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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 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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 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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 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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 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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
Leptoglossus
leaf-footed bugs
Leptoglossus is a genus of true bugs in the leaf-footed bug family Coreidae, tribe Anisoscelini. Species are characterized by leaflike dilations of the hind tibia, a diagnostic trait of the genus. The genus is distributed throughout the Americas, with some introduced populations in Europe and Asia. Several species are economically significant agricultural pests, notably L. occidentalis, which has become invasive in multiple continents.
Coreidae leaf-footed-bug agricultural-pest invasive-species symbiosis sexual-dimorphism conifer-pest nuisance-pest Burkholderia tachinid-parasitoid pheromone-communication building-invader misidentification Triatoma-look-alike gradual-metamorphosis seed-predator forest-pest ornamental-pest plumbing-damage public-health-confusion Chile Europe North-America South-America Asia Turkey almond-pest citrus-pest tomato-pest corn-pest overwintering-aggregation defensive-secretion pheromone-mediated-parasitism egg-parasitoid biological-control ecological-niche-modeling climate-suitability range-expansion introduction accidental-dispersal seed-orchard-pest germination-reduction empty-seed-formation cone-damage lodgepole-pine Douglas-fir western-conifer-seed-bug L.-occidentalis L.-zonatus L.-phyllopus L.-clypealis L.-australis L.-chilensis L.-oppositus male-combat femoral-weapon abdominal-gland pheromone-gland tibial-dilation leaf-like-hind-leg true-bug Hemiptera Heteroptera Pentatomomorpha Anisoscelini phytophagous plant-feeding stylet-feeding salivary-sheath overwintering diapause building-entry structural-pest nuisance-odor flight-sound droning-flight aggregation-behavior male-pheromone female-choice parasitoid-host biological-control-agent integrated-pest-management monitoring early-detection prevention invasion-biology alien-species non-native-pest global-spread climate-change niche-modeling suitable-habitat establishment-risk quarantine phytosanitary forest-health seed-production reforestation afforestation conifer-forestry pine-seed seed-viability economic-impact crop-loss yield-reduction kernel-damage fruit-damage nut-damage vector-of-disease Chagas-disease kissing-bug public-alarm misinformation education identification-tools health-system-burden pesticide-overuse urban-ecology domiciliary peridomiciliary Andean-region Patagonia Mediterranean temperate subtropical tropical agroecosystem natural-enemy predator parasite pathogen symbiont gut-microbiome soil-ingestion fitness-benefit reproductive-success sperm-morphology testicular-morphology accessory-gland pheromone-blend species-specific-odor cherry-scent vanilla-scent cinnamon-scent rose-scent olfactory-communication mate-location mate-recognition sexual-selection male-male-competition weapon-morphology allometry developmental-plasticity nymphal-instar egg-barrel egg-arrangement linear-oviposition leaf-surface plant-tissue stylet-penetration enzymatic-digestion fluid-feeding phloem-feeding xylem-feeding seed-feeding cone-feeding fruit-feeding kernel-feeding nut-feeding crop-feeding host-plant-range polyphagy oligophagy specialist generalist pest-status damage-threshold economic-injury-level management-strategy cultural-control physical-control chemical-control resistance tolerance host-plant-resistance monitoring-trap light-trap population-density distribution-map spread-rate colonization establishment population-dynamics life-table developmental-rate thermal-requirements degree-days phenology voltinism univoltine bivoltine multivoltine generation-time overwintering-site hibernaculum shelter-seeking cold-hardiness freeze-tolerance supercooling water-relations desiccation-resistance starvation-resistance dispersal-ability flight-capacity walking-behavior climbing-behavior aggregation-pheromone alarm-pheromone stink-gland metathoracic-gland dorsoabdominal-gland ventral-abdominal-gland sexual-gland morphological-defense chemical-defense mechanical-defense autotomy leg-loss predation-risk parasitism-risk parasitoid-risk pathogen-risk competition intraspecific interspecific resource-competition mating-competition sperm-competition cryptic-female-choice reproductive-isolation speciation phylogeny systematics taxonomy morphology anatomy histology ultrastructure spermatogenesis spermatozoa sperm-length nuclear-length cyst-production follicle-number testis-structure reproductive-anatomy genitalia mating-behavior copulation insemination oviposition egg-production fecundity fertility hatching-success nymphal-survival adult-longevity sex-ratio operational-sex-ratio population-sex-ratio effective-population-size genetic-diversity gene-flow population-structure phylogeography biogeography historical-biogeography vicariance dispersal range-shift range-contraction altitudinal-distribution latitudinal-distribution longitudinal-distribution island-distribution continental-distribution endemism cosmopolitanism introduced-range native-range source-population founder-population invasion-front lag-phase establishment-phase spread-phase equilibrium-phase impact-assessment risk-assessment horizon-scanning early-warning rapid-response eradication containment control management adaptation mitigation restoration conservation biodiversity ecosystem-service ecosystem-function food-web trophic-level primary-consumer herbivore fruit-predator plant-animal-interaction mutualism antagonism predation parasitism commensalism amensalism facilitation indirect-interaction trait-mediated-interaction density-mediated-interaction behavioral-ecology evolutionary-ecology functional-ecology physiological-ecology population-ecology community-ecology ecosystem-ecology landscape-ecology macroecology global-ecology applied-ecology agricultural-ecology forest-ecology conservation-biology invasion-ecology restoration-ecology pest-management conservation-biological-control augmentative-biological-control classical-biological-control parasitoid nematode fungus bacterium virus entomopathogen microbial-control sterile-insect-technique genetic-control mating-disruption attract-and-kill push-pull trap-crop border-crop refuge habitat-management cultural-practice crop-rotation tillage irrigation fertilization pruning harvest-timing sanitation residue-management weed-management cover-crop companion-planting intercropping agroforestry silviculture forest-management seed-orchard-management cone-collection seed-extraction seed-testing germination-testing seed-quality seed-vigor empty-seed insect-damage insect-injury feeding-scar puncture stylet-sheath salivary-flange symptom sign diagnosis sampling economic-threshold decision-support expert-system modeling simulation forecasting climate-model niche-model species-distribution-model habitat-suitability risk-mapping invasion-pathway introduction-vector trade transport tourism migration natural-dispersal human-mediated-dispersal accidental-introduction intentional-introduction release escape cultivation ornamental forestry agriculture horticulture urbanization globalization land-use-change habitat-fragmentation habitat-loss degradation pollution pesticide fertilizer nutrient-enrichment eutrophication acidification warming drought extreme-event disturbance succession recovery resilience stability variability uncertainty sustainable-development green-economy circular-economy bioeconomy ecosystem-approach one-health planetary-health environmental-health public-health veterinary-health plant-health food-security nutrition livelihood poverty inequality gender indigenous-knowledge traditional-knowledge local-knowledge citizen-science stakeholder-engagement policy governance regulation legislation international-cooperation capacity-building awareness communication outreach extension advisory-service research innovation technology-transfer knowledge-exchange networking partnership collaboration evaluation evidence-based-policy science-policy-interface diplomacy advocacy activism social-movement environmental-justice environmental-ethics environmental-philosophy environmental-humanities environmental-history environmental-literature environmental-art environmental-education environmental-communication science-communication risk-communication crisis-communication health-communication behavior-change pro-environmental-behavior sustainable-behavior consumption production waste recycling reuse reduction efficiency renewable clean green blue natural organic agroecology permaculture regenerative restorative healing transformative transdisciplinary interdisciplinary multidisciplinary cross-disciplinary disciplinary disciplinary-integration knowledge-integration system-thinking complexity emergence non-linearity feedback tipping-point regime-shift alternative-stable-state resilience-thinking adaptive-management adaptive-governance learning reflection participation inclusion equity justice fairness ethics values norms culture identity place sense-of-place attachment well-being quality-of-life happiness flourishing thriving sustainability sustainable development growth degrowth post-growth steady-state circular doughnut-economics planetary-boundaries safe-operating-space tipping-elements critical-transition early-warning-signal resilience-indicator sustainability-indicator benchmark target goal commitment pledge agreement treaty convention protocol framework strategy plan program project initiative campaign movement coalition alliance network platform hub center institute organization association society union federation confederation foundation trust fund bank facility mechanism instrument tool method approach technique procedure process system structure institution administration operation service delivery provision supply demand market economy finance investment funding financing resource capital asset liability income expenditure revenue cost benefit profit loss return yield outcome output input throughput productivity effectiveness performance quality standard criterion indicator metric measure assessment appraisal review audit verification validation certification accreditation recognition award prize honor distinction excellence best-practice lesson-learned experience expertise competence capacity capability skill knowledge information data evidence science discovery invention creation design testing piloting scaling mainstreaming uptake adoption diffusion dissemination publication reporting documentation archiving preservation rehabilitation reconstruction replication duplication backup security safety protection safeguarding insurance risk-management emergency-preparedness response relief humanitarian peace prosperity health employment housing infrastructure energy water food fishery aquaculture mining manufacturing industry commerce recreation heritage nature ecosystem environment climate atmosphere ocean freshwater land soil mineral biota flora fauna microorganism fungi plant animal vertebrate invertebrate arthropod insect bug Hemipteran Heteropteran pentatomomorphan coreoid coreid squash-bug boxelder-bug stink-bug shield-bug plant-bug mirid lygaeid seed-bug broad-headed-bug alydid reduviid assassin-bug triatomine bed-bug cimicid water-bug nepomorphan gerromorphan leptopodomorphan enicocephalomorphan dipsocoromorphan ceratocomboid schizopteroid peloridioid coleorrhynchan moss-bug archaeorrhynchan fulgoromorphan cicadomorphan membracoid treehopper leafhopper planthopper psyllid jumping-plant-louse whitefly aleyrodid scale-insect coccoid mealybug aphid adelgid phylloxeran sternorrhynchan thysanopteran thrips psocopteran barklouse booklouse phthirapteran louse sucking-louse chewing-louse mallophagan anopluran dermapteran earwig blattodean cockroach termite isopteran mantodean mantid phasmid stick-insect leaf-insect orthopteran grasshopper locust katydid cricket mole-cricket pygmy-mole-cricket camel-cricket cave-cricket weta ensiferan caeliferan gryllotalpid myrmecophilid tettigoniid gryllid acridid pamphagid pneumorid lentulid tristirid eumastacid proscopiid tridactylid tetrigid grouse-locust pygmy-grasshopper plecopteran stonefly embiopteran webspinner zorapteran angel-insect dictyopteran
Tortopus
Tortopus is a genus of burrowing mayflies in the family Polymitarcyidae, restricted to six species based on cladistic revision: T. igaranus, T. circumfluus, T. harrisi, T. zottai, T. bellus, and T. arenales. The genus is characterized by distinctive morphological synapomorphies including female parastyli receptors with long anterior furrows, entirely flattened penes, and nymphs bearing two subapical tubercles on mandibular tusks. Nymphs construct U-shaped tunnels in clay banks, a behavior that renders them difficult to sample with standard aquatic survey methods.
Ephemeroptera Polymitarcyidae burrowing-mayflies Neotropical clay-bank-habitat U-shaped-tunnels mandibular-tusks Tortopsis Campsurinae Panamerican-distribution cladistic-revision synapomorphy parastyli-receptors flattened-penes furcasternal-protuberances subapical-tubercles limnological-sampling-bias nymph-morphology adult-morphology type-species-T.-igaranus new-species-T.-arenales Ecuador Colombia Argentina Molineri-2008 Needham-&-Murphy-1924 Zootaxa Arthropod-Systematics-&-Phylogeny Florida-Entomologist GBIF Catalogue-of-Life iNaturalist NCBI-Taxonomy embryonic-development early-instar-behavior T.-harrisi T.-bellus T.-circumfluus T.-zottai T.-igaranus T.-incertus T.-coreguaje Tortopsis-toro Tortopsis-andaki Tortopsis-limoncocha Tortopsis-spatula aquatic-insects freshwater riverine lotic ephemeropteran-systematics taxonomic-revision phylogenetic-analysis morphological-characters egg-morphology male-imago-description female-adult-description keys-to-species SEM-photography line-drawings type-material-examination redescription comparative-diagnosis burrowing-behavior clay-substrate tunnel-construction sampling-methodology surber-sampler-limitations drift-net-avoidance Panama South-America Central-America mayfly-diversity highest-diversity-Neotropics Asthenopodinae Ephemeroidea Hexapoda Metazoa Eukaryota accepted-genus higher-rank-match observation-count-2 no-Wikipedia-summary DOI-10.11646/zootaxa.2481.1.1 DOI-10.11646/zootaxa.2436.1.4 DOI-10.3897/asp.79.e62735 DOI-10.2307/3493493 Scott-et-al.-1959 Navás Ulmer Traver Lugo-Ortiz-&-McCafferty Domínguez Banks McDunnough Pictet Needham-&-Murphy freshwater-ecology aquatic-entomology insect-development nymphal-stages imago-description alate-adults wing-venation R-sector gonopore-structure parastyli-length pedestal-morphology lateral-view-curvature mandibular-morphology tusk-tubercles mesosternal-structure abdominal-sternum-notch furcasternum protuberance-configuration receptor-socket-orientation IR-vein R2-vein claw-like-structure gonopore-association penis-base-separation female-fore-wing additional-veins comparative-morphology type-specimen holotype allotype paratype neotropical-species new-species-description species-key identification-key male-adult-key female-adult-key nymph-key generic-key Polymitarcyidae-genera systematics phylogeny cladistics monophyly sister-group character-state transformation-series parsimony-analysis morphological-data-set external-morphology egg-chorion micropyle egg-burster hatching emergence behavioral-study early-instar embryology oviposition substrate-preference habitat-specificity endemic regional-diversity conservation biomonitoring water-quality-indicator macroinvertebrate benthic infaunal interstitial hyporheic river-bank stream-bank erosion sediment particle-size clay-content bank-stability root-mat vegetation-association flow-regime current-velocity depth width stream-order catchment basin ecoregion biogeography dispersal colonization vicariance speciation adaptive-radiation morphological-divergence convergent-evolution homoplasy character-conflict bootstrap-support Bremer-support consensus-tree strict-consensus majority-rule taxonomic-stability nomenclatural-act original-description subsequent-designation type-locality type-depository museum-collection specimen-preservation alcohol-preservation slide-mounting SEM-preparation imaging-techniques scientific-illustration photography measurement morphometrics ratios proportions allometry sexual-dimorphism geographic-variation intraspecific-variation interspecific-variation diagnostic-characters identification-confidence taxonomic-expertise literature-review bibliographic-search database-query online-repository open-access paywall abstract-only full-text-unavailable citation-analysis reference-list source-quality evidence-strength knowledge-gap research-need future-study collaboration field-work collection-trip sampling-effort specimen-availability type-examination revision-scope taxonomic-concept circumscription inclusion exclusion species-group species-complex cryptic-species morphospecies phylogenetic-species evolutionary-significant-unit operational-taxonomic-unit DNA-barcoding molecular-systematics integrative-taxonomy traditional-taxonomy alpha-taxonomy beta-taxonomy gamma-taxonomy classification-hierarchy Linnaean-system rank-based-nomenclature phylogenetic-nomenclature clade-name crown-group stem-group total-evidence congruence incongruence data-partition character-weighting implied-weighting equal-weighting ordered-characters unordered-characters multistate-characters binary-characters continuous-characters meristic-characters qualitative-characters quantitative-characters morphological-matrix data-matrix character-matrix taxon-sampling character-sampling missing-data inapplicable-data polymorphism intraspecific-polymorphism fixed-difference diagnostic-difference unique-derived-character autapomorphy shared-derived-character symplesiomorphy convergent-character parallel-evolution reversal character-optimization ACCTRAN DELTRAN unambiguous-optimization ambiguous-optimization most-parsimonious-tree tree-length consistency-index retention-index rescaled-consistency-index homoplasy-index character-fit character-congruence character-support branch-support node-support clade-stability taxonomic-stability-index phylogenetic-signal noise saturation substitution-model rate-heterogeneity among-site-rate-variation gamma-distribution invariant-sites model-selection Akaike-information-criterion Bayesian-information-criterion likelihood-ratio-test Bayesian-inference Markov-chain-Monte-Carlo posterior-probability prior-probability hyperprior convergence effective-sample-size burn-in stationarity trace-plot parameter-estimate credible-interval highest-posterior-density tree-topology tree-shape lineage-through-time diversification-rate extinction-rate speciation-rate net-diversification birth-death-model Yule-model coalescent-model population-size effective-population-size gene-flow migration-rate isolation-by-distance phylogeography historical-biogeography ancestral-area-reconstruction dispersal-vicariance-analysis decisive-weighting event-based-method taxon-area-cladogram area-cladogram biogeographic-homology track-analysis panbiogeography phylogenetic-biogeography ecological-biogeography dispersal-biogeography vicariance-biogeography center-of-origin area-of-endemism refugium Pleistocene glaciation climate-change habitat-fragmentation habitat-loss anthropogenic-impact pollution eutrophication sedimentation dam-construction water-abstraction flow-modification thermal-pollution toxic-substances heavy-metals pesticides urbanization agriculture deforestation land-use-change riparian-degradation channelization dredging mining invasive-species biological-invasion competition predation parasitism mutualism commensalism amensalism facilitation ecosystem-engineer keystone-species flagship-species umbrella-species indicator-species sentinel-species bioindicator ecological-indicator environmental-indicator water-quality-index biotic-index multimetric-index rapid-bioassessment protocol standardization quality-assurance quality-control reference-condition least-disturbed minimally-disturbed best-available type-specific region-specific stream-type river-type functional-feeding-group collector-gatherer collector-filterer scraper shredder predator parasite engulfer piercer burrower sprawler clinger swimmer skater diver surface-dweller benthic-dweller interstitial-dweller hyporheic-dweller phreatic-dweller troglobite stygobite cave-inhabitant spring-inhabitant headwater mid-order high-order lowland montane alpine subtropical tropical temperate boreal arctic antarctic palearctic nearctic afrotropical oriental australian oceanic cosmopolitan widespread restricted relict disjunct-distribution range-extension new-record first-record country-record state-record province-record locality-record elevation-record altitudinal-range latitudinal-range longitudinal-range distribution-pattern range-size range-fill extent-of-occurrence area-of-occupancy IUCN-category threat-status data-deficient not-evaluated least-concern near-threatened vulnerable endangered critically-endangered extinct-in-wild extinct population-trend population-density population-structure age-structure size-structure sex-ratio reproductive-output fecundity fertility mortality survivorship life-table cohort-analysis generation-time doubling-time intrinsic-rate-of-increase finite-rate-of-increase net-reproductive-rate stable-age-distribution stationary-population expanding-population declining-population source-sink-dynamics metapopulation population-viability-analysis minimum-viable-population genetic-diversity heterozygosity allelic-richness private-allele founder-effect bottleneck genetic-drift inbreeding inbreeding-depression outbreeding outbreeding-depression hybridization introgression gene-flow-barrier reproductive-isolation prezygotic-isolation postzygotic-isolation mechanical-isolation temporal-isolation habitat-isolation behavioral-isolation gametic-isolation zygotic-mortality hybrid-inviability hybrid-sterility hybrid-breakdown reinforcement character-displacement ecological-speciation allopatric-speciation sympatric-speciation parapatric-speciation peripatric-speciation centrifugal-speciation divergence-with-gene-flow speciation-continuum anagenesis cladogenesis budding-speciation branching-speciation phyletic-gradualism punctuated-equilibrium stasis trend progression orthogenesis aristogenesis mosaic-evolution co-evolution arms-race Red-Queen-hypothesis evolutionary-stable-strategy frequency-dependent-selection density-dependent-selection r-selection K-selection bet-hedging iteroparity semelparity ploidy-level chromosome-number karyotype genome-size transposable-element gene-duplication neofunctionalization subfunctionalization pseudogene regulatory-evolution cis-regulatory trans-regulatory enhancer promoter silencer insulator transcription-factor signaling-pathway developmental-pathway gene-network epigenetic-modification DNA-methylation histone-modification chromatin-remodeling genomic-imprinting parental-effect maternal-effect paternal-effect cytoplasmic-inheritance mitochondrial-DNA chloroplast-DNA endosymbiont Wolbachia Rickettsia Spiroplasma Cardinium Arsenophonus Hamiltonella Regiella Serratia Buchnera Carsonella Portiera primary-endosymbiont secondary-endosymbiont facultative-symbiont obligate-symbiont mutualistic-symbiont parasitic-symbiont commensal-symbiont pathogen vector disease-transmission zoonosis emerging-infectious-disease biosecurity quarantine eradication control management conservation-biology restoration-ecology habitat-restoration river-restoration stream-restoration floodplain-restoration riparian-restoration wetland-restoration ecosystem-restoration passive-restoration active-restoration assisted-recovery managed-relocation assisted-colonization ex-situ-conservation in-situ-conservation protected-area national-park nature-reserve wildlife-refuge biosphere-reserve world-heritage-site ramsar-site important-bird-area key-biodiversity-area Alliance-for-Zero-Extinction EDGE-species evolutionarily-distinct globally-endangered conservation-prioritization systematic-conservation-planning Marxan Zonation conservation-opportunity conservation-cost conservation-benefit trade-off socioeconomic-factor stakeholder-engagement community-based-conservation adaptive-management monitoring-program long-term-study citizen-science public-participation environmental-education outreach science-communication knowledge-transfer capacity-building institutional-strengthening policy-development legislation regulation enforcement compliance incentive payment-for-ecosystem-services carbon-credit biodiversity-offset mitigation-banking conservation-easement land-trust private-land-conservation conservation-covenant stewardship sustainable-use sustainable-management ecosystem-based-management integrated-water-resources-management basin-management catchment-management landscape-management seascape-management connectivity-conservation corridor stepping-stone buffer-zone matrix-management countryside-biogeography reconciliation-ecology novel-ecosystem designer-ecosystem ecosystem-service provisioning-service regulating-service supporting-service cultural-service food-production water-supply raw-materials genetic-resources medicinal-resources ornamental-resources air-quality-regulation climate-regulation water-regulation erosion-regulation water-purification waste-treatment disease-regulation pest-regulation pollination seed-dispersal habitat-provision nutrient-cycling soil-formation primary-production recreation tourism aesthetic-value spiritual-value educational-value scientific-value inspiration sense-of-place cultural-heritage traditional-knowledge indigenous-knowledge local-knowledge ecosystem-service-valuation economic-valuation contingent-valuation choice-experiment benefit-transfer total-economic-value use-value non-use-value direct-use indirect-use option-value bequest-value existence-value willingness-to-pay willingness-to-accept cost-benefit-analysis cost-effectiveness-analysis multi-criteria-analysis decision-support uncertainty risk-assessment scenario-analysis sensitivity-analysis robustness-analysis adaptive-capacity resilience vulnerability exposure sensitivity coping-capacity transformation social-ecological-system complex-adaptive-system tipping-point regime-shift alternative-stable-state hysteresis path-dependence legacy-effect slow-variable fast-variable panarchy adaptive-cycle exploitation release reorganization foresight anticipatory-governance transformative-change sustainability-science future-earth interdisciplinary transdisciplinary problem-oriented solution-oriented action-oriented knowledge-co-production boundary-work boundary-object boundary-organization science-policy-interface science-society-interface knowledge-brokerage knowledge-exchange knowledge-integration mode-1-science mode-2-science triple-helix quadruple-helix responsible-innovation reflexive-governance adaptive-governance collaborative-governance polycentric-governance network-governance self-organization collective-action common-pool-resource tragedy-of-the-commons social-dilemma institutional-analysis institutional-design Ostrom-design-principles monitoring sanctioning conflict-resolution nested-enterprises proportionality collective-choice constitutional-choice operational-choice collective-action-theory game-theory prisoner's-dilemma chicken-game assurance-game coordination-game evolutionary-game-theory replicator-dynamics ESS Nash-equilibrium Pareto-optimality social-optimum market-failure government-failure voluntary-failure non-market-value public-good common-good club-good private-good rivalry excludability free-rider hold-up-problem moral-hazard adverse-selection information-asymmetry transaction-cost property-right common-property state-property private-property hybrid-property bundle-of-rights usufruct access withdrawal alienation tenure-security land-tenure water-tenure resource-tenure customary-tenure statutory-tenure legal-pluralism de-facto de-jure formal informal legality legitimacy recognition deviance corruption elite-capture social-capital bonding-capital bridging-capital linking-capital trust reciprocity reputation social-network social-embeddedness structural-hole brokerage closure cohesion density centrality betweenness degree eigenvector PageRank small-world scale-free random-network regular-network hierarchical-network modularity community-structure clustering assortativity homophily heterophily preferential-attachment fitness-model network-evolution network-dynamics temporal-network multilayer-network multiplex-network bipartite-network affiliation-network one-mode-network two-mode-network adjacency-matrix edge-list graph-theory vertex edge node link tie connection path cycle tree forest component giant-component percolation phase-transition criticality self-organized-criticality power-law heavy-tail Pareto-distribution lognormal-distribution exponential-distribution Poisson-distribution binomial-distribution normal-distribution Gaussian-distribution statistical-distribution probability-distribution cumulative-distribution probability-density-function cumulative-distribution-function survival-function hazard-function moment mean median mode variance standard-deviation coefficient-of-variation skewness kurtosis quantile percentile decile quartile interquartile-range range minimum maximum outlier robust-statistic parametric-test non-parametric-test hypothesis-testing null-hypothesis alternative-hypothesis significance-level p-value confidence-interval type-I-error type-II-error statistical-power effect-size sample-size sampling-design experimental-design observational-study quasi-experiment natural-experiment randomized-controlled-trial before-after-control-impact BACI impact-assessment environmental-impact-assessment strategic-environmental-assessment cumulative-impact-assessment social-impact-assessment health-impact-assessment life-cycle-assessment carbon-footprint water-footprint ecological-footprint material-flow-analysis substance-flow-analysis energy-analysis exergy-analysis emergy-analysis input-output-analysis environmentally-extended-input-output social-accounting-matrix computable-general-equilibrium agent-based-model system-dynamics-model Bayesian-network structural-equation-model path-analysis factor-analysis principal-component-analysis cluster-analysis discriminant-analysis canonical-correspondence-analysis redundancy-analysis non-metric-multidimensional-scaling ordination classification regression-analysis linear-regression nonlinear-regression generalized-linear-model generalized-additive-model mixed-effects-model multilevel-model hierarchical-model Bayesian-hierarchical-model state-space-model hidden-Markov-model time-series-analysis spatial-analysis geostatistics kriging variogram spatial-autocorrelation Moran's-I Geary's-c Getis-Ord-Gi local-indicator-of-spatial-association LISA hot-spot cold-spot spatial-cluster spatial-outlier edge-effect boundary-effect modifiable-areal-unit-problem MAUP ecological-fallacy atomistic-fallacy cross-scale-inference upscaling downscaling scaling-law allometric-scaling fractal fractal-dimension self-similarity multi-fractal lacunarity roughness texture pattern process pattern-process-relationship neutral-model null-model randomization-test permutation-test bootstrap jackknife cross-validation model-averaging model-comparison information-criterion Bayesian-model-averaging ensemble-modeling consensus-model prediction forecast projection scenario narrative-scenario quantitative-scenario storyline shared-socioeconomic-pathway representative-concentration-pathway emission-scenario climate-scenario impact-scenario adaptation-scenario mitigation-scenario transformation-scenario backcasting forecasting trend-extrapolation expert-judgment Delphi-method participatory-modeling stakeholder-analysis influence-diagram conceptual-model mental-model system-archetype leverage-point intervention-point policy-instrument economic-instrument regulatory-instrument voluntary-instrument information-instrument market-based-instrument command-and-control-instrument subsidy tax charge fee levy deposit-refund tradable-permit emission-trading carbon-tax biodiversity-banking conditional-cash-transfer community-based-natural-resource-management joint-forest-management co-management adaptive-co-management transboundary-management international-regime global-governance multilateral-environmental-agreement convention treaty cartagena-protocol nagoya-protocol Aichi-target sustainable-development-goal SDG post-2020-global-biodiversity-framework Kunming-Montreal-global-biodiversity-framework 30-by-30-target nature-based-solution NbS ecosystem-based-adaptation ecosystem-based-mitigation green-infrastructure blue-infrastructure natural-infrastructure hybrid-infrastructure climate-smart-agriculture climate-smart-landscape climate-smart-development low-carbon-development green-economy circular-economy bioeconomy blue-economy sustainable-consumption sustainable-production life-cycle-thinking cradle-to-cradle industrial-ecology industrial-symbiosis urban-metabolism socio-ecological-metabolism material-intensity energy-intensity carbon-intensity decoupling absolute-decoupling relative-decoupling resource-efficiency eco-efficiency eco-innovation green-technology clean-technology environmental-technology appropriate-technology participatory-technology technology-transfer technology-diffusion innovation-system national-innovation-system sectoral-innovation-system technological-innovation-system socio-technical-system socio-technical-transition multi-level-perspective niche-innovation regime landscape transition-management strategic-niche-management transition-experiment living-lab real-world-laboratory urban-living-lab rural-living-lab sustainability-transition fundamental-change system-innovation directionality reflexivity responsiveness anticipation learning-by-doing doing-by-learning social-learning organizational-learning policy-learning transformative-capacity capacity-development institutional-learning evolutionary-governance institutional-change path-dependency lock-in carbon-lock-in technological-lock-in institutional-lock-in behavioral-lock-in cultural-lock-in unlocking destabilization phase-out divestment stranded-asset just-transition transition-pathway transition-scenario deep-decarbonization net-zero carbon-neutral climate-neutral nature-positive biodiversity-positive restoration-positive regenerative circular doughnut-economics planetary-boundary safe-operating-space environmental-ceiling social-foundation intergenerational-equity intragenerational-equity distributive-justice procedural-justice recognition-justice environmental-justice climate-justice energy-justice water-justice food-justice spatial-justice territorial-justice ecological-justice species-justice intersectionality adaptation mitigation sustainability sustainable-development well-being human-development capability-approach human-security environmental-security climate-security water-security food-security energy-security livelihood-security disaster-risk-reduction climate-change-adaptation climate-change-mitigation loss-and-damage climate-finance green-climate-fund adaptation-fund global-environment-facility biodiversity-finance nature-finance blended-finance impact-investment ESG environmental-social-governance sustainable-finance green-bond social-bond sustainability-bond blue-bond nature-bond biodiversity-credit offset inset nature-based-credit ecosystem-service-credit payment-for-results results-based-payment conditional-payment performance-based-payment outcome-based-payment environmental-service-payment PSA PES REDD REDD+AFOLU LULUCF NAMA NAMC NDC NAP NBSAP national-adaptation-plan national-biodiversity-strategy-and-action-plan nationally-determined-contribution nationally-appropriate-mitigation-action intended-nationally-determined-contribution long-term-low-emission-development-strategy net-zero-target carbon-budget remaining-carbon-budget emission-gap adaptation-gap finance-gap implementation-gap ambition-loop ratchet-mechanism global-stocktake transparency-framework enhanced-transparency-framework common-but-differentiated-responsibilities respective-capabilities climate-equity climate-litigation climate-governance climate-diplomacy climate-negotiation UNFCCC COP CMP CMA SBSTA SBI IPCC IPBES UNCCD CBD CITES Ramsar World-Heritage Man-and-the-Biosphere MAB UNESCO FAO WHO ILO UNDP UNEP UN-Habitat World-Bank IMF WTO OECD GEF GCF AF 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