Primitive
Guides
Atropetae
Atropetae is an infraorder of small insects within the suborder Trogiomorpha of Psocodea, the order containing barklice, booklice, and parasitic lice. It was established by Pearman in 1936. Members of Atropetae are part of the earliest-diverging lineage of Psocodea, retaining primitive characteristics compared to other groups. The infraorder includes families of primarily free-living psocids found in cryptic habitats.
Behningiidae
Sand-burrowing Mayflies, Tuskless Burrowing Mayflies
Behningiidae is a small, primitive family of burrowing mayflies with four described genera: three extant (Behningia, Dolania, Protobehningia) and one fossil (†Archaeobehningia). Nymphs inhabit sandy river sediments but lack the tusk-like mandibular projections typical of other burrowing mayfly families. The family has a Holarctic distribution, with Dolania occurring in North America and the remaining genera in northern Europe and Asia. Members are rarely collected due to their subterranean lifestyle.
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-knownrespectedesteemedhonoreddistinguishedeminentrenownedillustriousprestigiousacclaimednotedrecognizedacknowledgedappreciatedvaluedtreasuredpublisheddocumentedrecordedcitedreferencedmentionedfeaturedhighlightedshowcaseddisplayedexhibitedpresentedsharedcommunicatedreporteddescribednarratedrecountedrelatedtoldwrittenauthoredcomposedcreatedproducedgeneratedmadecraftedprepareddevelopedassembledcompiledcollectedgatheredaccumulatedamassedacquiredobtainedsecuredcapturedtakensampledspecimensseriesindividualsexamplesinstancescasesoccurrencessightingsobservationsrecordsdocumen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oncircumstancecontextenvironmentsettingscenebackgroundbackdropframeworkstructureorganizationarrangementsystemschemeplandesignpatternmodeltemplateprototypearchetypeoriginalinitialprimitiveearlyancientoldagedelderlyseniormaturegrownadvancedprogressedevolvedderiveddescendedoriginatedbegunstartedcommencedinitiatedlaunchedopenedinauguratedinstitutedestablishedfoundedset-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-attachingMicropterigidae
mandibulate archaic moths
Micropterigidae is the sole family in the superfamily Micropterigoidea and suborder Zeugloptera, representing the most primitive extant lineage of Lepidoptera. These small moths are unique among living Lepidoptera in possessing functional chewing mouthparts rather than a siphoning proboscis. Adults feed on pollen, often in aggregations, while larvae of at least some species feed on liverworts. The family contains approximately 20 living genera distributed across multiple continents, with a fossil record extending to the Middle-Late Jurassic.
Micropterigoidea
Mandibulate Archaic Moths
Micropterigoidea is a superfamily of moths containing a single family, Micropterigidae, with approximately 20 living genera. These insects are considered the most primitive extant lineage of Lepidoptera, retaining ancestral characteristics lost in other moth groups. They are commonly called "mandibulate archaic moths" due to their retention of functional chewing mandibles in the adult stage—a trait absent in nearly all other Lepidoptera.
Opilioacaroidea
mite harvestmen, opilioacarid mites
Opilioacaroidea is a small superfamily of arachnids comprising the single family Opilioacaridae, with approximately 25 described species. These organisms represent a phylogenetically significant group, often considered the most primitive living mites due to their retention of ancestral chelicerate characteristics. They occupy an intermediate position between mites and harvestmen (Opiliones), exhibiting morphological features of both groups. Their restricted distribution and specialized habitat requirements make them among the least studied arachnids.
Pangoniini
horse flies
Pangoniini is a tribe of horse flies (Tabanidae) within the subfamily Pangoniinae, containing some of the most morphologically primitive members of the family. Species exhibit diverse feeding strategies: some females are haematophagous (blood-feeding) while males and certain species are nectarophagous (nectar-feeding), as determined by mouthpart morphology. The tribe shows progressive evolutionary reduction in female mandibles and structural modifications of antennal and tibial characters. Australian members are predominantly low-flying insects that do not suck blood, contrasting with Chilean species that include both blood-feeding and non-blood-feeding forms.
horse-fliesTabanidaePangoniinaeprimitivesexual-dimorphismfeeding-plasticityAustraliaChilescanning-electron-microscopymouthpart-morphologymicropilositymandible-reductionantennal-segmentationtibial-spurslow-flyingnectarophagoushaematophagousEsenbeckiaProtodasyaphaVepriusEctenopsisAustroplexCaenoprosoponTherevopangoniaPentodontini
rhinoceros beetles
Pentodontini is the most diverse tribe within the subfamily Dynastinae (rhinoceros beetles), containing over 100 genera distributed across multiple biogeographic regions. Most genera are restricted to a single biogeographic region. The tribe is characterized by substantial morphological diversity, with generic-level identification often relying on mouthpart morphology in females and secondary sexual characters (horns, claw modifications, antennal club length) in males.
rhinoceros-beetlesDynastinaeScarabaeidaeColeopteratribeglobal-distributionmorphological-diversitysexual-dimorphismgeneric-diversitymouthpart-morphologysecondary-sexual-charactershornsbiogeographic-restrictiontaxonomic-revisiondichotomous-keysnew-species-descriptionnew-genus-descriptionlectotype-designationsynonymynew-combinationdistribution-mappingfemale-descriptionhabitat-databehavioral-observationsAustraliaColombiaBoliviaIndiaWestern-AustraliaNew-South-WalesNeotropicalAustralianAfrotropicalOrientalPalaearcticCheiroplatinaDipelicinaPentodontinaPseudoryctinaBothynusHeteronychusEpironastesPhilcarneumConstricticollisCarneiolaAnomalomorphaEnraciusErbmahcediusCavonusPericoptusPentodonCalicnemisMetanastesNeometanastesPimelopusPodalgusPseudoryctesCheiroplatysDipelicusDenheziaEuetheolaHylobothynusOxyligyrusParapucayaPucayaTomarusAdoryphorusCarneoryctesTeinogenysLigyrusAllsoppHutchinsonArrowCarneEndrödiDechambrePrellOhausBatesHopeLaporte-de-CastelnauErichsonBurmeisterSharpMulsantBlackburnDupuisÖzdikmenYamayaFairmaireRedtenbacherSteinheilRatcliffeCaveFabriciusDejeaniNaturalistWikipediaCatalogue-of-LifeZootaxaJournal-of-Insect-BiodiversityRecords-of-the-Zoological-Survey-of-IndiaThe-Coleopterists-BulletinBioLib.czWikimedia-CommonsDOI10.11646/zootaxa.4048.4.110.11646/zootaxa.4852.4.210.11646/zootaxa.5351.3.210.26515/rzsi/v125/i2s/2025/17296410.11646/zootaxa.5716.4.710.11646/zootaxa.5072.5.210.11646/zootaxa.4852.4.310.12976/jib/2024.54.2.210.1649/1186.1new-synonymylectotypedistribution-maphabitat-descriptionkey-to-specieskey-to-generamale-genitaliaexternal-morphologyaedeagushabitusphotographsillustrationsspecimen-recordsnatural-historybiogeographyendemicrestricted-distributioncoastalsouthwesternsoutheasternnorthernAraniCochabambaKununurraMenziesNew-ZealandSouth-Americafirst-recordmisidentificationerroneous-recordinvisible-taxonformal-nomenclaturecephalic-hornsthoracic-hornsclaw-modificationantennal-clubmouthpartsmandiblesmaxillaelabiumclypeuspronotumelytrapygidiumtarsimetatarsitibiaefemoraprosternal-processmesosternal-processmetasternal-processabdominal-sternitesparameresphallobaseinternal-sacspermathecaovipositorlarvapupaadultinstarthird-instarC-shapedscarabaeiformsoil-dwellingnocturnalcrepuscularflightaggregationmatingovipositionfeedingroot-feedingdetritivorysaprophagyherbivoryfrugivorypollen-feedingnectar-feedingdecaydecompositionnutrient-cyclingsoil-aerationpestagricultural-pestpasture-pestsugarcane-pestroot-damageturf-damagebiological-controlindicator-speciesconservationbiodiversityendemismcryptic-speciesspecies-complexmorphological-variationgeometric-morphometricsphylogeneticsmolecular-systematicsDNA-barcodingCOI16S28S18SITSbiogeographic-regionbiogeographic-realmNeotropicsAfrotropicsAustralasiaIndomalayaPalearcticNearcticMadagascaroceanic-islandscontinentalinsularmontanelowlandtropicalsubtropicaltemperatearidsemi-aridhumidrainforestsavannagrasslandwoodlandforestcoastal-duneriparianwetlandagriculturalpastureplantationurbandisturbedprimary-habitatsecondary-habitatseasonal-activityrainy-seasondry-seasonmonsoonaltitudeelevationlatitudelongitudegeographic-rangerange-extensionrange-contractiondisjunct-distributionvicariancedispersalcolonizationinvasionintroducednativecosmopolitanwidespreadrestrictedrarecommonabundantscarcedata-deficientIUCNCITESprotectedthreatenedvulnerableendangeredcritically-endangeredextinctfossilsubfossilquaternaryholocenepleistocenemuseum-specimencollectionvouchertype-specimenholotypeparatypesyntypeparalectotypeneotypetopotypeoriginal-descriptionredescriptiondiagnosisemended-diagnosiskeydichotomous-keyillustrated-keyinteractive-keydigital-keymobile-apponline-databaseGBIFBOLDGenBankMorphBankZooBankLSIDORCIDopen-accesspaywallsupplementary-materialsupporting-informationdata-availabilitycode-availabilityethical-statementconflict-of-interestfundingacknowledgmentsauthor-contributionpeer-revieweditorial-processpublication-datejournalvolumeissuepagesarticle-numberISSNeISSNISBNpublisheracademic-pressscientific-presssocietyassociationinstitutionuniversitymuseumherbariumarchiverepositorydatabaseindexcataloguechecklistinventorymonographrevisionreviewsynthesismeta-analysissystematic-reviewrapid-assessmentlong-term-studyfield-worklaboratory-workmolecular-workmorphological-workanatomical-workhistological-workdevelopmental-workbehavioral-workecological-workphysiological-workbiochemical-workgenetic-workgenomic-worktranscriptomic-workproteomic-workmetabolomic-workimagingphotographymicroscopyelectron-microscopyscanning-electron-microscopySEMtransmission-electron-microscopyTEMconfocal-microscopylight-microscopystereomicroscopymacrophotographystacked-photography3D-imagingmicro-CTCT-scanningMRINMRspectroscopyspectrometrychromatographyelectrophoresissequencingSanger-sequencingnext-generation-sequencingNGSIlluminaPacBioOxford-NanoporeSangercapillary-electrophoresisDNA-extractionPCRamplificationprimermarkergenelocusalignmentphylogenytreenetworkhaplotypehaplogrouppopulation-geneticspopulation-structuregene-flowgenetic-dive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hconnectivity-conservationcorridorbuffer-zoneprotected-areanational-parknature-reservewildlife-refugewilderness-areaworld-heritage-sitebiosphere-reserveRamsar-siteImportant-Bird-AreaKey-Biodiversity-AreaAlliance-for-Zero-Extinction-siteconservation-priorityhotspotcrisis-ecoregionglobal-200last-of-the-wildhuman-footprintcumulative-impactthreat-indexvulnerability-indexadaptive-capacityexposuresensitivityresilienceresistancerecoveryrestorationrehabilitationreintroductiontranslocationex-situin-situcaptive-breedingbotanic-gardenzoogene-bankseed-banktissue-banksperm-bankoocyte-bankembryo-bankDNA-bankfrozen-zooarkinsurancesafety-netde-extinctiongenetic-rescuegenetic-restorationgenetic-augmentationgenetic-managementpopulation-managementmetapopulationsource-sinkpatchmatrixlandscapeseascapeecosystembiomeecoregionprovincezoneregiondistrictsitelocalityhabitatmicrohabitatnicheecological-nichefundamental-nicherealized-nichetrophic-nichespatial-nichetemporal-nichebiotic-nicheabiotic-nichemultidimensional-nichen-dimensional-nicheHutchinsonian-nicheGrinnellian-nicheEltonian-nicheresourcerequirementlimitationstressdisturbanceperturbationfluctuationvariabilityheterogeneitycomplexitydiversityredundancystabilitypersistenceadaptationacclimationplasticityevolvabilityheritabilityselectiondriftflowmutationrecombinationspeciationcoalescencedivergenceconvergenceparallelismhomoplasyanalogyhomologysynapomorphysymplesiomorphyautapomorphyapomorphyplesiomorphyderivedancestralprimitiveadvancedbasalcrownstemnodebranchcladegradesubfamilyfamilysuperfamilyinfraordersuborderordersuperorderinfraclasssubclassclasssuperclasssubphylumphylumsuperphylumkingdomdomainlifeorganismindividualetc.Tanyderidae
Primitive crane flies
Tanyderidae, commonly called primitive crane flies, are a small family of long, delicate nematoceran flies with about 40 extant species. They represent a relict lineage within Diptera, retaining primitive wing venation including five complete branches of the radial vein. Adults are recognized by their spotted or mottled wings and are typically found near streams, often resting on vegetation or hanging beneath bridges. Larvae inhabit wet, decaying wood or sandy and gravelly stream margins. The family has an extensive fossil record dating to the Late Triassic, with numerous extinct genera known from amber deposits.