social wasp family order found united_states mexico bahamas parts canada solitary nest founding queens extended range past decades covers eastern half united_states north expansion typically attributed changing global climate temperatures specific including males workers queens dominance hierarchy distinguished age older wasp higher ranking colony nests queen lays eggs colony similarities worker queen led experiments attempting distinguish characteristics determined males easily identifiable characteristics live relatively small open nests subject predators parasites birds defense recognition strategies help protect predators parasites taxonomy phylogeny subfamily family paper second largest containing species composed entirely social tribes genus currently split distributed world new world subgenus closely related description identification forms united_states typical variable typical form found south eastern united_states inhabits following states north_carolina south_carolina georgia alabama florida texas louisiana oklahoma arkansas kansas colorado arizona form color vary yellow coloring marked yellow normally yellow stripes yellow lines yellow markings head wings yellow color yellow tinged like purple color typical form confused minor variable form red brown coloring middle base abdominal segments outer mid hind black colored wings instead yellow markings variable form pale ivory white markings dispersed body antennae banded red black yellow paper antennae color females fore wing length range males fore wings range extent markings variable distinguishing structural features found separate workers queens male easily identified eyes slender antennae males variable external morphology compared females possible given male females distribution habitat distributed united_states following areas states new_jersey florida florida texas west nebraska colorado arizona california found mexico bahamas observed ontario canada wasp nests typically paper single layer cells open nest size varies upper limit cells outer envelope paper nests indicated nests approximately circular single center usually located nest cells near oldest particularly able new sites disperse new territories found nests located near man structures tend readily build nests near man structures wasp species occupied artificial nesting sites prefers lit open sites nests built wood fiber collected posts plant stems fiber formed paper like cells nests downward held observed occupying artificial nests researchers consisting colony cycle colony cycle longer northern wasp species shorter tropical wasp species respects intermediate cycle occurs march september brood workers emerges july takes fourteen days eggs develop eggs smaller nests laid later season longer develop thirteen eighteen days develop thirteen days develop total weeks egg develop adult reproductive emerge later august september queen queen characteristics lives society queen lays eggs females morphology capability taking egg laying necessary queen death queen migration queen morphology rest workers colony raises question queen role assigned makes special studies found queens males higher levels workers leading higher levels use work increasing concentration cells results greater colder weather found queens survival rate weather compared survival workers day test period queens layer fat surrounding bodies allows live longer winter possibly extend mating season queens ones capable queen development determination aren external morphological differences queens workers internal traits signals wasp social status found determination determined temperature immature stages development experiment emerging june kept long day chambers chambers set hours light hours darkness individuals larger active compared short day chambers set hours light hours darkness thought phenomenon occur juvenile determines suppression workers environmental factors given nests destroyed suffer death queen natural circumstances necessary specific system replacement queen current system place best described furthermore closely related dominance system colony dominance hierarchy older individuals higher rank younger individuals serve lower ranks older female workers generally aggressive forage frequently reflection dominance ranking behavior female workers seen queen longer present colony eldest worker new queen eldest queen replacement queen affects overall females brood allowing future brood related adult workers queen role unlike tend smaller nests typically fewer individuals given nest size nest queen tends active individual nest queen monitor directly control activities nest addition reproductive role queen act nest worker activity points recently disputed workers self organized observed queen act aggressively individuals active development reproduction sex determination system insects species including means males produce identical sperm females produce eggs consequences genetic colonies sisters receive identical sperm share father normal half identical contribution mother results sex biased conflicts queen workers respect sex ratio workers preferring female biased sex ratio queens preferring equal investment queens males sexual investment equal sex ratio obtained investment devoted females female times larger male study found sexual investment female biased especially years high predation nests generally successful seen females produced males female investment bias result ability females worker reproductive nest gives nest increased flexibility compared males important overall nest success females provide colony greater nest defense furthermore original queen longer present sexual investment biased females expected given genetic implications new queen early males workers emerge july emergence reproductive males emerge brood called early males males provide great adaptation allow presence reproductive males nests greater number early males produce greater number workers cells uncertain queen deaths common frequently majority original queens dead july eggs autumn reproductive laid emerged late august september queens remain alive post june order birth future able eldest worker new queen reproductive early males produced deaths queens colony die able continue sexual attraction insects use sexual attract members opposite sex researchers attempted determine exact role sexual play sexual attraction paper set wind tunnel males females exposed sexual isolated males females taken extracts bodies females male extracts taken seventh found opposite sex attracted scent results intensified fan turned range scent found metres nature observed males venture away nest order attract females males press perch attract females females observed visit branches sample found occasions males attracted male behavior ecology presence brood smaller nests colonies compared certain species common nest destroyed queen die prior end season nest fail means predation worker mortality leading workers help rear related sister remain till adulthood allowing female reproductive capabilities plastic predicted worker act workers brood present begin develop characteristics brood present exhibiting aforementioned found true conducted study experimental group eggs larvae removed experimental group began behave like future queens developed characteristic layer fat common queens control group continued behave workers indicative presence brood affects female workers exhibits reproductive recognition nest mate recognition widespread insects physical differences diet nesting materials longer nest mate recently indicated insects ability relatives having lived identical environments non relative counterpart studies indicated nest areas located close near nests following spring social bees able recognize relatives indicated discrimination relatives non relatives having lived identical environments possible found allen experiment different nests box raised identical environments number unrelated groups increased formed box indicates capable relative ability defense nest species aggregate winter large frequently found protected places called varied cracks rocks beneath bark trees walls buildings natural artificial structures provides protection found organ pipe nests mud nest provide certain level protection thought actually house winter worker mortality typical worker live days oldest wasp observed natural colony days old queens typically live times longer workers variation observed life typically attributed colony origin noted variation colonies typically variation original nest satellite nest variation inside single colony occur presence different frequently travel outside nest lead life tend lower life stay nest observed colonies high foraging rates higher reproductive rates compensate lower life interesting aspect observed nest increase female population led lower longevity females focused competing queen foraging fulfilling roles worker death important factor colony failure cause colony failures small colony size colonies small worker longevity crucial survival nest range extension 1950s 1960s extending range spreading united_states following states new state records illinois indiana kentucky maryland missouri new_jersey new_mexico tennessee virginia basis range expansion based exceptional capability colonization new sites solitary nest founding behavior exhibits capability assisted associated tendencies disperse newly available nesting sites species social reproduce near place origin extend range far satellite nests satellite nests common queen fly away original nest settle new location nests create satellites july similar wasp species constructing multitude nests clusters order create protection effect satellites insurance attacks predators nests knocked birds satellites colony survived moving satellite compared reproductive success nests satellite attacked prevalence satellites didn increase effect introduced gave place escape workers older typically start satellite nests developed younger lesser developed workers join satellite establishment distribution workers main nest satellite important needed establishment satellite continued running main nest months main nest abandoned workers follow initial worker new nest likely fail satellite interaction species live social nests open leaves nests susceptible attacks predators attack occur nest brood attempt insert offspring host nest common invasions detect violently bite sting location passed cause vibrations inside nest inside alarmed phenomenon known parasite dance continue hours detected found eaten immediately likely happen lays eggs nest hatch nest result wasp aborted commonly seen late summer abundant second common disastrous effects nest hatch single cell males emerge exit nest wait females emerge female emerge males mate reproduce quickly destroying population cases invading hide larvae nest find important survival hosts attempt remove possible prevent spreading nearby nests defense mechanism proven effective larvae hosts experiment conducted found nests lost brood observed larger satellites greater risk fact large nests typically older relationship age seen fact older queens nests tend younger queens nests predation different predators dangerous predators birds fly knock nest ground phenomenon discovered disappearance nests wasp nests found ground near bird nest found birds eat larvae nest attack kills nest isn effective adults colony attacked adults fly away colony satellite colony attack different attacks birds knocking nest ants nest remove brood ants destroy entire brood unable kill adults predators attack adults foraging traveling satellite nests difficult determine number adults killed away nest difficult track observed queen flight satellite nest isn dangerous reduce idea group living discussed costs benefits alexander predicted cost large groups likely suffer time benefit defense predation theory supported case increase nest size predation rate independent size colony defense facial patterns animals assess rivals specific characteristics relevant fighting abilities animals instead use conventional characteristics determine competitive ability rivals conventional characteristics signals indicative rival abilities reflection animal physical abilities uses facial patterns conventional characteristic determine abilities competitive rivals larger greater brown use determine viability challenging rival larger likely willing challenge rival smaller willing challenge rival assists costs conflict dominance competition nest founding queens alarm response exhibit alarm response typical higher level previously indicated nest mates alarm rest nest movement wings capable rest nest attracting attacking chemical means releasing non species specific alarm chemical signal warn attack releases response post initial attack predator venom released sting venom capable warning coordinating response nest attracting nearby females nearby colonies attack predator helps deter predator attacking especially defense nest certain insects able adapt release means communicating alarm indicating primitive state evolution complex systems communication alarm seen higher social insects experiment extracted female spread venom paper found females attracted venom paper attempted sting response reacted found wasp couldn release response individuals alarmed response rest nest scent strong reach nests neighboring nests alarmed physical movements wing queen response invaded queen forced leave nest satellite established queen queen different nest gives protection nest having invest valuable resources queen use resources new brood
Polistes exclamans