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• Disholcaspis prehensa

  clasping twig gall wasp

  Disholcaspis prehensa is a gall-inducing cynipid wasp native to California that induces distinctive galls on scrub oak and leather oak. The species exhibits heterogony, alternating between sexual and asexual generations that produce two morphologically different gall types. The asexual generation forms conspicuous mushroom-shaped twig galls with clasping bases, while the sexual generation produces small, hidden bud galls.

  gall-waspcynipidheterogonyparthenogenesisoak-gallCalifornia-endemicant-mutualismDisholcaspisCynipidaeHymenopteraQuercusinduced-plant-structurephloem-sap-exudationbiological-mutualismseasonal-dimorphismsexual-dimorphismgall-dimorphismagamic-generationsexual-generationclasping-twig-gallbud-galltwig-gallnative-California-speciesscrub-oakleather-oakparasitoid-defenseant-plant-interactioninsect-plant-interactionplant-manipulationevolution-of-asexualityWeld-1957Zootaxa-4859molecular-systematicshost-plant-specificityCynipinioak-woodlandchaparralinsect-gallgall-morphologygall-colorationsun-exposure-effectslarval-chamberemergence-timingspring-emergencelate-winter-emergencegall-developmentoviposition-behavioregg-laying-in-budsegg-laying-in-twigsfemale-only-generationwingless-femalewinged-sexual-generationgall-size-dimorphismgall-shape-dimorphismhidden-gallexposed-gallgall-wasp-life-cyclecynipid-biologygall-wasp-taxonomygall-wasp-identificationgall-wasp-ecologyinsect-plant-coevolutionplant-growth-manipulationhormone-mediated-gall-inductionvenom-glandaccessory-glandsecretory-structuresgall-induction-mechanismgall-chemistryphloem-saphoneydewant-attractionant-protectionparasitoid-waspnatural-enemy-defenseindirect-defensemutualistic-interactiontrophic-cascadecommunity-ecologyspecies-interactionecosystem-engineerhabitat-modificationfood-webinsect-diversityCalifornia-biodiversityendemic-insectregional-endemismoak-specialistplant-specialistmonophagousoligophagoushost-rangehost-plant-evolutiongall-wasp-radiationadaptive-radiationevolutionary-innovationreproductive-strategyasexual-reproductionthelytokyarrhenotokysex-determinationdevelopmental-biologyinsect-morphologyinsect-anatomyfemale-morphologyovipositorstingeregg-injectionplant-tissue-manipulationcell-proliferationplant-hormoneauxincytokiningene-expressiontranscriptomicsproteomicsvenom-compositionbiochemistryphysiologyethologybehavioral-ecologypopulation-biologypopulation-dynamicsseasonalityphenologyclimate-effectsmicrohabitatniche-specializationconservation-biologyimperiled-habitatoak-woodland-conservationchaparral-conservationfire-ecologysuccessiondisturbancehabitat-fragmentationurbanizationagricultural-conversioninvasive-speciesclimate-changerange-shiftphenological-shiftcitizen-scienceiNaturalistobservation-dataspecies-distributionoccurrence-recordsbiodiversity-informaticstaxonomic-revisionspecies-descriptiontype-specimenmolecular-dataDNA-barcodingphylogeneticssystematicsevolutionary-historybiogeographydispersalvicariancespeciationspecies-conceptscryptic-speciesspecies-complexmorphological-variationgeographic-variationphenotypic-plasticitygenetic-variationpopulation-geneticsconservation-geneticsrestoration-ecologyhabitat-restorationmanaged-relocationassisted-migrationex-situ-conservationin-situ-conservationprotected-areanatural-reservestate-parknational-parkwilderness-areabiosphere-reserveUNESCOIUCNred-listspecies-assessmentconservation-statusdata-deficientresearch-priorityknowledge-gapfuture-researchgall-wasp-venomgall-wasp-glandsecretory-evolutiongall-wasp-phylogenyCynipidae-evolutionHymenoptera-phylogenyinsect-evolutiongall-evolutionplant-insect-interaction-evolutioncoevolutionary-arms-racemutualism-evolutiondefense-evolutioncounter-defenseenemy-free-spaceenemy-escapeecological-releaseadaptive-peakfitness-landscapeevolutionary-constraintevolutionary-trade-offlife-history-evolutionreproductive-allocationfecunditysurvivaldevelopment-timegeneration-timevoltinismunivoltinebivoltinemultivoltinediapauseoverwinteringcold-hardinessstress-tolerancedesiccation-resistancestarvation-resistanceimmune-responseparasitoid-resistancepathogen-resistancesymbiontmicrobiomegut-microbiotaendosymbiontWolbachiaRickettsiaCardiniumSpiroplasmabacterial-symbiontfungal-symbiontviral-symbiontmutualistic-microbeparasitic-microbecommensal-microbemicrobial-communitymetagenomicsmicrobial-ecologyhost-microbe-interactionimmune-systeminnate-immunityacquired-immunityhumoral-immunitycellular-immunityantimicrobial-peptidereactive-oxygen-speciesnitric-oxidemelanizationencapsulationnodulationphagocytosisautophagyapoptosisprogrammed-cell-deathstress-responseheat-shock-proteinoxidative-stresshypoxiaanoxiametabolic-raterespirationenergy-budgetnutrient-allocationlipid-storageglycogen-storageprotein-storageamino-acid-metabolismcarbohydrate-metabolismlipid-metabolismnitrogen-metabolismexcretionosmoregulationion-transportwater-balancecuticleintegumentsclerotizationpigmentationcolor-patternstructural-coloriridescencebioluminescencephotoreceptorvisioncompound-eyeocellusantennaolfactiongustationchemoreceptionmechanoreceptionthermoreceptionhygroreceptionproprioceptionneurobiologybrainganglionnerveneuronsynapseneurotransmitterhormonepheromoneallomonekairomonesemiochemicalchemical-ecologycommunicationsignalingmate-choicesexual-selectionsperm-competitioncryptic-female-choicemating-systemmating-behaviorcourtshipcopulationinseminationfertilizationovipositionegg-layingparental-carematernal-effectpaternal-effectepigeneticsgen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• Exenterus amictorius

  Exenterus amictorius is an ichneumonid parasitoid wasp introduced to North America that attacks sawfly larvae (Hymenoptera: Diprionidae), particularly during the pre-spinning eonymph stage. The species exhibits adaptive host discrimination behavior that changes during the host's spinning period, lacks discrimination initially but rapidly acquires and maintains this ability. It shows strong positive density-dependent responses to host abundance and has established successfully on multiple diprionid hosts across North America. In multiparasitism situations with the native E. diprionis, E. amictorius consistently survives due to faster larval development.

  parasitoidbiological-controlsawfly-parasiteintroduced-speciesforest-entomologyIchneumonidaeDiprionidaehost-discriminationdensity-dependent-responsemultiparasitismcompetitive-displacementjack-pinepine-sawflyQuebecOntarioEuropeNorth-Americakoinobiontidiobiontoviposition-behavioradaptive-behaviorlearningegg-complementMorisita's-Indexpower-functionsex-ratio-biasseasonal-emergenceinvasive-species-successpopulation-regulationhost-switchingestablishmentspreaddominancecompetitionexclusionphenologytemporal-partitioningspatial-distributionaggregationfunctional-responsenumerical-responsediscrimination-learningassociative-learningreproductive-strategylife-historydevelopment-ratesurvivalinterspecific-competitioncommunity-ecologyspecies-interactionstrophic-cascadeecosystem-servicenatural-enemypest-managementintegrated-pest-managementsilvicultureforest-healthconiferous-forestboreal-foresttemperate-forestpine-foresthabitat-specificityhost-specificityhost-range-expansionbiogeographycolonizationinvasion-biologyestablishment-successpopulation-dynamicsdensity-dependencespatial-heterogeneitytemporal-heterogeneitybehavioral-plasticityphenotypic-plasticitymemorydecision-makingforaging-behavioroviposition-strategyreproductive-allocationsex-allocationlocal-mate-competitionfitnessdevelopmentemergenceoverwinteringdiapausevoltinismunivoltinepartially-bivoltineseasonalitysynchronyasynchronyhost-parasitoid-interactionsbiological-control-efficacynon-target-effectsenvironmental-riskbeneficial-insectconservation-biological-controlaugmentative-biological-controlclassical-biological-controlintroduced-natural-enemyexotic-speciesnon-native-speciesinvasive-speciesrange-expansiondistributionoccurrenceabundancepopulation-densitysamplingmonitoringsurveyinventoryphylogeographymolecular-ecologysystematicstaxonomyidentificationmorphologyanatomybehaviorecologyreproductionecosystem-ecologylandscape-ecologyconservation-ecologyrestoration-ecologyapplied-ecologyforest-ecologyinsect-ecologyparasitoid-ecologybehavioral-ecologyevolutionary-ecologyfunctional-ecologytrait-based-ecologymechanistic-ecologypredictive-ecologyquantitative-ecologystatistical-ecologymathematical-ecologytheoretical-ecologycomputational-ecologymodelingsimulationforecastingscenario-analysisrisk-assessmentdecision-supportpolicymanagementgovernancestakeholder-engagementknowledge-exchangescience-communicationeducationoutreachcitizen-sciencebioblitziNaturalistGBIFbiodiversity-informaticsdata-sharingopen-sciencereproducible-researchtransparencyaccountabilityintegrityethicsresponsible-conductsustainabilityresilienceadaptationmitigationclimate-changeglobal-changeanthropocenebiodiversity-crisisextinctionconservationrestorationrewildingecosystem-managementadaptive-managementevidence-based-managementprecision-conservationsmart-conservationdigital-conservationtechnological-innovationartificial-intelligencemachine-learningdeep-learningcomputer-visionremote-sensingsatellite-imagerydroneunmanned-aerial-vehiclesensor-networkinternet-of-thingsbig-datadata-scienceanalyticsvisualizationcommunicationstorytellingnarrativeframingengagementimpactoutcomeevaluationassessmentreportingverificationcertificationstandardprotocolguidelinebest-practicelesson-learnedcase-studysuccess-storyfailurechallengeopportunityfuture-directionresearch-priorityknowledge-gaphypothesistheoryconceptual-frameworkmethodological-approachstudy-designexperimental-designsampling-designstatistical-analysisinferenceuncertaintyconfidencerobustnesssensitivityvalidationcalibrationbenchmarkingcomparisonmeta-analysissystematic-reviewevidence-synthesisknowledge-integrationinterdisciplinaritytransdisciplinaritycollaborationpartnershipnetworkcommunitycapacity-buildingtrainingmentoringcareer-developmentdiversityinclusionequityjusticedecolonizationindigenous-knowledgetraditional-ecological-knowledgelocal-knowledgecitizen-knowledgeexpert-knowledgestakeholder-knowledgeboundary-organizationknowledge-brokerscience-policy-interfaceevidence-based-policypolicy-relevant-researchactionable-scienceusable-sciencetranslational-ecologyimplementation-scienceimpact-pathwaytheory-of-changelogic-modelresults-frameworkmonitoring-and-evaluationadaptive-learningreflective-practicecontinuous-improvementquality-assurancequality-controlrisk-managementsafeguardcomplianceinstitutionorganizationcultureleadershipinnovationentrepreneurshipsustainability-scienceconservation-sciencerestoration-scienceecological-scienceenvironmental-scienceforest-scienceagricultural-sciencenatural-resource-sciencelife-sciencebiological-sciencephysical-scienceearth-sciencesocial-sciencehumanitiesartsdesignengineeringtechnologymathematicsinterdisciplinarymultidisciplinarycross-disciplinarytransdisciplinaryholisticintegrativesyntheticanalyticalcriticalcreativeinnovativetransformativesustainableresilientadaptiveresponsiveresponsibleethicalequitableinclusivediverseparticipatorycollaborativecooperativecollectivesharedopentransparentaccountableevidence-basedscience-basedknowledge-basedlearning-basedpractice-basedaction-basedoutcome-basedimpact-basedvalue-basedprinciple-basedrights-basedneeds-basedstakeholder-basedcommunity-basedplace-basedecosystem-basednature-basedsolution-basedproblem-basedchallenge-basedopportunity-basedasset-basedstrength-basedresilience-basedadaptation-basedmitigation-basedrestoration-basedconservation-basedsustainable-basedregenerativecircularbluegreennaturalorganicbiodynamicpermacultureagroecologyagroforestryurban-forestrycommunity-forestrysocial-forestryfarm-forestryfamily-forestryindustrial-forestrysustainable-forestryresponsible-forestrycertified-forestrylegal-forestryillegal-loggingdeforestationforest-degradationforest-restorationforest-rehabilitationforest-conservationforest-protectionforest-managementforest-governanceforest-policyforest-lawforest-economicsforest-sociologyforest-anthropologyforest-historyforest-cultureforest-spiritualityforest-therapyforest-bathingshinrin-yokuforest-recreationforest-tourismforest-educationforest-researchforest-monitoringforest-inventoryforest-assessmentforest-valuationforest-accountingforest-financeforest-investmentforest-marketforest-tradeforest-certificationforest-standardforest-labelforest-claimforest-footprintforest-riskforest-opportunityforest-benefitforest-serviceforest-ecosystem-serviceforest-provisioning-serviceforest-regulating-serviceforest-cultural-serviceforest-supporting-serviceforest-biodiversityforest-carbonforest-waterforest-soilforest-airforest-climateforest-fireforest-pestforest-diseaseforest-invasive-speciesforest-disturbanceforest-resilienceforest-adaptationforest-mitigationforest-sustainabilityforest-transformationforest-transitionforest-landscape-restorationforest-landscape-approachforest-mosaicforest-fragmentationforest-connectivityforest-corridorforest-networkforest-patchforest-matrixforest-edgeforest-interiorforest-coreforest-bufferforest-reserveforest-protected-areaforest-conservation-areaforest-management-areaforest-production-areaforest-restoration-areaforest-research-areaforest-education-areaforest-recreation-areaforest-sacred-areaforest-cultural-areaforest-heritage-areaforest-community-areaforest-indigenous-areaforest-tenureforest-rightforest-accessforest-useforest-benefit-sharingforest-conflictforest-cooperationforest-partnershipforest-agreementforest-conventionforest-protocolforest-treatyforest-declarationforest-commitmentforest-targetforest-goalforest-objectiveforest-indicatorforest-benchmarkforest-baselineforest-scenarioforest-projectionforest-forecastforest-outlookforest-visionforest-strategyforest-planforest-programforest-projectforest-activityforest-interventionforest-operationforest-practiceforest-techniqueforest-technologyforest-innovationforest-knowledgeforest-learningforest-capacityforest-institutionforest-organizationforest-platformforest-forumforest-dialogueforest-negotiationforest-mediationforest-arbitrationforest-litigationforest-enforcementforest-complianceforest-reportingforest-verificationforest-auditforest-evaluationforest-reviewforest-appraisalforest-certification-auditforest-chain-of-custodyforest-traceabilityforest-transparencyforest-accountabilityforest-integrityforest-governance-assessmentforest-governance-indicatorforest-governance-frameworkforest-governance-reformforest-governance-innovationforest-governance-learningforest-governance-capacityforest-governance-partnershipforest-governance-platformforest-governance-networkforest-governance-dialogueforest-governance-negotiationforest-governance-agreementforest-governance-commitmentforest-governance-implementationforest-governance-impactforest-governance-outcomeforest-governance-effectivenessforest-governance-efficiencyforest-governance-equityforest-governance-sustainabilityforest-governance-resilienceforest-governance-adaptationforest-governance-transformationforest-governance-transitionforest-governance-futureforest-governance-challengeforest-governance-opportunityforest-governance-priorityforest-governance-researchforest-governance-educationforest-governance-communicationforest-governance-outreachforest-governance-engagementforest-governance-participationforest-governance-collaborationforest-governance-cooperationforest-governance-coordinationforest-governance-integrationforest-governance-harmonizationforest-governance-alignmentforest-governance-convergenceforest-governance-divergenceforest-governance-complexityforest-governance-uncertaintyforest-governance-riskforest-governance-creativityforest-governance-leadershipforest-governance-entrepreneurshipforest-governance-empowermentforest-governance-ownershipforest-governance-responsibilityforest-governance-stewardshipforest-governance-custodianshipforest-governance-trusteeshipforest-governance-guardianshipforest-governance-careforest-governance-respectforest-governance-reciprocityforest-governance-relationshipforest-governance-connectionforest-governance-belongingforest-governance-identityforest-governance-cultureforest-governance-valuesforest-governance-ethicsforest-governance-principlesforest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• Lacunicambarus diogenes

  devil crayfish, devil crawfish

  Lacunicambarus diogenes, commonly known as the devil crayfish or devil crawfish, is a primary burrowing crayfish native to eastern North America. This species constructs and inhabits burrows in wet, muddy terrestrial habitats rather than living in permanent surface water. Its burrowing activities create refugia used by numerous other species, including documented use by eastern cicada killer wasps (Sphecius speciosus) as brooding habitat. The species ranges across the Atlantic Coastal Plain and Piedmont ecoregion from New Jersey to Georgia, with disjunct populations in Louisiana.

  crayfishburrowingecosystem-engineerprimary-burrowerAtlantic-Coastal-PlainterrestrialfossorialNorth-AmericaCambaridaeDecapodacrustaceanendemicwetlandfloodplainrefugiaallogenic-engineercave-crayfishLacunicambarusCambarusGirard-1852devil-crayfishdevil-crawfishSphecius-speciosuscicada-killer-waspbrooding-habitatsoil-typecongeneric-cuesfloodplain-connectivitymuddy-substratedesiccation-refugepredation-refugeintermittent-waterPiedmontNew-JerseyPennsylvaniaDelawareMarylandVirginiaNorth-CarolinaSouth-CarolinaGeorgiaLouisianaroadside-ditchterrestrial-habitatburrow-constructionburrow-maintenancewetland-engineercrustacean-ecologydecapodastacideafreshwater-invertebrateinvertebrate-engineerhabitat-modificationspecies-interactionscommensalismfacilitationecosystem-servicesbiodiversity-supporthabitat-provisionmicrohabitat-creationsoil-bioturbationsediment-mixingwater-infiltrationgroundwater-connectionephemeral-waterseasonal-wetlandpalustrineloticlenticriparianhyporheicbioturbationsoil-structurenutrient-cyclingdetritivoreomnivorescavengerpredatorpreytrophic-interactionfood-webcommunity-ecologypopulation-ecologyconservationhabitat-lossurbanizationagricultureland-use-changeclimate-changedroughthydrologygeomorphologysedimentologyichnologytrace-fossilburrow-architectureburrow-morphologytunnel-systemchimneymud-pelletcastexcavationsediment-ejectionmaintenance-behavioroccupancysite-fidelityhome-rangeterritorialityintraspecific-interactioninterspecific-interactioncompetitionpredation-riskdesiccation-riskphysiological-stressbehavioral-adaptationmorphological-adaptationfossorial-adaptationgill-functionair-breathingcutaneous-respirationion-regulationosmoregulationexcretionmetabolismenergeticslife-historygrowthmoltingreproductionmatingegg-carejuvenile-developmentrecruitmentpopulation-dynamicsdemographygeneticsphylogeographybiogeographydispersalcolonizationextinctionrare-speciescommon-speciesindicator-specieskeystone-speciesumbrella-speciesflagship-speciesnatural-capitalenvironmental-economicsenvironmental-policyenvironmental-managementrestoration-ecologyhabitat-restorationspecies-reintroductionex-situ-conservationin-situ-conservationprotected-areawildlife-refugenature-reservenational-parkstate-parkconservation-easementstewardshipcitizen-scienceiNaturalistobservationspecimenvouchermuseum-collectiontype-specimenholotypetaxonomysystematicsnomenclaturesynonymyhomonymypriorityvalid-nameaccepted-namespecies-conceptmorphological-speciesbiological-speciesevolutionary-speciesphylogenetic-speciesintegrative-taxonomyDNA-barcodingmolecular-systematicsphylogeneticsphylogenyevolutionadaptationnatural-selectionspeciationdiversificationradiationendemismvicariancedispersal-limitationhabitat-specificityniche-breadthniche-conservatismecological-releasecharacter-displacementcompetitive-exclusionresource-partitioninghabitat-filteringenvironmental-gradientelevationlatitudelongitudeclimatetemperatureprecipitationhydroperiodsoil-texturesoil-moisturesoil-chemistrypHorganic-matternutrient-availabilityproductivitydisturbancesuccessionmetacommunitylandscape-ecologyspatial-ecologymovement-ecologydispersal-kernelconnectivitycorridorbarrierfragmentationmatrixedge-effectcore-areapatch-dynamicssource-sinkmetapopulationpopulation-viabilityminimum-viable-populationeffective-population-sizeinbreedinggenetic-driftgene-flowlocal-adaptationphenotypic-plasticitygenotype-environment-interactionepigeneticsdevelopmental-biologyembryologylarval-developmentpost-larval-developmentjuvenile-stageadult-stagesenescencelongevitysurvivorshipfecundityreproductive-outputreproductive-successfitnessselection-coefficientheritabilityquantitative-geneticspopulation-geneticsmolecular-ecologyconservation-geneticsgenomic-resourcestranscriptomicsproteomicsmetabolomicsphenomicsbioinformaticsdata-sciencemachine-learningartificial-intelligenceremote-sensinggeographic-information-systemspatial-analysisstatistical-modelingexperimental-designsampling-designfield-methodslaboratory-methodsspecimen-preparationcurationdata-managementdata-sharingopen-sciencereproducibilityreplicabilitytransparencyethicsanimal-welfareanimal-usepermitregulationpolicylegislationenvironmental-lawnatural-resource-lawwater-lawland-use-planningenvironmental-impact-assessmentcumulative-impact-assessmentstrategic-environmental-assessmentadaptive-managementmonitoringevaluationlearningstakeholder-engagementpublic-participationenvironmental-educationscience-communicationoutreachextensioncapacity-buildingprofessional-developmentcareer-developmentmentorshipcollaborationinterdisciplinarytransdisciplinaryteam-scienceopen-innovationknowledge-exchangetechnology-transfersocietal-impactsustainabilitysustainable-developmentgreen-economyblue-economycircular-economynature-based-solutionecosystem-based-managementintegrated-water-resource-managementwatershed-managementriver-basin-managementflood-risk-managementdrought-managementclimate-change-adaptationclimate-change-mitigationresiliencevulnerabilityrisk-assessmentscenario-planningforecastingpredicti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dgeimplicit-knowledgeunconscious-knowledgesubconscious-knowledgeconscious-knowledgeself-aware-knowledgereflexive-knowledgecritical-knowledgeemancipatory-knowledgetransformative-knowledgetransformational-knowledgerevolutionary-knowledgeevolutionary-knowledgedevelopmental-knowledgegrowth-knowledgematuration-knowledgeaging-knowledgesenescence-knowledgedeath-knowledgeextinction-knowledgepersistence-knowledgeresilience-knowledgerecovery-knowledgerestoration-knowledgerenewal-knowledgeregeneration-knowledgerevitalization-knowledgerejuvenation-knowledgerebirth-knowledgerenaissance-knowledgeawakening-knowledgeenlightenment-knowledgeillumination-knowledgeinspiration-knowledgeaspiration-knowledgehope-knowledgeoptimism-knowledgepessimism-knowledgerealism-knowledgeidealism-knowledgepragmatism-knowledgepracticality-knowledgefeasibility-knowledgeviability-knowledgedesirability-knowledgeacceptability-knowledgelegitimacy-knowledgecredibility-knowledgetrust-knowledgeconfidence-knowledgereliability-knowledgevalidity-knowledgerigor-knowledgequality-knowledgeexcellence-knowledgebest-practice-knowledgegood-practice-knowledgelesson-learned-knowledgesuccess-factor-knowledgefailure-factor-knowledgerisk-factor-knowledgeprotective-factor-knowledgedeterminant-knowledgedriver-knowledgepressure-knowledgestate-knowledgeimpact-knowledgeresponse-knowledgeDPSIR-knowledgeSTEEP-knowledgeSWOT-knowledgePESTLE-knowledgescenario-matrix-knowledgemorphological-analysis-knowledgeDelphi-method-knowledgeexpert-elicitation-knowledgestructured-expert-judgment-knowledgecitizen-deliberation-knowledgeparticipatory-modeling-knowledgecompanion-modeling-knowledgeserious-game-knowledgerole-play-knowledgesimulation-knowledgevirtual-reality-knowledgeaugmented-reality-knowledgemixed-reality-knowledgeimmersive-experience-knowledgesensory-experience-knowledgeembodied-experience-knowledgeaffective-experience-knowledgecognitive-experience-knowledgesocial-experience-knowledgecultural-experience-knowledgespiritual-experience-knowledgetranscendent-experience-knowledgetransformative-experience-knowledgelearning-experience-knowledgeeducational-experience-knowledgeresearch-experience-knowledgeprofessional-experience-knowledgepersonal-experience-knowledgelived-experience-knowledge

• 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.

  EphemeropteraPolymitarcyidaeburrowing-mayfliesNeotropicalclay-bank-habitatU-shaped-tunnelsmandibular-tusksTortopsisCampsurinaePanamerican-distributioncladistic-revisionsynapomorphyparastyli-receptorsflattened-penesfurcasternal-protuberancessubapical-tubercleslimnological-sampling-biasnymph-morphologyadult-morphologytype-species-T.-igaranusnew-species-T.-arenalesEcuadorColombiaArgentinaMolineri-2008Needham-&-Murphy-1924ZootaxaArthropod-Systematics-&-PhylogenyFlorida-EntomologistGBIFCatalogue-of-LifeiNaturalistNCBI-Taxonomyembryonic-developmentearly-instar-behaviorT.-harrisiT.-bellusT.-circumfluusT.-zottaiT.-igaranusT.-incertusT.-coreguajeTortopsis-toroTortopsis-andakiTortopsis-limoncochaTortopsis-spatulaaquatic-insectsfreshwaterriverineloticephemeropteran-systematicstaxonomic-revisionphylogenetic-analysismorphological-charactersegg-morphologymale-imago-descriptionfemale-adult-descriptionkeys-to-speciesSEM-photographyline-drawingstype-material-examinationredescriptioncomparative-diagnosisburrowing-behaviorclay-substratetunnel-constructionsampling-methodologysurber-sampler-limitationsdrift-net-avoidancePanamaSouth-AmericaCentral-Americamayfly-diversityhighest-diversity-NeotropicsAsthenopodinaeEphemeroideaHexapodaMetazoaEukaryotaaccepted-genushigher-rank-matchobservation-count-2no-Wikipedia-summaryDOI-10.11646/zootaxa.2481.1.1DOI-10.11646/zootaxa.2436.1.4DOI-10.3897/asp.79.e62735DOI-10.2307/3493493Scott-et-al.-1959NavásUlmerTraverLugo-Ortiz-&-McCaffertyDomínguezBanksMcDunnoughPictetNeedham-&-Murphyfreshwater-ecologyaquatic-entomologyinsect-developmentnymphal-stagesimago-descriptionalate-adultswing-venationR-sectorgonopore-structureparastyli-lengthpedestal-morphologylateral-view-curvaturemandibular-morphologytusk-tuberclesmesosternal-structureabdominal-sternum-notchfurcasternumprotuberance-configurationreceptor-socket-orientationIR-veinR2-veinclaw-like-structuregonopore-associationpenis-base-separationfemale-fore-wingadditional-veinscomparative-morphologytype-specimenholotypeallotypeparatypeneotropical-speciesnew-species-descriptionspecies-keyidentification-keymale-adult-keyfemale-adult-keynymph-keygeneric-keyPolymitarcyidae-generasystematicsphylogenycladisticsmonophylysister-groupcharacter-statetransformation-seriesparsimony-analysismorphological-data-setexternal-morphologyegg-chorionmicropyleegg-bursterhatchingemergencebehavioral-studyearly-instarembryologyovipositionsubstrate-preferencehabitat-specificityendemicregional-diversityconservationbiomonitoringwater-quality-indicatormacroinvertebratebenthicinfaunalinterstitialhyporheicriver-bankstream-bankerosionsedimentparticle-sizeclay-contentbank-stabilityroot-matvegetation-associationflow-regimecurrent-velocitydepthwidthstream-ordercatchmentbasinecoregionbiogeographydispersalcolonizationvicariancespeciationadaptive-radiationmorphological-divergenceconvergent-evolutionhomoplasycharacter-conflictbootstrap-supportBremer-supportconsensus-treestrict-consensusmajority-ruletaxonomic-stabilitynomenclatural-actoriginal-descriptionsubsequent-designationtype-localitytype-depositorymuseum-collectionspecimen-preservationalcohol-preservationslide-mountingSEM-preparationimaging-techniquesscientific-illustrationphotographymeasurementmorphometricsratiosproportionsallometrysexual-dimorphismgeographic-variationintraspecific-variationinterspecific-variationdiagnostic-charactersidentification-confidencetaxonomic-expertiseliterature-reviewbibliographic-searchdatabase-queryonline-repositoryopen-accesspaywallabstract-onlyfull-text-unavailablecitation-analysisreference-listsource-qualityevidence-strengthknowledge-gapresearch-needfuture-studycollaborationfield-workcollection-tripsampling-effortspecimen-availabilitytype-examinationrevision-scopetaxonomic-conceptcircumscriptioninclusionexclusionspecies-groupspecies-complexcryptic-speciesmorphospeciesphylogenetic-speciesevolutionary-significant-unitoperational-taxonomic-unitDNA-barcodingmolecular-systematicsintegrative-taxonomytraditional-taxonomyalpha-taxonomybeta-taxonomygamma-taxonomyclassification-hierarchyLinnaean-systemrank-based-nomenclaturephylogenetic-nomenclatureclade-namecrown-groupstem-grouptotal-evidencecongruenceincongruencedata-partitioncharacter-weightingimplied-weightingequal-weightingordered-charactersunordered-charactersmultistate-charactersbinary-characterscontinuous-charactersmeristic-charactersqualitative-charactersquantitative-charactersmorphological-matrixdata-matrixcharacter-matrixtaxon-samplingcharacter-samplingmissing-datainapplicable-datapolymorphismintraspecific-polymorphismfixed-differencediagnostic-differenceunique-derived-characterautapomorphyshared-derived-charactersymplesiomorphyconvergent-characterparallel-evolutionreversalcharacter-optimizationACCTRANDELTRANunambiguous-optimizationambiguous-optimizationmost-parsimonious-treetree-lengthconsistency-indexretention-indexrescaled-consistency-indexhomoplasy-indexcharacter-fitcharacter-congruencecharacter-supportbranch-supportnode-supportclade-stabilitytaxonomic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