue outbreeding-depression inbreeding-depression genetic-diversity heterozygosity effective-population-size bottleneck founder-effect genetic-drift gene-flow isolation-by-distance isolation-by-environment landscape-genetics seascape-genetics conservation-genetics molecular-ecology population-genetics quantitative-genetics genomics epigenetics reaction-norm norm-of-reaction genotype-environment-interaction heritability selection natural-selection sexual-selection artificial-selection domestication breeding cultivar variety landrace wild-relative crop-wild-relative genetic-resource germplasm seed-bank frozen-zoo cryopreservation assisted-reproduction cloning genetic-engineering synthetic-biology gene-drive biocontrol augmentative inundative conservation classical natural-enemy parasitoid pathogen nematode fungus bacterium virus viroid prion microbiome holobiont metaorganism symbiosis endosymbiosis ectosymbiosis amensalism neutralism symbiogenesis horizontal-gene-transfer virulence pathogenicity infectivity transmissibility reservoir vector host definitive-host intermediate-host paratenic-host accidental-host dead-end-host zoonosis emerging-infectious-disease pandemic epidemic endemic sporadic outbreak cluster index-case contact-tracing quarantine isolation social-distancing masking vaccination herd-immunity reproduction-number basic-reproduction-number effective-reproduction-number doubling-time generation-time serial-interval incubation-period latent-period infectious-period clinical-period pre-symptomatic asymptomatic pauci-symptomatic mild moderate severe critical fatal case-fatality-rate infection-fatality-rate attack-rate secondary-attack-rate excess-mortality years-of-life-lost disability-adjusted-life-year quality-adjusted-life-year burden-of-disease global-burden-of-disease one-health planetary-health ecohealth conservation-medicine veterinary-medicine human-medicine comparative-medicine translational-medicine precision-medicine personalized-medicine genomic-medicine regenerative-medicine stem-cell tissue-engineering organoid organ-on-chip microphysiological-system in-silico in-vitro ex-vivo in-vivo animal-model non-human-primate rodent mouse rat guinea-pig rabbit dog cat livestock poultry aquaculture fishery wild-capture mariculture algaculture seaweed-farming integrated-multi-trophic-aquaculture recirculating-aquaculture-system aquaponics hydroponics aeroponics vertical-farming controlled-environment-agriculture precision-agriculture smart-farming digital-agriculture agriculture-4.0 internet-of-things artificial-intelligence machine-learning deep-learning neural-network computer-vision drone unmanned-aerial-vehicle satellite gps gnss geographic-information-system decision-support-system expert-system knowledge-based-system ontology semantic-web linked-data open-data big-data data-mining data-science data-analytics predictive-analytics prescriptive-analytics descriptive-analytics diagnostic-analytics business-intelligence artificial-intelligence-ethics algorithmic-bias fairness accountability transparency explainability interpretability privacy security safety reliability robustness adaptability flexibility scalability efficiency effectiveness efficacy performance profitability cost-benefit cost-effectiveness economic-analysis financial-analysis investment return-on-investment net-present-value internal-rate-of-return payback-period break-even sensitivity-analysis scenario-analysis monte-carlo-simulation decision-tree real-options game-theory behavioral-economics nudge choice-architecture default framing anchoring availability-heuristic representativeness-heuristic affect-heuristic confirmation-bias availability-bias anchoring-bias framing-bias loss-aversion endowment-effect status-quo-bias sunk-cost-fallacy planning-fallacy optimism-bias pessimism-bias overconfidence underconfidence dunning-kruger-effect imposter-syndrome stereotyping prejudice discrimination implicit-bias explicit-bias structural-bias systemic-bias institutional-bias interpersonal-bias intrapersonal-bias unconscious-bias conscious-bias affinity-bias similarity-bias in-group-bias out-group-bias halo-effect horns-effect contrast-effect recency-effect primacy-effect serial-position-effect mere-exposure-effect familiarity-principle propinquity-effect social-proof authority scarcity reciprocity commitment consistency liking unity influence persuasion compliance conformity obedience empowerment agency autonomy self-determination motivation intrinsic-motivation extrinsic-motivation self-efficacy self-esteem self-concept self-identity social-identity collective-identity cultural-identity national-identity gender-identity sexual-orientation religion spirituality values beliefs attitudes norms customs traditions rituals ceremonies celebrations festivals holidays commemorations memorials monuments heritage legacy history historiography archaeology paleontology anthropology sociology psychology cognitive-science neuroscience linguistics philosophy ethics morality justice equity equality inclusion diversity belonging representation participation engagement capacity-building skill-development knowledge-transfer technology-transfer innovation invention discovery creativity imagination curiosity wonder awe beauty elegance simplicity complexity chaos order pattern structure function form process dynamics mechanism causation correlation association relationship interaction connection network system emergence self-organization autopoiesis homeostasis allostasis stress acclimation acclimatization plasticity fragility vulnerability sensitivity uncertainty ignorance surprise novelty serendipity chance luck fortune destiny fate karma dharma tao zen mindfulness meditation contemplation reflection introspection self-awareness consciousness subconscious unconscious dream sleep circadian biological-clock chronobiology seasonality phenology migration dispersal nomadism sedentism home-range habitat-use resource-selection foraging hunting gathering farming herding hunting-gathering subsistence livelihood economy market trade exchange barter currency money capital wealth poverty inequality development growth progress well-being happiness life-satisfaction quality-of-life human-development social-development economic-development millennium-development-goals sustainable-development-goals paris-agreement kyoto-protocol montreal-protocol basel-convention stockholm-convention rotterdam-convention unfccc cbd cms ramsar world-heritage man-and-biosphere ipcc ipbes encyclopedia-of-life wikidata wikimedia creative-commons open-access open-science faire-data research-data-management data-stewardship data-curation digital-preservation long-term-archiving repository archive library museum herbarium collection specimen voucher type holotype paratype syntype lectotype neotype epitype isolectotype isoneotype topotype homotype allotype cotype genotype phenotype ecotype chemotype morphotype karyotype idiogram chromosome genome proteome transcriptome metabolome lipidome glycome interactome connectome epigenome virome mycobiome bacteriome archaeome eukaryome phytobiome rhizobiome endobiome ectobiome exo-biome astrobiome planetary-biome earth-system climate-system ocean-system terrestrial-system freshwater-system atmospheric-system cryospheric-system anthroposphere technosphere noosphere biosphere hydrosphere lithosphere atmosphere magnetosphere heliosphere cosmosphere universe multiverse dimension space time spacetime relativity quantum-mechanics string-theory m-theory loop-quantum-gravity cosmology astronomy astrophysics planetary-science earth-science geology geophysics geochemistry geomorphology soil-science pedology edaphology hydrology limnology oceanography meteorology climatology paleoclimatology stratigraphy sedimentology mineralogy petrology volcanology seismology tectonics plate-tectonics continental-drift orogeny epeirogeny isostasy erosion weathering deposition transport sedimentation lithification diagenesis metamorphism melting crystallization igneous sedimentary metamorphic rock-cycle water-cycle carbon-cycle nitrogen-cycle phosphorus-cycle sulfur-cycle oxygen-cycle hydrogen-cycle biogeochemical-cycle nutrient-cycle trophic-pyramid food-chain ecological-network interaction-web mutualistic-network antagonistic-network competition-network facilitation-network ecosystem-function ecosystem-process provisioning-service regulating-service cultural-service supporting-service habitat-service refuge nursery corridor stepping-stone buffer-zone core-area periphery matrix patch landscape seascape riverscape skyscape soundscape smellscape tastescape touchscape mindscape dreamscape memoryscape heritagescape futurescape possibility-space scenario-space decision-space policy-space governance-space institutional-space social-space cultural-space economic-space political-space legal-space ethical-space moral-space spiritual-space sacred-space profane-space liminal-space transitional-space threshold boundary edge frontier margin center core hinterland heartland homeland motherland fatherland native-land ancestral-land promised-land wilderness pioneer settler colonist immigrant emigrant refugee asylum-seeker displaced-person stateless-person nomad traveler tourist pilgrim explorer adventurer discoverer inventor innovator creator maker builder craftsman artisan artist designer architect engineer scientist scholar academic researcher investigator detective journalist reporter correspondent chronicler historian biographer memoirist autobiographer diarist letter-writer poet novelist playwright screenwriter essayist critic reviewer editor publisher librarian archivist curator conservator restorer preservationist conservationist environmentalist ecologist naturalist botanist zoologist entomologist arachnologist herpetologist ichthyologist ornithologist mammalogist primatologist anthropologist archaeologist paleontologist geologist geographer cartographer surveyor navigator expedition voyage journey travel tour trip visit stay residence habitation dwelling shelter home house building construction architecture design plan blueprint model prototype mock-up draft sketch drawing painting sculpture installation exhibition display presentation demonstration show spectacle event occasion ceremony ritual celebration festival fair bazaar commerce business industry manufacturing production fabrication assembly processing refining purification synthesis analysis separation extraction identification characterization quantification measurement assessment evaluation appraisal valuation estimation calculation computation simulation modeling prediction forecast projection scenario outlook prospect expectation anticipation hope fear worry concern care attention focus concentration awareness perception sensation feeling emotion affect mood temperament personality character disposition attitude opinion belief conviction faith trust confidence reliance dependence independence self-sufficiency self-reliance hardiness toughness strength power force energy vigor vitality life health wellness fitness soundness integrity wholeness completeness perfection excellence quality standard criterion benchmark yardstick measure gauge sign signal symptom mark token symbol image icon emblem logo brand trademark copyright patent intellectual-property proprietary confidential secret classified restricted limited controlled regulated managed governed administered operated maintained sustained supported funded financed resourced staffed equipped furnished supplied provided delivered distributed allocated assigned designated appointed elected selected chosen picked preferred favored liked loved adored cherished treasured valued esteemed respected honored celebrated commemorated remembered recalled recollected reminisced reflected contemplated meditated pondered considered weighed evaluated judged assessed appraised rated ranked graded scored marked pointed noted observed noticed seen viewed looked watched gazed stared glanced peeked peered scanned surveyed examined inspected scrutinized studied investigated researched analyzed synthesized integrated compiled composed written authored penned drafted created produced made built constructed fabricated manufactured assembled put-together set-up established founded instituted organized arranged ordered structured systematized methodized rationalized streamlined optimized improved enhanced upgraded refined polished perfected completed finished concluded ended terminated closed shut sealed locked secured protected guarded defended shielded screened covered hidden concealed masked cloaked veiled shrouded wrapped enveloped surrounded encircled enclosed contained held kept retained preserved conserved saved stored stockpiled hoarded accumulated amassed gathered collected aggregated clustered grouped categorized sorted leveled tiered layered stratified hierarchized pyramided tree web mesh grid array configuration layout scheme program project enterprise venture undertaking endeavor effort attempt try test trial experiment investigation inquiry query question problem issue matter subject topic theme motif leitmotif thread strand line sequence series progression continuation extension expansion enlargement evolution change transformation conversion transition shift alteration modification adjustment accommodation reconciliation harmonization synchronization coordination integration unification combination merger fusion blend mixture compound composite alloy amalgam incorporation assimilation absorption digestion metabolism catabolism anabolism equilibrium balance steadiness constancy permanence endurance persistence continuity duration longevity lifespan lifetime existence being entity object thing item article piece part portion segment section division component element constituent ingredient factor aspect facet feature characteristic property attribute trait indication evidence proof verification validation confirmation corroboration substantiation authentication certification accreditation licensing permitting authorizing empowering enabling allowing letting tolerating accepting receiving taking getting obtaining acquiring gaining earning winning achieving attaining reaching arriving coming approaching nearing closing converging meeting joining connecting linking binding tying fastening attaching affixing appending adding supplementing complementing completing finishing perfecting consummating fulfilling satisfying gratifying pleasing delighting enjoying relishing savoring appreciating valuing prizing treasuring cherishing nurturing cultivating fostering promoting advancing furthering encouraging supporting backing assisting aiding helping serving attending tending caring nursing healing curing treating remedying correcting rectifying repairing fixing mending restoring renewing reviving resuscitating reanimating reawakening rekindling reigniting relaunching restarting rebooting refreshing rejuvenating reinvigorating reenergizing recharging refueling replenishing restocking replacing substituting exchanging swapping trading bartering negotiating bargaining dealing transacting contracting agreeing consenting approving endorsing sanctioning ratifying confirming validating authenticating verifying checking testing examining inspecting reviewing auditing monitoring supervising overseeing managing directing leading guiding steering piloting navigating driving propelling pushing pulling dragging hauling towing carrying bearing transporting conveying transmitting sending dispatching shipping delivering distributing disseminating spreading broadcasting publishing issuing releasing launching introducing presenting showing exhibiting displaying demonstrating illustrating exemplifying typifying representing symbolizing signifying meaning denoting connoting implying suggesting indicating pointing referring alluding hinting intimating insinuating inferring deducing concluding reasoning thinking cogitating cerebrating pondering musing reflecting contemplating meditating concentrating focusing listening hearing heeding noting noticing observing watching looking seeing viewing regarding considering weighing evaluating assessing judging appraising estimating guessing speculating hypothesizing theorizing postulating proposing recommending advising counseling consulting conferring discussing debating arguing disputing contesting challenging opposing resisting defying confronting facing encountering experiencing undergoing suffering enduring withstanding surviving living existing subsisting persisting persevering continuing remaining staying abiding residing inhabiting occupying possessing owning holding keeping retaining maintaining preserving conserving protecting guarding defending securing sheltering harboring entertaining welcoming greeting saluting hailing acclaiming applauding praising commending lauding extolling celebrating honoring glorifying exalting magnifying amplifying enlarging expanding extending stretching diffusing dispersing scattering allocating assigning allotting apportioning dividing partitioning separating detaching disconnecting disjoining disuniting dissociating disengaging withdrawing retreating receding retrograding reversing inverting turning rotating revolving spinning twirling whirling swirling circling orbiting looping coiling winding twisting bending curving arching vaulting bridging spanning crossing traversing passing moving going traveling journeying voyaging sailing flying gliding soaring hovering floating drifting flowing streaming pouring rushing gushing spurting spraying splashing splattering spattering uniting combining merging fusing blending mixing mingling intermingling intermixing integrating incorporating assimilating absorbing digesting metabolizing transforming converting transmuting transfiguring metamorphosing evolving developing growing maturing ripening aging declining deteriorating degenerating decaying rotting decomposing disintegrating crumbling collapsing falling failing dying perishing expiring departing leaving exiting retiring resigning abdicating relinquishing surrendering yielding submitting succumbing capitulating acquiescing complying conforming adapting adjusting accommodating reconciling harmonizing synchronizing coordinating unifying standardizing normalizing regularizing systematizing methodizing rationalizing streamlining simplifying clarifying elucidating explaining interpreting translating rendering paraphrasing summarizing abstracting condensing compressing shrinking reducing decreasing diminishing lessening lowering dropping descending sinking submerging diving plunging plummeting crashing colliding impacting hitting striking knocking tapping patting touching sensing perceiving detecting discovering finding locating positioning placing putting setting laying standing sitting lying resting relaxing reposing sleeping dreaming imagining fantasizing daydreaming woolgathering praying worshiping revering venerating adoring loving nourishing feeding eating drinking consuming ingesting energizing powering fueling motivating inspiring stimulating exciting arousing awakening alerting warning cautioning informing notifying telling reporting announcing declaring stating asserting claiming contending proving revealing disclosing exposing uncovering unmasking unveiling opening unlocking unsealing unfastening untying unbinding unwrapping unpacking unloading emptying clearing cleaning washing bathing showering rinsing soaking steeping immersing drowning suffocating choking strangling throttling squeezing pressing centering aiming targeting zeroing aligning tuning calibrating balancing leveling equalizing stabilizing steady mooring docking landing touching-down alighting perching roosting nesting denning burrowing tunneling digging excavating mining quarrying extracting removing eliminating eradicating exterminating destroying demolishing ruining wrecking damaging harming injuring hurting wounding maiming crippling laming disabling incapacitating paralyzing stunning shocking surprising astonishing amazing astounding staggering stupefying dazing bewildering confusing puzzling perplexing baffling mystifying enigmatic cryptic obscure abstruse recondite esoteric arcane occult mysterious unknown unfamiliar strange odd peculiar curious queer weird bizarre fantastic extraordinary remarkable notable noteworthy significant important momentous consequential weighty grave serious crucial vital essential necessary required needed wanted desired wished hoped expected anticipated awaited looked-for searched-for sought hunted pursued chased tracked trailed followed shadowed stalked crept crawled climbed scaled ascended mounted rose lifted elevated raised hoisted heaved hauled pulled tugged dragged drawn attracted lured enticed tempted seduced charmed enchanted fascinated captivated engrossed absorbed immersed involved engaged occupied busy active energetic dynamic vigorous forceful powerful strong mighty potent effective efficient productive fruitful fertile fecund prolific abundant plentiful ample sufficient adequate enough satisfactory acceptable tolerable bearable endurable sustainable maintainable manageable controllable governable rule govern reign dominate command direct lead guide steer pilot navigate drive propel push thrust shove press squeeze compress condense concentrate aim point target zero align adjust tune calibrate level equalize standardize normalize stabilize fix secure anchor moor dock land touch-down alight perch roost nest den burrow tunnel dig excavate mine quarry extract remove eliminate eradicate exterminate destroy demolish ruin wreck damage harm injure hurt wound maim cripple lame disable incapacitate paralyze stun shock astonish amaze astound stagger stupefy daze bewilder confuse puzzle perplex baffle mystify enigma
Phortica variegata
variegated fruit fly
Phortica variegata is a small vinegar fly in the family Drosophilidae, distinguished by its unusual zoophilic behavior. Unlike most drosophilids, adult males feed on the lacrimal secretions (tears) of mammals, including humans, dogs, foxes, and other carnivores. This lachryphagous behavior makes it the primary European vector of Thelazia callipaeda, a zoonotic eyeworm that causes ocular infections in mammals. The species is native to Europe and has been introduced to the United States, where it has demonstrated vector competence for the parasite. Climate change is projected to alter its distribution significantly, with suitable habitats shifting toward Mediterranean coastal regions while declining in central-southern-eastern European transitional areas.
vector zoonotic-disease lachryphagy Steganinae Thelazia-callipaeda climate-change-sensitive emerging-infectious-disease olfactory-ecology
Sarcoptidae
Itch Mites
Sarcoptidae is a family of parasitic mites in the order Sarcoptiformes, commonly known as itch mites. The family includes the genus Sarcoptes, whose members cause sarcoptic mange in mammals. These mites are obligate parasites that burrow into host skin, causing intense pruritus, crusting lesions, and potentially fatal secondary infections. Sarcoptes scabiei, the most studied species, infests over 200 million humans worldwide and numerous domestic and wild mammal species.
parasite mite scabies mange zoonosis veterinary wildlife-disease neglected-tropical-disease burrowing epidermal-parasite direct-life-cycle host-specific-strains immunology emerging-infectious-disease biodiversity-threat cross-species-transmission feral-dog-reservoir quantitative-surveillance tissue-digestion-protocol ivermectin-treatment crusted-scabies ordinary-scabies pruritus alopecia hyperkeratosis secondary-bacterial-infection WHO-neglected-tropical-disease over-200-million-human-cases mammalian-host-specificity Astigmata Sarcoptiformes Arachnida Acari
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 nitrogen-cycle-change phosphorus-cycle-change carbon-cycle-change water-cycle-change hydrological-change soil-change vegetation-change ecosystem-change landscape-change seascape-change biosphere-change noosphere-change technosphere-change anthroposphere-change sociosphere-change econosphere-change politosphere-change culturosphere-change semiosphere-change infosphere-change digisphere-change cybersphere-change urban-sphere-change agricultural-sphere-change industrial-sphere-change extractive-sphere-change manufacturing-sphere-change service-sphere-change financial-sphere-change knowledge-sphere-change innovation-sphere-change creativity-sphere-change culture-sphere-change identity-sphere-change meaning-sphere-change value-sphere-change purpose-sphere-change vision-sphere-change mission-sphere-change strategy-sphere-change tactic-sphere-change operation-sphere-change action-sphere-change practice-sphere-change habit-sphere-change routine-sphere-change ritual-sphere-change 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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