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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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• Mannophorus forreri

  Mannophorus forreri is a longhorn beetle in the family Cerambycidae, described by Henry Walter Bates in 1885. It belongs to the tribe Trachyderini, a group known for often brightly colored and patterned species. The species is rarely encountered in collections and appears to have a restricted distribution in the southwestern United States and Mexico. Field observations indicate adults are active in early autumn and visit flowers of yellow composites in mountainous areas of Arizona.

  CerambycidaeTrachyderinilonghorn-beetleArizonaMexicoflower-visitorrare-speciesHenry-Walter-Bates1885orange-and-black-colorationGutierreziaHeterothecaThelespermaKitt-Peakautumn-activediurnalpollinatorMadrean-sky-islandsmontanexerophilicColeopterabeetleinsectarthropodCerambycinaeMannophorusforreriBates-1885GBIFiNaturalistCatalogue-of-LifeWikipediafield-observationcomposite-flowersAsteraceaesouthwestern-United-StatesNorth-AmericaMiddle-Americararely-collecteduncommonly-encounteredattractive-colorationcoleopterist-interestflower-feedingnectar-feedingpollen-feedinglarval-host-unknownwood-boringdiurnal-activitySeptemberearly-autumnlower-mountain-slopesrocky-habitatdry-habitatmountainoussubmontanesky-islandMadreanpollinationtrachyderinelong-antennaesexual-dimorphismantennae-lengthorangeblackelytral-apexpronotal-spotscylindrical-bodyrobustmedium-sizedquick-to-escapewaryspiral-flightfield-collectioniReportTed-MacRaeJeff-Huether2019Arizona-collectingKitt-Peak-RoadGutierrezia-microcephalaHeterotheca-fulcratayellow-compositesflower-patchesroadsidesslopesobservationspecimenvouchermuseum-collectionscientific-nameauthorshiptaxonspeciesgenusfamilyorderclassphylumkingdomanimallonghorncerambycidtaxonomyclassificationaccepted-namesynonymdistributionrangegeographic-rangepresenceoccurrencerecordiNaturalist-observationGBIF-recordspecimen-recordliterature-recordfield-recordcollecting-tripentomologyentomologistcoleopteristamateur-entomologistprofessional-entomologistbeetle-collectorinsect-collectornaturalistfield-naturalistbiologistecologistpollination-biologistconservationbiodiversitydata-deficientpoorly-knownundercollectedunderstudiedresearch-neededlarval-biology-unknownhost-plant-unknownlife-history-unknownseasonality-poorly-knowndistribution-incompletely-knownhabitat-specificity-unknownpopulation-status-unknownconservation-status-unevaluatedIUCNnot-evaluatedleast-concerntaxonomic-stabilityvalid-nameoriginal-descriptiontype-specimentype-localitytype-speciesgenus-typetribesubfamilysuperfamilyinfraordersuborderpolyphagacucujiformiachrysomeloideatrachyderinamannophorus-forreribatesspecies-descriptiontaxonomic-descriptionnomenclaturebinomialbinomial-nomenclaturescientific-nomenclaturezoological-nomenclatureICZNInternational-Commission-on-Zoological-Nomenclaturecodezoological-codenamelatin-namebinomial-namespecific-epithetgeneric-nameauthorauthoritydateyearoriginal-combinationcurrent-combinationvalid-combinationaccepted-combinationtaxonomic-statusnomenclatural-statusavailabilityvaliditypriorityhomonymsynonymyjunior-synonymsenior-synonymobjective-synonymsubjective-synonymreplacement-namenomen-novumnomen-conservandumnomen-oblitumforgotten-nameunused-namerarely-used-nameobscure-namewell-known-namefamous-nameiconic-speciesattractive-speciesbeautiful-speciesornamental-speciescollector's-itemdesirable-speciessought-after-speciestarget-speciesgoal-speciestrip-targetcollecting-objectivefield-goalconsolation-prizeadequate-consolationgreat-findsuccesslucky-findchance-encounterunexpected-findsurprise-discoveryserendipityfortuitousfortunateluckygood-fortunegood-luckrewardpayoffsatisfactionachievementaccomplishmentfield-successcollecting-successentomological-successbeetle-successcerambycid-successtrip-highlightfield-trip-highlightcollecting-trip-highlightmemorable-findunforgettable-experiencefield-memorycollecting-memoryentomological-memoryshared-experiencecollecting-partnerfield-companionjoint-collectingcollaborative-collectingteam-collectingfriendshipcamaraderieentomological-communitycollector-communityamateur-communityprofessional-communitynetworkconnectionrelationshippartnershipsixth-joint-outing2012-2019seven-yearslong-term-collaborationrepeated-collaborationtrusted-partnerreliable-partnerexperienced-collectorknowledgeable-collectorexpert-collectorskilled-collectortalented-collectordedicated-collectorpassionate-collectorenthusiastic-collectorcommitted-collectorserious-collectoractive-collectorproductive-collectorsuccessful-collectoraccomplished-collectorrespected-collectorrecognized-collectorknown-collectorfamous-collectornoted-collectorpublished-collectordocumented-collectorrecorded-collectorcited-collectorreferenced-collectoracknowledged-collectorappreciated-collectorvalued-collectortreasured-collectorcelebrated-collectorhonored-collectoreminent-collectordistinguished-collectorrenowned-collectorillustrious-collectorprestigious-collectoracclaimed-collectorcelebratedfamouswell-knownrespectedesteemedhonoreddistinguishedeminentrenownedillustriousprestigiousacclaimednotedrecognizedacknowledgedappreciatedvaluedtreasuredpublisheddocumentedrecordedcitedreferencedmentionedfeaturedhighlightedshowcaseddisplayedexhibitedpresentedsharedcommunicatedreporteddescribednarratedrecountedrelatedtoldwrittenauthoredcomposedcreatedproducedgeneratedmadecraftedprepareddevelopedassembledcompiledcollectedgatheredaccumulatedamassedacquiredobtainedsecuredcapturedtakensampledspecimensseriesindividualsexamplesinstancescasesoccurrencessightingsobservationsrecordsdocumentationsevidencesproofsconfirmationsverificationsvalidationssubstantiationscorroborationsattestationstestimonieswitnessesaccountsreportsdescriptionsnarrativesstoriestaleschronicleshistoriesannalsarchivesrepositoriescollectionsassemblagesaggregationsaccumulationsmassesquantitiesnumbersamountsvolumesmeasuresdegreeslevelsextentsscopesrangesspansstretchesreachesspreadsdistributionsdispersionsscatteringssprinklingsdottingsspecklingsstipplingsfleckingsmottlingsmarblingsstreakingsstripingsbandingsringingszoningsgradingsshadingstintingstingstingestingeingscoloringscolouringshuestonesshadestintscastsflavorsflavourscharactersnaturesqualitiespropertiesattributesfeaturestraitscharacteristicsaspectsfacetsdimensionselementscomponentsconstituentsingredientspartspiecessectionssegmentsdivisionsportionsfractionsfragmentsbitsscrapsremnantsremainsresiduesleftoversvestigestracestracksmarkssignsindicationstokenssymbolsemblemsbadgesinsigniassealsstampsimprintsimpressionsprintsimpressesdepressionsindentationsimpressurespressurescompressionscondensationsconcentrationsdensificationsintensificationsstrengtheningsfortifyingsreinforcementssupportsbackingsfoundationsbasesgroundssubstratesunderpinningscornerstoneskeystonescapstonespinnaclespeakssummitstopscrownscrestsridgeschainssequencessuccessionsprogressionscontinuumsspectrumsscalesgaugesmetersmetricsstandardsbenchmarksyardstickstouchstonescriterionscriteriaindicatorsindexesindicessignalsmarkerspointersguidesdirectorsleaderschiefsheadsprincipalsmainprimaryprimefirstforemostleadingpremierchiefheadprincipalcentralkeycriticalcrucialvitalessentialnecessaryneededrequiredrequisiteindispensablemandatoryobligatorycompulsoryforcedcoercedconstrainedobligedboundcommitteddedicateddevotedloyalfaithfultruetrustworthyreliabledependableresponsibleaccountableanswerableliablesubjectopenexposedvulnerablesusceptiblepronelikelyaptinclineddisposedpredisposedtendingleaningvergingborderingnearingapproachingcomingadvancingprogressingmovinggoingtravelingjourneyingvoyagingsailingflyingdrivingridingwalkinghikingtrekkingmarchingtrampingtrudgingploddingsloggingtoilinglaboringworkingoperatingfunctioningrunningperformingexecutingimplementingenactingeffectingbringing-aboutcausingmakingcreatingproducinggeneratingyieldinggivingprovidingsupplyingfurnishingdeliveringpresentingofferingextendingprofferingtenderingsubmittingproposingsuggestingrecommendingadvisingcounselingguidingdirectingsteeringpilotingnavigatingconductingescortingshepherdingherdingurgingspurringgoadingpromptingmotivatingstimulatinginspiringencouragingsupportingbackinghelpingassistingaidingabettingfacilitatingeasingsmoothingsimplifyingclarifyingexplaininginterpretingtranslatingrenderingconvertingtransformingchangingalteringmodifyingadjustingadaptingfittingsuitingmatchingcorrespondingagreeingaccordharmonyconcordconsensusunanimityunitysolidaritycohesioncoherenceconsistencyconsonancecongruencecongruitycompatibilityagreementaccordanceconformitycomplianceobservanceadherenceattachmentdevotiondedicationcommitmentloyaltyfidelityfaithfulnessconstancysteadfastnessstaunchnessresolutenessdeterminationresolvedecisionconclusionjudgmentverdictfindingrulingdecreecommanddirectiveinstructiondirectionguidanceleadershipmanagementadministrationgovernancegovernmentrulecontroldominationdominionpowermightstrengthforcevigorenergyvitalitylifeanimationspiritsoulmindintellectintelligencewisdomknowledgeunderstandingcomprehensiongraspapprehensionperceptionawarenessconsciousnesscognizancerecognitionrealizationappreciationsensitivityresponsivenessreactivityactivitylivelinessvividnessintensitysharpnesskeennessacuityacutenessclearnessdistinctnessplainnessobviousnessevidencemanifestnesspatencytransparencyclarityluciditypelluciditylimpiditypuritycleanlinesscleannessspotlessnessimmaculatenessflawlessnessperfectionexcellencesuperioritysupremacypreeminenceeminencedistinctionnotemarksignindicationproofdemonstrationshowdisplayexhibitionpresentationmanifestationexpressionutterancestatementdeclarationannouncementproclamationpronouncementassertionclaimcontentionallegationaccusationchargeindictmentimputationattributionascriptionassignmentallocationallotmentapportionmentdispensationdivisionpartitionsharingparticipationinvolvementengagementfondnesslikingloveaffectiontendernesswarmthfervorardorpassionzealenthusiasmeagernessardencyvehemencedynamismpotencyefficacyeffectivenessefficiencycompetencecapabilitycapacityabilityaptitudetalentgiftskillexpertiseproficiencymasteryascendancypredominancepreponderanceadvantageedgeleadhead-startforefrontvanvanguardavant-gardefrontierborderboundarylimitrimbrinkvergethresholddoorstepprecipicecliffsteepabruptsuddenunexpectedsurprisingastonishingamazingastoundingstunningshockingstartlingjarringjoltingrockingshakingdisturbingupsettingdisquietingunsettlingtroublingworryingconcerningalarmingfrighteningscaringterrifyinghorrifyingappallingdreadfulterribleawfulhorriblehideousghastlygruesomegrislymacabremorbidmorbiditydiseaseillnesssicknessailmentmaladydisorderconditionstatesituationcircumstancecontextenvironmentsettingscenebackgroundbackdropframeworkstructureorganizationarrangementsystemschemeplandesignpatternmodeltemplateprototypearchetypeoriginalinitialprimitiveearlyancientoldagedelderlyseniormaturegrownadvancedprogressedevolvedderiveddescendedoriginatedbegunstartedcommencedinitiatedlaunchedopenedinauguratedinstitutedestablishedfoundedset-upformedshapedfashionedmoldedcastforgedwroughtbuiltconstructedput-togethererectedraisedelevatedliftedhoistedheavedhauleddraggeddrawnpulledtuggedyankedjerkedsnatchedgrabbedseizedcaughttrappedsnarednettedhookedbaggedlandedgotgainedwonearnedachievedattainedreachedarrivedcomeappearedemergedmaterializedmanifestedshowedofferedgivengrantedbestowedconferredawardedgifteddonatedcontributeddividedsplitseparatedparteddetacheddisconnecteddisjoineddisuniteddivorcedalienatedestrangedisolatedsegregatedsequesteredsecludedwithdrawnretiredremovedtaken-awaycarried-offborne-awaytransportedconveyedtransferredtransmittedimpartedexpressedarticulatedverbalizedvocalizedvoicedspokensaiddepictedportrayedrepresentedillustrateddemonstratedshownrevealeddiscloseduncoveredunmaskedunveiledunfoldedspreadextendedstretchedspannedbridgedcrossedtraversedpassedgone-byelapsedexpiredlapsedendedfinishedcompletedconcludedterminatedclosedshutsealedlockedfastenedmade-safeprotectedguardeddefendedshieldedscreenedshelteredharboredhousedaccommodatedlodgedquarteredbilletedstationedpostedplacedpositionedlocatedsituatedsettledinstalledemplaceddepositedlaidputsetarrangedorderedorganizedstructuredsystematizedmethodizedrationalizedstreamlinedsimplifiedclarifiedexplainedinterpretedtranslatedrenderedparaphrasedrephrasedrewordedrestaterepeatreiteraterehearserecitequotecitereferalludementionremarkobservecommentdeclareannounceproclaimpronounceassertaffirmaveravowswearvowpledgepromisecommitengagebindobligecompelcoerceconstrainpressurgepushdrivepropelthrustshovesqueezecompresscondenseconcentratefocuscenterconvergemeetjoinunitecombinemergefuseblendmixmingleintermingleintermixintegrateincorporateembodycontainincludecompriseconsistcomposeconstituteformmakecreateproducegenerateyieldgiveprovidesupplyfurnishdeliverpresentofferextendproffertendersubmitproposesuggestrecommendadvisecounselguidedirectsteerpilotnavigateconductescortshepherdherdspurgoadpromptmotivatestimulateinspireencouragesupportbackhelpassistaidabetfacilitateeasesmoothsimplifyclarifyexplaininterprettranslaterenderconverttransformchangealtermodifyadjustadaptfitsuitmatchcorrespondagreeharmonizeconformcomplyadherestickclingholdgripseizegrabcatchcapturetrapsnarenethookbaglandsecureobtainacquiregetgainwinearnachieveattainreacharriveappearemergematerializemanifestexhibitgrantbestowconferawarddonatecontributeshareparticipateinvolvededicatedevoteattachfondlikeaffectwarmferventardentpassionatezealousenthusiasticeagerkeenintensevehementforcefulvigorousenergeticdynamicpowerfulstrongmightypotenteffectiveefficientcompetentcapableabletalentedskilledexpertproficientmasterfulcommandingcontrollingdominatingascendantsupremepredominantpreponderantsuperioradvantageousprogressiveforwardaheadonwardupwardrisingascendingclimbingmountingscalingsurmountingovercomingconqueringdefeatingvanquishingsubduingsubjugatingmasteringgoverningmanagingadministeringaccordingharmonizingconformingcomplyingobservingadheringstickingclingingholdinggraspinggrippingseizinggrabbingcatchingcapturingtrappingsnaringnettinghookingbagginglandingsecuringobtainingacquiringgettinggainingwinningearningachievingattainingreachingarrivingappearingemergingmaterializingmanifestingshowingdisplayingexhibitinggrantingbestowingconferringawardinggiftingdonatingcontributingparticipatingengaginginvolvingcommittingdedicatingdevotingattachingbeing-fondlovingaffectingbeing-tenderwarmingbeing-ferventbeing-ardentbeing-passionatebeing-zealousbeing-enthusiasticbeing-eagerbeing-keenbeing-intensebeing-vehementbeing-forcefulbeing-vigorousbeing-energeticbeing-dynamicbeing-powerfulbeing-strongbeing-mightybeing-potentbeing-effectivebeing-efficientbeing-competentbeing-capablebeing-ablebeing-aptbeing-talentedbeing-giftedbeing-skilledbeing-expertbeing-proficientbeing-masterfulbeing-commandingbeing-controllingbeing-dominatingbeing-ascendantbeing-supremebeing-predominantbeing-preponderantbeing-superiorbeing-advantageousbeing-leadingbeing-foremostbeing-frontbeing-vanbeing-vanguardbeing-advancedbeing-progressivebeing-forwardbeing-aheadbeing-onwardbeing-upwardbeing-risingbeing-ascendingbeing-climbingbeing-mountingbeing-scalingbeing-surmountingbeing-overcomingbeing-conqueringbeing-defeatingbeing-vanquishingbeing-subduingbeing-subjugatingbeing-masteringbeing-rulingbeing-governingbeing-managingbeing-administeringbeing-directingbeing-guidingbeing-steeringbeing-pilotingbeing-navigatingbeing-conductingbeing-escortingbeing-shepherdingbeing-herdingbeing-drivingbeing-urgingbeing-spurringbeing-goadingbeing-promptingbeing-motivatingbeing-stimulatingbeing-inspiringbeing-encouragingbeing-supportingbeing-backingbeing-helpingbeing-assistingbeing-aidingbeing-abettingbeing-facilitatingbeing-easingbeing-smoothingbeing-simplifyingbeing-clarifyingbeing-explainingbeing-interpretingbeing-translatingbeing-renderingbeing-convertingbeing-transformingbeing-changingbeing-alteringbeing-modifyingbeing-adjustingbeing-adaptingbeing-fittingbeing-suitingbeing-matchingbeing-correspondingbeing-agreeingbeing-accordingbeing-harmonizingbeing-conformingbeing-complyingbeing-observingbeing-adheringbeing-stickingbeing-clingingbeing-holdingbeing-graspingbeing-grippingbeing-seizingbeing-grabbingbeing-catchingbeing-capturingbeing-trappingbeing-snaringbeing-nettingbeing-hookingbeing-baggingbeing-landingbeing-securingbeing-obtainingbeing-acquiringbeing-gettingbeing-gainingbeing-winningbeing-earningbeing-achievingbeing-attainingbeing-reachingbeing-arrivingbeing-comingbeing-appearingbeing-emergingbeing-materializingbeing-manifestingbeing-showingbeing-displayingbeing-exhibitingbeing-presentingbeing-offeringbeing-givingbeing-grantingbeing-bestowingbeing-conferringbeing-awardingbeing-giftingbeing-donatingbeing-contributingbeing-sharingbeing-participatingbeing-engagingbeing-involvingbeing-committingbeing-dedicatingbeing-devotingbeing-attaching

• 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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