the flame ( axylia putris ) is a moth of the family noctuidae .

it is found throughout europe then east across the palearctic to armenia , western siberia and amur , korea and japan .

the range extends into northern india .

this species has creamy-buff forewings ( sometimes tinged with red ) with black streaking along the costa .

the hindwings are whitish with a dark line along the margin .

the wingspan is 30 – 36 mm .

unusually for a noctuid , this moth rests with its wings wrapped tightly around its body making it resemble a broken twig .

it flies at night in june and july ( sometimes a second brood is produced which flies in september ) and is attracted to light .

flame ( moth )

the crimson flame / first flame moth - as a hunter , defeat 1 flame moth in the ancient ruins . ( 150 stars )
flame moth furore / flame moth frenzy - as a hunter , defeat 3 flame moths in the ancient ruins . ( 400 stars )
episode 3 : the moth and the flame - professor . . . - clickview
ancient ruins - a moth queen may be in its place instead . should this happen , the player may leave the area and re - enter to attempt spawning the flame moth .
moth + flame guides brands entering the vr space through each step of the process including creative , production , distribution strategy and app publishing .
ulgamoth is a combination of ウルカヌス urukanusu ( vulcan ) , 蛾 ga ( moth ) , and moth .
official video for mmddyyyy featuring brandon robbins of the moth & the flame . directed and edited by matt walker , derick smith and brian doherty filmed by d . . .
till recently the flame brocade had been an irregular migrant to the downs around eastbourne . then in 2011 the moth was first recorded in some numbers around horseshoe plantation to the ne of birling gap .
looking for a specific moth species ? enter just part of the name below .
the flame , very common except in the extreme north , flies june - july with an occasional second generation in the autumn . the larvae feed on low growing herbaceous plants such as dock and bedstraw . the flame has a habit of rolling it’s wings to disguise itself as a twig during the day , as the photograph below shows .
the eastern spruce budworm in its adult moth phase , when it migrates to other forest stands .
the new study advances our understanding of the evolution of moth pheromone sensory systems in general and primitive moths in particular .
hopefully this moth can really establish its self on the downs as there is plenty of habitat available for it in the vicinity .
another common micro moth and one of the so - called ‘greys’ which i seem to be getting in large numbers now . this is
or the bramble shoot moth is another common micro that found it’s way into my trap during the night . easily recognised by the chocolate blotch on the forewings , this little moth is on the wing in june and july . the larvae feed on , you guessed it , bramble .
this giant moth with blazing wings has a devastating attack known as < firestorm > , which damages all its foes while also lowering their p . def .
volcarona is based on a moth , particularly the atlas moth or cecropia moth . conceptually , it seems to be derived from the phrase
like a moth to a flame
, which refers to the observed positive phototaxis demonstrated by moths . it may be based on a solar deity . its wings may be based on the petals of a tiger lily , having the same number of extensions , black spots , and red - orange coloring . the black spots on its wings may also represent the dark spots on the sun . the head and antennae of volcarona bear a resemblance to the headpiece of hathor , which depicts the horns of the goddess cradling a disk representing the sun in between . volcarona ' s appearance may also be loosely based on a seraph , a type of angel that is often associated with fire , as its name literally translates to
burning one
. like volcarona , seraphim are often depicted with six red wings and no legs .
in moths , there are four major groups of pheromones classified by their chemistry and how the compounds are biosynthesized . the pheromones of old moth lineages , type 0 , are thought to represent the ancestral state of moth pheromones . type 0 pheromones all have short carbon chains and they are remarkably similar to many common plant volatiles .
it wouldn’t be moth diaries without the introduction of yet another new pug moth . this time it’s the turn of the grey pug . what is there to say ? it’s a pug and it’s grey . it flies in may and june and sometimes again in august . the larvae eat the leaves and flowers of a number of herbaceous plants .
this page lists the moth species recorded at the site , mostly on the unimproved limestone grassland slopes of rough bank sssi but also any additional species recorded in the two pasture fields which were included in bc ' s purchase . these fields both have wetland areas with plants associated with such habitat and supporting colonies of some rush - feeding moth species .
now for the first time , jothi yuvaraj and colleagues at lund university , sweden , have identified the corresponding pheromone receptors ( prs ) from a primitive leafminer moth , called eriocrania semipurpurella .
or the garden grass - veneer . flying in june and july it is probably britains most common grass moth . the larvae feed on the base of a number of grasses and grass stems .
in moths , there are four major groups of pheromones classified by their chemistry and how the compounds are biosynthesized . the pheromones of old moth lineages , type 0 , are thought to . . .
or the hook - streak grass - veneer , a micro moth common throughout britain and one of many ‘grass moths’ . flies may - august with the larvae feeding on a range of grass stems .
or the crescent bell is a fairly common micro moth throughout britain and wherever birch is present . on the wing in june and july the larvae feed on the catkins of birch throughout the winter .
altermatt and ebert suggest this might be good news for moths and other nocturnal insects . if insects can evolve reduced attraction to light , perhaps light pollution won ' t be as dangerous for these species . but they caution that their results also show that as light pollution has caused systematic changes in animal behaviour to evolve , it may take several generations before moths see the benefits of efforts to reduce light pollution . in addition , it is unclear how the reduction in attraction of the moths to light might impact the insect ' s ecology . our light pollution has taught the moth to avoid the flame , but perhaps at a cost .
you met ‘dark arches’ in episode 8 , well now we meet ‘light arches’ . flying june - august it’s a fairly common moth . the larvae feed on the roots and stems of a range of grasses .
the authors , therefore , propose that not only have the pheromone receptors of this basal moth evolved from ors that recognize plant odorants but that the same might be true of the canonical pheromone receptors of more derived moths .
a stunning moth and one of my favourites , the swallow - tailed moth is actually fairly common , but being strictly nocturnal and having quite a short emergence period from late june and throughout july , it is not often encountered by the non - enthusiast . the first two of the year ( the one ( poorly ) photographed above included ) were found in my trap this very morning . the larvae feed on a range of trees but have a preference for ivy .
meet the ‘setaceous hebrew character’ . you might remember the ‘hebrew character’ from my march moth diaries but this is an altogether different moth , although sharing the black arch pattern which gives both moths their name . like the hebrew character , the setaceous version is very common too . small numbers appear in may and july with far more on the wing in august and september . in the north there is just a single generation , flying july - august . the larvae feed on nettle .
the common white wave is an elegant , fairy - like moth common throughout britain . it occupies woodland , copses and similar habitats , on the wing from may – august in two generations , the larvae feed on deciduous trees , mainly birch .
i found this common footman lurking in my trap this morning . a smooth - scaled alien looking moth , it is on the wing usually from july to august . the larvae feed on lichens , which makes grave yards a particularly favourable habitat .
butterfly conservation ' s rough bank reserve is a site known by some local butterfly enthusiasts for many years , but until 2012 ( the year when bc purchased rough bank ) there had been few reports of moths found there apart from daytime observations on a few occasions . one such record in 1993 was of a larva of narrow - bordered bee hawk - moth found feeding on devil ' s - bit scabious . five nationally scarce micro - moth species had also been recorded at the site in the past .
the species name links in the table below are to the species entries on the uk moths website . please note that the gloucestershire moth distribution maps are not updated frequently , so some species listed below may not yet be shown as recorded in the rough bank tetrad so90e . also some informal reports of species from rough bank are listed below , but such records will only be mapped if the recorder sends them to the vc33 ( east gloucestershire ) moth recorder - for contact details see the distribution maps page .
or green oak tortrix is a micro moth . it is a very common moth , on the wing from may to july , inhabiting oak woodland , it can become a pest . although oak is the main food plant , other deciduous trees are also used , the larvae feeding in a rolled - up or folded leaf . these abundant larvae sometimes completely defoliate trees . curiously , the same day this moth hit my trap , it was also recorded for the first time this year by other leicester mothers , including the county recorder , adrian russell , who caught a total of 34 in one night despite usually only catching single figures each year . there is very little oak in mine or adrian’s vicinity , suggesting some kind of local movement or a switch to a new food - plant .
robinson , g . s . , ackery , p . r . , kitching , i . j . , beccaloni , g . w . & hernández , l . m . 2001 . hostplants of the moth and butterfly caterpillars of the oriental region . 744 pp .
or the poplar shoot . you might think that you’ve seen me post this moth before , especially being as there are many of these ‘bird dropping’ type micro moths . i definitely haven’t though , as this is a real local rarity , with only two other recent records for the whole of leicestershire and rutland . not that i knew it , as i emptied out the trap this morning , setting it aside as just another micro moth ‘to be id’d’ . flying in june and july it is a relatively local species , occurring sporadically throughout england and is classed nationally as ‘scarce’ . the larvae feed on poplar leaves as the name would suggest . despite the beauty of many of the moths above , it seems this minuscule moth is easily the best catch of the day , and one of the most important records i’ve had to date .
for each species the table below lists the agassiz , beavan & heckford checklist number , the bradley list number , scientific and common names , national status , earliest and most recent years recorded and typical uk larval food . the list will be updated at least annually using records sent to the current vc33 ( east gloucestershire ) moth recorder , robert homan . it is updated more frequently with my own records from the site , and sometimes with additional species reported to me . records prior to 2012 are from the former gloucestershire moth recorder roger gaunt ' s database .
since 2012 a total of 628 moth species have been recorded on the site ( plus 4 other species only recorded in earlier years ) . three of these were the first records for the species in gloucestershire : trifurcula headleyella and coleophora therinella in 2012 , and aphelia viburnana in 2014 .
the phoenix is a beautifully marked moth . it is found throughout most of the british isles but is not particularly common anywhere , although at least one has found my garden . usually flies from july - august so this one was early . the larvae feed on the leaves of red and black currant .
robinson , g . s . , ackery , p . r . , kitching , i . j . , beccaloni , g . w . & hernández , l . m . 2001 . hostplants of the moth and butterfly caterpillars of the oriental region . 744 pp . southdene sdn bhd , kuala lumpur .
the phrase is a simple allusion to the well - known attraction that moths have to bright lights . the word moth was used the the 17th century to mean someone who was apt to be tempted by something that would lead to their downfall . this is referred to by shakespeare in the merchant of venice , 1596 :
robinson , g . s . , ackery , p . r . , kitching , i . j . , beccaloni , g . w . & hernández , l . m . 2002 . hostplants of the moth and butterfly caterpillars of america north of mexico . memoirs of the american entomological institute , 69 : 1 - 824 .
robinson , g . s . , ackery , p . r . , kitching , i . j . , beccaloni , g . w . & hernández , l . m . 2002 . hostplants of the moth and butterfly caterpillars of america north of mexico . 824 pp . [ memoirs of the american entomological institute , volume 69 . ]
but that’s not the case . originally not a british species , this moth was accidentally introduced with fruit and vegetables and appears to be established and expanding its range . uncommon but increasing in leicestershire , the larvae feed , strangely , on a mix of mosses , dead insects , detritus , decaying vegetable matter and also on seedpods of tree lupin and tans ; no wonder it’s thriving .
it ' s a common literary trope , but moths really are attracted to the flame . and that usually goes poorly for the insect , with death a common outcome – either by burning or by predators that use the light source as a buffet table . as humans occupy an increasing share of the planet , we are lighting up ever more portions of the night sky , which may affect nocturnal species like moths that are attracted to light . swiss researchers florian altermatt and dieter ebert , based in zürich and basel , thought that moths may be able to evolve changes in their behaviour to resist anthropological changes in light conditions . as artificial light can be so dangerous to moths , they hypothesized that moths from areas with bright lights at night might be less attracted to lethal light than moths from dark areas as a result of evolutionary changes in their behaviour .
despite the joy of finding the swallow - tailed moths , i was also a little gutted this morning . when i checked the trap prior to bed last night , a stunning lime hawk - moth has found its way in , however by morning it appears to have found it’s way out again meaning i was left a little bereft and without the standard ‘look how big this hawk - moth is on my hand’ photo - shot . the picture above shows the beauty , but not the scale . occupying suburbia it flies in may and june and is fairly common in the southern half of the country , including leicestershire . famously common in london , owing to an array of still tree - line streets and avenues , in recent years its distribution has spread northwards and is now regularly found well into north yorkshire and beyond . the larvae feed on lime but also birch , alder and elm .
the coronet ( photos below ) appears in two distinct varieties , a grey - scale or the mossy green as below . i was thankful for the mossy green ; what a stunning moth ! it’s also a local rarity , though increasing year on year in numbers , so it’s a good record to have . the habitat preferences are woodland , commons , downland and marshy places , and the flight period is during june and july . the larvae feed on ash and privet .
hosts brings together an enormous body of information on what the world ' s butterfly and moth ( lepidoptera ) caterpillars eat . the web - based version presented here offers a synoptic data set drawn from about 180 , 000 records comprising taxonomically ' cleaned ' hostplant data for about 22 , 000 lepidoptera species drawn from about 1600 published and manuscript sources . it is not ( and cannot be ) exhaustive , but it is probably the best and most comprehensive compilation of hostplant data available .
volcarona is a large , moth - like pokémon with four small feet . it appears to have compound eyes that are two shades of blue , and there is a red horn - like structure on each side of its face . the fuzz on its upper body is smoky white and tangled . its abdomen is black in the center and light blue with black dots on the outside . it has six leaf - like wings that are orange in color and have small black spots .
cousin to the heart and dart , this recent arrival is the heart and club . unlike the heart and dart ( which has been filling my trap in droves ) , this moth is distributed mainly in the south and south - east of england . whilst it is common in places , it has a scattered distribution elsewhere . it is frequent but not common in leicestershire and rutland . flying in june - july the larvae feed on growing leaves and roots of knot - grass and clover .
the elephant in the room and resting gently on my finger , it is of course the elephant hawk - moth . this little gem is one of our more common hawk - moths and can be found in hedgerows , gardens , woodland edges and heathland , and on rough grassland and sand - dunes . practically anywhere ! it is a real feast for the eyes . flying from may - july the larvae feed on rosebay willowherb and bedstraws . the large body is also vibrant pink on the underside .
to test this , the researchers set out collecting young caterpillars of the ermine moth yponomeuta cagnagella from cities and towns in switzerland and france that varied in their night - time light levels . they reared these caterpillars to adult moths in a laboratory where all the animals received the same light levels . they then set up an indoor flight cage with a light trap at one end , left the moths overnight to flutter to the light , and counted how many were collected by the trap in the morning to test how attracted to the light the moths were .
now , i know it’s only been a week since the last moth diaries ( and you are probably all getting very bored ) but , as i warned with the onset up summer , the moths are arriving thick and fast . 28 new species have made the garden a food stop since a week ago , including 2 county rarities , taking my garden total to 128 species for the year so far . again , many of the pictures below are stock images unless otherwise stated and the information is kept to a minimum so that it doesn’t drag on…enjoy !
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gaden s . robinson , phillip r . ackery , ian j . kitching , george w . beccaloni and luis m . hernández
records of caterpillar hostplants are scattered through published and manuscript sources worldwide and are difficult to retrieve . many rearing records are never published and so are not accessible to other entomologists . but collected hostplant records form a valuable scientific resource that can be used eventually to answer broader biological questions about how lepidoptera and plants interact ( eg , letourneau , hagen & robinson , 2001 ) . it provides information of immediate relevance to agriculture , ecology , forestry , conservation and taxonomy .
we hope that it will be useful to a wide range of biologists and that it will act as a spur to further recording and analysis of caterpillar - plant interactions .
hosts can be searched in two ways , using text search and drill - down search modes .
in text search mode , use either lepidoptera or hostplant criteria or a combination of the two . hosts operates only using scientific names . it is not possible to search for the hostplants of the red admiral butterfly but a search for hostplants using its scientific name , vanessa atalanta , will be successful . enter the generic name ( vanessa ) in the - lepidoptera criteria - genus box ; enter atalanta in the species : box ; click search .
hint : leave the ' starts with ' command as the default and in the species entry box omit the last few letters of the species - name ( eg , atalant ) . this will get around the problem of variable gender - endings . hosts uses original orthography of species - group names as far as possible , but some checklists follow the convention of altering the species - group name to accord with the presumed gender of the generic name ( eg flava , flavus ) .
restrict or refine searches using additional criteria ; choosing ' usa ' from the drop - down location box and entering [ starts with ] ' urt ' in the hostplant family box will return hostplant records of vanessa atalanta from urticaceae in the usa .
hint : restricting location may deliver a very incomplete subset of records : in the previous example all records specified as from the nearctic region ( usa + canada ) would be missed .
drill - down search mode allows the user to home in from the starting - point of either the lepidoptera or the plant family . choose a family group from the drop - down box and allow time for all genera of that family in the database to load to the genus drop - down box ; choose a genus and wait for the species to load . the search button can be pressed at any time , but the record delivery limit may be exceeded at higher taxonomic levels . drill - down search allows the user to see by scrolling all the taxa that are represented in the database . following the previous example , choose nymphalidae , then choose vanessa from the genus drop - down box and then atalanta from the ten possible species of vanessa included in hosts .
the preliminary search results pages give family , genus and species of the caterpillar and its hostplant . note that many hostplants are recorded as a plant genus only . clicking the caterpillar name will give the full record . the full record listing gives , additionally , the author of the lepidoptera species , subspecific information on the insect and the plant , if available , together with details of larval damage . laboratory rearings where the food utilised may not be the natural hostplant are indicated . the status of the record ( whether considered true , erroneous or suspect ) is not currently implemented in this version of hosts .
leguminosae ( c ) - caesalpinioideae . leguminosae ( m ) - mimosoideae . leguminosae ( p ) - papilionoideae .
all lepidoptera groups are covered and subspecific taxa are differentiated . the original source of the record , original form of the names used ( prior to taxonomic ' cleaning ' and standardisation of nomenclature ) and validation and verification fields are not included . this information is , however , retained in the databases used to generate this site . while we have included species that do not feed on green plants , the known food substrates of these are not listed exhaustively . such species comprise detritophages and predators and include , for example , most tineidae and the stored - products pests . the published compilations from hosts ( see more detail - publications from hosts ) include record status and the sources of all records .
detailed published compilations from hosts are available in press . these books give greater detail than the website , together with comprehensive cross - indexes , record status and full bibliographies . they are indispensable tools for naturalists and professional entomologists .
beccaloni , g . w . , viloria , a . l . , hall , s . k . & robinson , g . s . 2008 . catalogue of the hostplants of the neotropical butterflies / catálogo de las plantas huésped de las mariposas neotropicales . m3m - monografías tercer milenio , volume 8 . zaragoza , spain : sociedad entomológica aragonesa ( sea ) / red iberoamericana de biogeografía y entomología sistemática ( ribes ) / ciencia y tecnología para el desarrollo ( cyted ) / natural history museum , london , u . k . ( nhm ) / instituto venezolano de investigaciones científicas , venezuela ( ivic ) . 1 - 536 pp . , 1 fig , 3 tabs .
gaden robinson was responsible for the overall project design and management of the hosts database , and for records of lepidoptera exclusive of butterflies and bombycoid moths . phillip ackery and george beccaloni were responsible for butterfly data , including data drawn from card catalogues developed by ackery , whilst ian kitching was responsible for hostplant data of bombycoid moths . luis m . hernández was responsible for abstracting in the latter two years of the project and for development of the bibliography for the hardcopy versions of the data .
we are extremely grateful to the many people who contributed their own rearing records of lepidoptera or personal accumulations of data for inclusion in the hosts database , particularly mike bigger ( uk ) , john w . brown ( usa ) , chris conlan ( usa ) , rob ferber ( usa ) , konrad fiedler ( germany ) , jeremy holloway ( uk ) , frank hsu ( usa ) , jurie intachat ( malaysia ) , alec mcclay ( canada ) , bill palmer ( australia ) , pierre plauzoles ( usa ) and the generous individuals who contributed rearing records through the worldwideweb and who are known to us only as an email address .
we are particularly grateful to julian donahue and the los angeles county museum of natural history for allowing us to include data on microlepidoptera from the card catalogue prepared by the late j . a . comstock and c . henne , and for access to manuscript records by noel mcfarland .
marian fricano ( santa clara university ) and aileen giovanello ( clark university , international internship 1996 ) made substantial contributions of abstracted data ; fran love ( north carolina ) painstakingly checked scanned texts and reformatted them for import to paradox .
much of the immense task of data abstracting and entry from printed and manuscript sources as well as preliminary editing and name - checking was carried out by volunteers . many of these were school students on work - experience placements during 1993 - 2000 from , initially , the coopers ' company and coborn school , upminster , and later from other schools in greater london : christopher andrewes , simon bennett ( 1994 nhm vacation studentship ) , steven bond , michael brownlow , emma causer , laurence cooper , ailsa cranfield ( 1998 nuffield studentship ) , emily dwiar , andrew enever , jane feehan , madeleine ferry , max friedman , edward gold , jennifer hodgkinson , christopher joint , fateha khatun ( 1996 nuffield studentship ) , james lowe , louisa marchant , gemma millward , christopher milne , carolyn oughton , william perkins , rebecca reith , eleanor resheph , clare sambidge , neil shaftain , stephen sloan , helen stevens , samuel tarry , david taylor and thomas yeatman .
we are indebted to all our helpers for their diligence , accuracy and patience , and for their unswerving faith that this daunting project would reach a conclusion .
our colleagues here and overseas provided substantial help , advice and assistance with the checking of names of lepidoptera and with many other aspects of this project : kim and david goodger and jeremy holloway ( macrolepidoptera families ) , martin honey ( noctuoidea ) , brian pitkin ( computing ) , malcolm scoble and linda pitkin ( geometroidea ) , klaus sattler ( gelechioidea ) , michael shaffer ( pyraloidea , thyridoidea , pterophoroidea ) , alma solis ( usda , washington - pyraloidea ) , fernley symons ( oxford university - technical support ) and kevin tuck ( tortricoidea ) . julie harvey and the staff of the bmnh entomology and general libraries provided sterling support in locating obscure source material and intuitively correcting bowdlerized references .
we thank the trustees of the loke wan tho memorial foundation for their generous support for this project .
robinson , g . s . , p . r . ackery , i . j . kitching , g . w . beccaloni & l . m . hernández , 2010 . hosts - a database of the world ' s lepidopteran hostplants . natural history museum , london . urltoken . ( accessed : 18 aug . 2010 ) .
brummitt , r . k . 1992 . vascular plant families and genera . [ vi ] + 804 pp . , royal botanic gardens , kew .
kartesz , j . t . 1994 . a synonymized checklist of the vascular flora of the unites states , canada and greenland . timber press , portland . 1 , checklist , lxi + 622 pp ; 2 , thesaurus , vii + 816 pp .
kitching , i . j . & cadiou , j . - m . 2000 . hawkmoths of the world : an annotated and illustrated revsionary checklist . xx + 500 pp . , cornell university press , ithaca .
kitching , i . j . & rawlins , j . e . 1999 . the noctuoidea . pp . 355 - 401 . in : kristensen , n . p . ( ed . ) lepidoptera , moths and butterflies . 1 . evolution , systematics and biogeography . handbook of zoology , 4 ( 35 ) . lepidoptera . x + 491 pp . de gruyter , berlin .
letourneau , d . k . , hagen , j . a . & robinson , g . s . 2001 . bt crops : evaluating benefits under cultivation and risks from escaped transgenes in the wild . pp . 33 - 98 . in : letourneau , d . k . & burrows , b . e . ( eds ) , genetically engineered organisms : assessing environmental and human health effects . [ viii ] + 438 pp . , crc press , boca raton .
mabberley , d . j . 1987 . the plant book . a portable dictionary of the higher plants . xii + 707 pp . , cambridge university press . [ the 1993 reprint edition is currently used in editing . ]
nielsen , e . s . , edwards , e . d . & rangsi , t . v . ( eds ) 1996 . checklist of the lepidoptera of australia . monographs on australian lepidoptera . 4 . xiv + 529 pp . , csiro , melbourne .
nye , i . w . b . ( ed . ) 1975 - 91 . the generic names of moths of the world . 1 : 568 pp . ( noctuoidea ( part ) - nye , i . w . b . , 1975 ) ; 2 : xiv + 228 pp . ( noctuoidea ( part ) - watson , a . , fletcher , d . s . & nye , i . w . b . , 1980 ) ; 3 : xx + 243 pp . ( geometroidea - fletcher , d . s . , 1979 ) ; 4 : xiv + 192 pp . ( bombycoidea to zygaenoidea - fletcher , d . s . & nye , i . w . b . , 1982 ) ; 5 : xv + 185 pp . ( pyraloidea - fletcher , d . s . & nye , 1984 ) ; 6 : xxix + 368 pp . ( microlepidoptera - nye , i . w . b . & fletcher , d . s . , 1991 ) . british museum ( natural history ) / the natural history museum , london .
rawlins , j . e . 1984 . mycophagy in lepidoptera . pp . 382 - 483 . in : wheeler , q . & blackwell , m . ( eds ) fungus - insect relationships . perspectives in ecology and evolution . 514 pp . , columbia university press .
robinson , g . s . 1999 . hosts - a database of the hostplants of the world ' s lepidoptera . nota lepidopterologica , 22 : 35 - 47 .
scoble , m . j . ( ed . ) 1999 . a taxonomic catalogue to the geometridae of the world ( insecta : lepidoptera ) . 2 vols . csiro publications , melbourne .
royal botanic garden edinburgh dipterocarpaceae database : http : / / 193 . 62 . 154 . 38 / diptero /
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a common species over most of britain , except in the far north where it is more local . the adults rest with the wings wrapped around the body and closely resemble a broken piece of twig .
it inhabits woodland fringes , hedgerows and suburban habitats , and flies mainly in june and july , sometimes with a partial second generation in the autumn .
ukmoths is built , run and maintained by ian kimber , with thanks to the many kind contributors who provide photos and information .
the ukmoths facebook page is a great place to post your identification queries . more often than not you ' ll get a positive id on most photos fairly quickly .
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from the album
hardwired . . . to self - destruct
available everywhere on november 18 . pre - order now at metallica . com . directed by tom kirk director of photography - josh blackman edited by nick cortes produced by hesper lucent vfx by keith simpson , james pina & rowan glenn © 2016 blackened recordings follow metallica : urltoken urltoken urltoken urltoken urltoken urltoken
kino v tsoi - kukushka ( cuckoo bird ) ( edited by . a . sargsyan # 2013 )
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cloud strife : i couldn ' t finish ' em . looks like this ' s gonna get complicated .
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you’ve survived an alien invasion and wound up in prison , where you are told your memories aren’t real . is your mind playing tricks on you , and why does your captor keep appearing in your visions ? an immersive look at what we believe and how we can be convinced it isn’t true .
to help mtv launch their first vr initiative at comic - con 2016 , teen wolf vr takes fans into the world of the hit series and puts them face to face with this season’s new villain . using our gaze - activated technology , the user will be place into a scene where the choices they make will determine if they live or die . . .
an immersive journey into the world of the music video where users can have a first - of - its - kind encounter with the song
style
in a way they had never imagined .
the keynote demo at 2016 vrla , believe vr was the first virtual reality piece to bring gaze - activated content into a live action , narrative scene . the piece places users inside a scene such that the actors in the scene change their behavior according to where the user looks .
to give fans a chance to witness marco marco’s annual drag queen fashion show , we shot this 360 video in unique way that both gave the immersive feel of being right in the middle of the action and at the same time take advantage of traditional cinematic editing techniques to match the pace and vibe of the high energy show .
to help launch her debut album , we produced a 360 music video for actress taryn manning ( orange is the new black ) . we used projection photography to develop an aesthetic for the piece unlike any other 360 video .
imagine dragons has some demons . . . glad we could share this moment together !
last show of the night visions european tour and our friend ben of imagine dragons pays us a special visit during our set . . .
bass player goes for the gig of his life . you won ' t believe what happens next . . .
i had the amazing privilege of singing guest vocals on this song , written by my good friend matthew walker for his band of1000faces . check out the music video for it below and maybe you ' ll see me try to do my best
blue steel
provo ut : tomorrow for halloween , some of my close friends and i are dressing up as pearl jam and performing their album
vitalogy
from front to back ! i ( brandon ) will be eddie vedder ( my idol ) and singing every song . it ' s a
costume party
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for moths , love is literally in the air through the action of pheromones to attract mates .
virgin females release a perfumery concoction , specially blended to attract males from the same species , even over long distances .
to date , little is known on how males evolved to heed their siren ' s call .
in general , pheromone compounds in moths and other insects are detected by specialized receptors that generally do not respond to plant volatiles .
pheromones and other odorants are detected by odorant receptors ( ors ) expressed in olfactory sensory neurons found most prominently within the insect antennae .
then , they show that these receptors also respond to plant odors and propose a scenario in which pheromone receptors evolved from plant odor receptors .
our results suggest that prs for type 0 pheromones have evolved from ors that detect structurally - related plant volatiles ,
said professor christer löfstedt .
they are unrelated to prs detecting pheromones in advanced lepidoptera , which , in turn , also independently may have evolved a novel function from ors detecting plant volatiles .
our results suggest that sex pheromone receptors in lepidoptera have evolved sex pheromone detecting functions from ors detecting plant volatiles on multiple occasions ,
said jothi yuvaraj .
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to date , little is known on how males evolved to heed their siren’s call .
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the duo found that female moths were much less likely to be trapped than male moths and that , in agreement with their hypothesis , moths from bright areas were much less likely to be trapped than moths from dark areas . this suggests that the moths from bright areas have undergone selection to avoid lethal bright city lights . as the researchers had collected the moths as young caterpillars , they think this response must be a genetic difference , rather than a difference in light exposure during development .
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photo credit : t . - c . francis pan some oyster larvae grow faster than others , but now donal t . manahan and colleagues reveal that the fastest growers are marked out by their high protein synthesis rates .
“one of the underappreciated benefits of fellowships is the act of applying for them , because you have to write and articulate your ideas . ”
in our latest early - career interview , we chat to simon sponberg , assistant professor at the georgia institute of technology , usa . he shares the story of his career , beginning with how he combined a love of physics and biology by studying how geckos stick to walls .
a new review from craig p . mcgowan and clint e . collins looks at the ecology , biomechanics and evolution of bipedal hopping in mammals , with a focus on why bipedal hopping has arisen in multiple clades of mammals .
photo credit : ari friedlaender not all orca species prey on aquatic mammals , so how do delphinids know when they are at risk ? a new study shows that pilot whales and risso’s dolphins flee from a subset of orca calls that share acoustic characteristics with other mammal alarm calls , including human screams . this article was featured in science magazine .
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the eastern spruce budworm in its young caterpillar ( larvae ) phase , feeding on fir tree needles .
in the last decade , 7 million hectares of boreal forest in eastern canada have been destroyed by the spruce budworm , leaving devastation and fires in its wake .
take a walk in the boreal forest of eastern canada these days , and chances are you ' ll hear the sound of millions of caterpillars taking a dump . they ' re the spruce budworm , and they ' re on the move .
it ' s a bit yucky ,
said patrick james , a spatial ecologist with a doctorate in forest ecology who ' s a professor of ecological modeling in the department of biological sciences at université de montréal .
if you walk in the woods in an area that ' s being severely defoliated , it sounds like rain ,
said james , 37 , who has a new scientific paper published on the budworm phenomenon .
it ' s all of their frass , the bug poo , falling through the canopy of the trees .
there ' s a huge consequence for the forest industry ,
james said .
the budworm changes the composition of the forest , it denudes the trees and leaves behind huge areas of standing dead and dry timber . most of those trees don ' t get harvested , they don ' t go to the sawmill , profits aren ' t made .
budworm outbreaks will cost the industry in new brunswick alone an estimated $ 3 billion to $ 4 billion over the next 30 years , according to a 2012 study by university of new brunswick researchers . timber revenue will be lost , so will jobs , and the consequences will amplify as budworm , grown into dime - size moths , head south .
in his study , published in march in the u . s . journal ecological applications , james shows that defoliation increases the risk of natural fires igniting eight to 10 years after a budworm outbreak – especially now , in the spring , before summer fire season starts . interestingly , this risk actually decreases in the years immediately after an outbreak , since the
greenup
of ground vegetation keeps the soil moist and less likely to ignite .
for his study , james looked at defoliation data stretching back to 1963 in two vast ecosystems in eastern and central ontario . these results suggest that fire management agencies , which normally rely on weather indicators , could also include defoliation data to better predict areas of high fire risk .
knowing when and where the fire risk is increased due to the budworm can lead to pro - active techniques like
salvage logging ,
harvesting dead trees from areas already defoliated .
if you can reduce the amount of budworm - killed forests in an area , you would then reduce the risk of fire ignition , which would then reduce the probability of having large forest fires ,
james said .
the issue of how budworm affects fire is expected to become even more important in the future as both fire and insect activity is expected to increase due to climate change . however , how they interact and what sort of damage may result , remains uncertain .
spruce budworm outbreaks happen every 35 years or so ; the last one peaked in the early 1980s and the latest one began in quebec in 2006 . natural resources canada warned last fall that while the impact is greatest on quebec ' s north shore and in the gaspé and saguenay - lac - saint - jean , where 7 million hectares have been defoliated , budworm has been spotted in new brunswick and may spread to the acadian forest .
the process works like this . first , the budworm larvae eat away at the tops of trees – balsam fir , black and white spruce . after several years , the dead treetops break off in the wind and their debris builds up on the branches below . lightning strikes the dry material , which ignites , while meanwhile down on the forest floor an accumulation of needles lies ready to ignite , too .
unseen in urban environments like montreal , the budworm are nevertheless getting closer . they can be spotted in quebec ' s mauricie region , an hour north of trois rivières , and around ville marie , south of rouyn - noranda in the abitibi region .
we ' re at an interesting point in the outbreak ,
said james , who continues to monitor the phenomenon .
last year it increased , but at a slower rate . in quebec ' s north shore region , around baie - comeau , the budworm ate all they could , and with radar we detected great clouds of them on the move , migrating on the wind to areas that hadn ' t been so heavily affected , largely towards the south to the lower st . lawrence , the gaspé and new brunswick .
the moths are now
hovering on the new brunswick border around campbellton ,
he added , referring to a mass infestation that hit there last summer , when millions of larvae turned into moths and flew into town or were carried there by the wind , covering everything for miles around .
with that development , i would expect there ' ll be more activity in northern new brunswick this year .
can ' t the budworm be killed before they do their damage ? yes , but it ' s very expensive .
in the 1950s , authorities tried to contain budworm outbreaks by spraying infested areas with ddt . with that toxic method no longer an option , authorities now resort to a budworm - specific bacteria called bt . the problem is price : ddt cost only a few cents per hectare to spray , whereas bt costs $ 40 per hectare .
if you have $ 1 million worth of wood , it will cost you $ 1 million to spray it ,
james said .
the cost is prohibitive .
three summers ago , canadian forest services appealed to the public to help track budworm in eastern canada . several hundred
citizen scientists
now lure , trap and count moths in six provinces , as well as maine , uploading the data on an app and sending the dead moths to natural resources ' main research laboratory in fredericton , n . b .
it ' s a kind of early - warning system , suggesting how and where moths
disperse
and in what number .
the value is two - fold ,
said james , who ' s involved in the program at an analytical level , using spatial modeling and genetics .
it helps us understand if budworm populations are increasingly locally , which is a problem , or are migrating from somewhere else , flying in and then dying , which is less of a problem .


macrothemis is a genus of dragonfly in the family libellulidae .

it contains the following species : macrothemis absimilis costa , 1991 macrothemis aurimaculata donnelly , 1984 macrothemis belliata belle , 1987 macrothemis brevidens belle , 1983 macrothemis calliste ( ris , 1913 ) macrothemis capitata calvert , 1909 macrothemis celeno ( selys in sagra , 1857 ) - antillean sylph macrothemis cynthia ris , 1913 macrothemis declivata calvert , 1909 macrothemis delia ris , 1913 macrothemis extensa ris , 1913 macrothemis fallax may , 1998 macrothemis flavescens ( kirby , 1897 ) macrothemis griseofrons calvert , 1909 macrothemis guarauno rácenis , 1957 macrothemis hahneli ris , 1913 macrothemis hemichlora ( burmeister , 1839 ) macrothemis heteronycha ( calvert , 1909 ) macrothemis hosanai santos , 1967 macrothemis idalia ris , 1919 macrothemis imitans karsch , 1890 - ivory-striped sylph macrothemis inacuta calvert , 1898 - straw-colored sylph macrothemis inequiunguis calvert , 1895 - jade-striped sylph macrothemis lauriana ris , 1913 macrothemis ludia belle , 1987 macrothemis lutea calvert , 1909 macrothemis marmorata hagen , 1868 macrothemis meurgeyi daigle , 2007 macrothemis mortoni ris , 1913 macrothemis musiva calvert , 1898 macrothemis newtoni costa , 1990 macrothemis nobilis rácenis , 1957 macrothemis pleurosticta ( burmeister , 1839 ) macrothemis polyneura ris , 1913 macrothemis proterva belle , 1987 macrothemis pseudimitans calvert , 1898 - white-tailed sylph macrothemis rochai navás , 1918 macrothemis rupicola rácenis , 1957 macrothemis taurepan de marmels , 2008 macrothemis tenuis hagen , 1868 macrothemis tessellata ( burmeister , 1839 ) macrothemis ultima gonzález , 1992 macrothemis valida ( navás , 1916 )

macrothemis

kento furui added an unknown common name in an unknown language to
macrothemis
.
straw - colored sylph macrothemis inacuta female eagle creek , near clifton , greenlee co . , arizona 25 august 2013
summary of the logistic model describing the probability of the focal male of macrothemis imitans ( n = 31 ) being the resident in relationship to the value difference between their traits and its non - focal pair .
probability of the focal male macrothemis imitans to be the resident in relation the residual muscle mass difference from its non - focal pair . overlapping points represent different samples with the same focal result and residual muscle mass difference . high quality figures are available online .
average proportion of time that the resident and substitute males of the dragonfly macrothemis imitans spent in each behavior during 10 min of behavioral observation . the proportion of time in which males remained resting is not shown . bars represent the standard error . high quality figures are available online .
the straw - colored sylph ( macrothemis inacuta ) occurs in mexico and southward into south america , but in the united states it has only been found in a few areas of south texas and arizona . in october , 2003 , bob behrstock discovered a single mature male straw - colored sylph at a location in edinburg , hidalgo co . , texas . in the days to follow , up to five of these dragons were found at the same location . the first two shots here , of a young male , were taken at this same spot also in october , 2003 , with a canon eos 10d and ef 70 - 200 mm f / 2 . 8 l is lens with a 2x extender and 550 ex flash .
the jade - striped sylph ( macrothemis inequiunguis ) is a seldom - seen tropical dragonfly which is actually fairly rare and quite local in texas , the only state in the u . s . where it is found . it ranges southward into central and northern south america where it is more common , however . the shot here , of a teneral female , was taken along the banks honey creek in the honey creek state natural area , comal co . , texas , in june , 2005 . this photo was a digital capture with a canon eos 1d mark ii and an ef 300mm f / 4 l is lens and 2x extender and 580ex flash . martin reid had found another jade - striped sylph at a nearby location a week or so earlier and he took john abbott and me back to the vicinity to look for others .
males of the dragonfly macrothemis imitans ( karsch ) ( odonata : libellulidae ) are found along creeks and stream banks , where they presumably roam in search for females among patches with macrophytes or , in most cases , among exposed rocks and sandbars ( personal observation ) . territorial fights and matings are frequently observed . despite the high occurrence of agonistic interactions , however , it is unknown what behaviors are used by rivals during the dispute and what characteristics determine male rhp . therefore , in this work we proposed to evaluate whether physical contact is a common property of the territorial disputes and to identify which morphological or physiological traits determine the winning chances of male m . imitans in disputes for territory ownership . if contests occur with physical contact , we hypothesize that features related to size will determine male rhp . alternatively , if males do not exhibit physical contact during the contests , we hypothesize that features that confer greater capacity to remain in flight will determine male rhp .
males of many animal species show intraspecific disputes for mating territories that range from displays without physical contact to physical fights with risk of injury . this variation motivated the proposition of different models that suggest possible rules used by rivals to decide the contest winner . to evaluate those models , it is necessary to identify how males behave during the fight and the individual attributes that determine their fighting ability ( resource holding potential ) . for this , males of the dragonfly macrothemis imitans ( karsch ) ( odonata : libellulidae ) were used to evaluate two hypotheses conditioned on the occurrence of physical contact during the fight : if the contests occur with physical contact , features related to size should determine male resource holding potential , and if males do not exhibit physical contact during the contests , features that confer greater endurance should determine resource holding potential . to assess these hypotheses , we collected males that had ownership of territories ( resident males ) and males that occupied the territory after we removed the resident males ( substitute males ) . after the capture , the resident and substitute males were transferred to the laboratory for measurements of wing area , dry weight , thoracic muscle mass , and fat content . the results showed that resident males do not differ in any measured trait from substitutes . because the fights occur with physical contact , it is intriguing that resident males do not possess higher fighting capacity than intruders . perhaps physical contact does not incur high costs during the fight , and other asymmetries , such as motivation associated with prior residency of the disputed territory , determine the contest winner .
amphibians & reptiles birds mammals dragonflies fishes plants world biomes bird wing & tail images library resources publications pacific nw moths ( external site ) bug guide ( external site ) a catalogue of butterflies of the united states and canada , j . pelham , 2012 usfws feather atlas an identification manual to the small mammals of british coumbia tags silphidae of washington state
this list , including english names originally proposed in 1978 and now revised several times , is offered as both a current north american check - list and a list of english names . the english names were generated by dennis r . paulson and sidney w . dunkle ( a few were already in use ) and approved , with some modification , by the membership of the dragonfly society of the americas . common names for species recently added to the list were coined by the common names committe and then the checklist committee of the dsa or by the original describer . the list is kept up to date with taxonomic changes , name changes , and species newly added to the fauna .
slater museum of natural history 1500 n . warner st . # 1088 tacoma , wa 98416 253 . 879 . 3356
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canal area , metropolitan park , 27th august 2012 . netta smith ' s photo . total length : 40 - 43 mm . hind wing : 29 - 33 mm . note the color of its eyes , a beautiful rare type of blue .
white - tailed sylph male held in the hand , metropolitan natural park panama city , 27th august 2012 . erland nielsen ' s photo . habitat : clear rivers and large streams with some current , usually rocky . in open or forested country . distribution : vagrant in arizona ( 2007 ) , ranges south to ecuador and brazil ( dragonflies and damselflies of the west , dennis paulson ) .
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urltoken
thank you for taking the time to provide feedback on the iucn red list of threatened species website , we are grateful for your input .
justification : extent of occurrence is 1 , 683 , 500 km² and the species is known from six locations although it is probably more widespread across poorly sampled amazon lowland forest .
occurs in brazil , peru and paraguay ( ris 1913 , d . r . paulson pers . comm . 2006 , r . w . garrison 2006 ) .
this species is known from six locations and thought to be found in unsampled lowland forest but there is no data available on population sizes .
lowland amazon forest species ; no information on breeding habitat available , adults have been found near rivers and lakes ( collection data : ris 1919 , d . r . paulson pers . comm . 2006 ) . larva unknown .
to make use of this information , please check the < terms of use > .
the next four shots of straw - colored sylph were taken in falcon state park , zapata co . , texas , in october , 2004 . most of the park is in starr co . , but the section where these photos were taken is in zapata co . these images were taken with a canon eos 1d mark ii and an ef 300mm f / 4 l is lens and 2x extender and canon flash . these next four shots all show females .
the female straw - colored sylph shown here was along the rio grande in hidalgo co . , texas , in september , 2008 . this shot was taken with a canon eos 1d mark iii and ef 300mm f / 4 l is lens and 2x extender and canon 580 flash .
the next 5 shots show an adult female jade - striped sylph . these images were taken in august , 2007 , in lockhart , caldwell co . , texas , with a canon eos 1d mark iii and ef 300mm f / 4 l is lens and 2x extender and 580 flash . these 5 shots were of a posed insect which had been netted . we ( john abbott , eric isley and i ) saw at least 4 different individuals , but they remained in flight during all of our observations , thus the posed images . we still hope to get some shots of a naturally perched one at some point . the discovery of this species at this particular location represents the eastern most record in texas and only the 5th county in texas , and the u . s . , where this species has been found .
persistence pays off . we made another trip to the caldwell co . location a few days later and after watching several males and females in flight for long periods , we finally were able to watch several males perch . curiously , sometimes they hung from a grass stem , such as in the shot on the immediate right , sometimes they perched on top of a blade of grass or a ragweed leaf , such as in the next 3 shots , and sometimes they perched at a 45 degree angle such as the 5th shot . note that the males shown here , at least two different individuals , lack the brown tipped wings of the female in previous shots .
the perched female shown here was also shot in august , 2007 , at the same spot as the shots just before . finally , after many trips , we got some shots of a naturally perched female .
the next 5 shots of jade - striped sylph were taken along candeleria creek , montell , uvalde co . , texas , in july , 2010 , with a canon eos 1d mark iv and an ef 300mm f / 4 l is lens and 2x extender and 580 flash . the first 2 shots show a very teneral male while the next 3 shots show an adult female .
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m . a . n . mourão 1 , 2 , a and p . e . c . peixoto 3 , b , *
copyright this is an open access paper . we use the creative commons attribution 3 . 0 license that permits unrestricted use , provided that the paper is properly attributed .
this is an open access article distributed under the terms of the creative commons attribution license ( urltoken ) , which permits unrestricted reuse , distribution , and reproduction in any medium , providedthe original work is properly cited .
in many animal species , males fight for ownership of territories for mating purposes ( andersson 1994 ) . the fighting behaviors used by males differ greatly among species ( arnott and el wood 2009 ) , varying from disputes without any contact between the rivals ( e . g . , alcock and bailey 1997 ) to interactions with physical contact , injuries , and risk of death ( e . g . , bean and cook 2001 ) . such variation is common even in species that fight for the possession of mating sites during flight , such as butterflies , dragonflies , and damselflies ( marden and waage 1990 , peixoto and benson 2011a , junior and peixoto 2013 ) .
many theoretical models aim to explain the rules used by males to decide the winner of a territorial contest ( enquist and leimar 1983 , mesterton - gibbons et al . 1996 , payne 1998 ) . despite differences among these models about the ways by which disputes are decided , all assume that the chance of winning depends on two parameters : ( 1 ) the individual’s fighting ability ( rhp , resource holding potential ; parker 1974 ) , which can be determined by physical ( e . g . , size and weight ) or physiological traits ( e . g . , fat , muscle , and immunocompetence ) ( see suhonen et al . 2008 ) ; and ( 2 ) the individual’s motivation to invest in the contest ( rv , resource value ; parker 1974 ) , which is often determined by the previous experience of the owner of the contested resource and by the local quality of the defended site ( e . g . , briffa and elwood 2001 , bergman et al . 2010 ) .
because all the models previously mentioned postulate that rhp differences between rivals determine the chance of winning , to be able to evaluate such models empirically it is necessary to identify which morphological or physiological traits determine the males’ most often used for this purpose is to establish correlations between specific male traits and the chances of winning a territorial contest . however , this approach is unsuitable in establishing the mechanism that connects the trait to how costs are accrued during the contest , and it can lead to the establishment of spurious relationships ( lailvaux and irschick 2006 ) .
to avoid these results , it is necessary to establish the functional relationship between male traits and fighting behavior ( irschick et al . 2007 ) . one way to establish this functional relationship is to classify the fighting behaviors according to the occurrence of physical contact between rivals ( e . g . , peixoto and benson 2011b ) . if the fights occur with physical contact , injuries should be the main form of cost accrual during the fight . in this sense , features such as body mass should increase the individual’s resistance to injuries from rivals or the ability to inflict damage . therefore , traits related to physical strength must be functionally important to determine male rhp ( e . g . , forslund 2000 ) . if fights occur without physical contact , however , the costs should be determined by the energy depletion during the fight . the winner must be the individual that is able to persist for a longer period in the contest . consequently , physiological features such as energetic reserves may be more important in determining the individual’s fighting ability ( e . g . , marden and waage 1990 ) .
in many dragonfly species , males establish mating territories and fight for the ownership of these sites ( corbet 1980 , córdoba - aguilar 2008 ) . the fighting behaviors usually comprise a combination of circular chase flights , upward spiral flights , and straight back - and - forth persecutions ( córdoba - aguilar 2008 ) . in some species , males may exhibit physical contact during disputes ( corbet 1999 , switzer and schultz 2000 , junior and peixoto 2013 ) , but in others , the males exhibit displays without contact ( waage 1988 , switzer 2004 ) . features such as energy reserves ( marden and rollins 1994 ) and parasitism ( contreras - garduño et al . 2006 ) seem to predominate as rhp determinants in this group .
the study was conducted in a 130 - m stretch of urubú stream ( 15°42 ' s , 47°51 ' w ) , located about 9 . 5 km from the center of brasilia , df , brazil . the average depth of the stream at that location was 60 cm , and the average width was 3 . 5 m . the weather is tropical - high altitude , with two distinct seasons : a wet and rainy summer , with an average temperature of 29 . 7°c ( sd = 0 . 72 ) and average rainfall of 203 . 9 mm ( sd = 27 . 2 ) , and a dry and cold winter with an average temperature of 12 . 5°c ( sd = 2 . 5 ) and average rainfall of 24 . 9 mm ( sd = 76 . 7 ) ( inmet 2011 ) .
to evaluate whether disputes occur with or without physical contact , samples were collected in campaigns composed of two consecutive days between november 2011 and march 2012 . on the first day of each campaign , several males were marked with a number written on one wing with a black permanent marker that allowed visual identification of males without recapture . on the second day , behavioral observations of males and the removal experiment were done to evaluate the traits associated with the chance of winning a contest ( see “removal experiments” below for details ) . behavioral observations were made on 31 resident males ( males that were defending the territories at the beginning of our observations ) and 31 substitute males ( males that occupied the territories after the resident males were experimentally removed ) only on sunny days from 09 : 00 to 14 : 00 , when individuals are more active . during the observations , the behaviors of individuals were recorded for 10 min using a mini - portable recorder , with particular reference to the occurrence of physical contact during territorial disputes exhibited by the observed males (
) . afterwards , the recordings were used to time how long each individual spent in each behavior .
after the behavioral observation , we performed a removal experiment to evaluate male traits that may determine the winning chances . for this experiment , whenever a male exhibited signals of territorial defense ( i . e . , fights with conspecific males or flight patrols ) , it was removed from the defended site ( resident male ) . after the male was removed , the territory remained under observation until a new male arrived ( substitute male ) . upon arriving on the observed site , the substitute male was captured to receive a mark , consisting of a point on one wing . this approach was done with only one male at a time , and it never took more than 1 min to avoid stress on the manipulated animal ( all marked males resumed the territorial defense after being released ) . after releasing the marked substitute male , it was observed for 10 min and then captured . after this , both males ( resident and substitute ) were measured for weight , wing area , fat content , and residual muscle mass ( for a detailed description , see “morphological and physiological measures” below ) .
removal experiments have been used successfully to evaluate male traits that are important for contest resolution ( peixoto and benson 2008 , 2011a ) . in these experiments , it is expected that resident males are individuals that have disputed the ownership of territories and won , whereas substitute males represent weaker individuals that were unable to obtain territories . thus , if fights occur with physical contact , it is expected that resident males have a greater wing area or greater body mass than the rival males . alternatively , if fights occur without physical contact , it is expected that resident males have a greater fat content or greater investment in thoracic muscles for flight than substitute males .
a caliper was used to measure the length and width of each wing . the wing length was measured as the distance between the wing insertion on the thorax and its apex . the width was recorded as the longest distance between the wing edges on the perpendicular direction to the length axis . wing area was estimated by calculating the ellipse area , considering the wing length as the largest diameter and width as the smallest diameter . then , the wing area of the four wings was added to estimate the total wing area for each male .
to estimate fat content , males were kept in an oven at ∽60°c for 48 hr . later , the thorax ( without wings and legs ) and abdomen were separately weighed on a balance ( accuracy 0 . 0001 g ) . the summed mass of thorax and abdomen after dehydration was used as a measure of male size ( dry mass ) . after weighing , both parts of each animal were submerged in a closed bottle containing 10 ml of chloroform for 48 hr to extract lipids . after removing them from the chloroform , the body parts were kept in an oven at 60°c for an additional 48 hr before being weighed again . to estimate the amount of thoracic muscle mass available for flight ( flight muscle ratio ) , the same procedure was used . however , only the thorax weight ( after lipid extraction ) was measured before and after being immersed in 0 . 2 m potassium hydroxide ( plaistow and siva - jothy 1999 ) . the mass difference before and after immersion in chloroform was used to estimate fat content , and the mass difference before and after the immersion in potassium hydroxide was used to estimate muscle mass . because larger individuals tend to have more fat and muscle mass than smaller ones , we calculated the residuals of a linear regression between the fat content and the dry body mass of individuals before the lipid extraction ( residual fat ) and between the muscle mass and dry body mass ( residual muscle mass ) to obtain values of fat content and muscle mass independent of size ( marden 1989 , marden and chai 1991 , marden and rollins 1994 ) .
a logistic regression was used to assess the effect of each trait ( wing area , dry weight , residual fat content , and residual muscle mass ) on the probability of a focal male to be territorial ( hosmer and lemeshow 1989 ) . because data were collected from pairs of males ( resident and substitute ) , we adopted a modification to maintain the pairing ( kemp 2000 ) . for this , 16 pairs were randomly chosen . for each pair , the resident individual was designated as the focal male ( focal state 1 ) . for the remaining 15 pairs , the substitute male was designated as focal individual ( focal state 0 ) . when the focal state was 1 , the values of resident male traits were subtracted from the values of its substitute pair . inversely , when the focal state was 0 , the values of the substitute male were subtracted from the values of its resident pair . thus , if resident males possess a higher fighting ability than substitutes , it was expected to find positive values for the differences between male traits associated with focal pairs whose state was 1 and negative values associated with focal pairs whose state was 0 ( e . g . , peixoto and benson 2008 ) . to identify the most parsimonious model describing the relationship between the focal status and differences in the male traits , we used a corrected version for small samples of the akaike information criterion ( burnham and anderson 2002 ) .
) . resident and substitute males stayed perched for the greatest proportion of the time ( resident males : mean = 519 . 2 s , sd = 41 . 1 ; substitute males : mean = 524 . 8 s , sd = 39 . 2 ) . when males were not perched , they spent most of their time in flight and flight patrol , respectively . although the 62 males ( resident and substitute combined ) had been involved in 180 disputes , the proportion of time spent in fights accounted for a smaller proportion of the time budget of males . on average , the fights lasted 10 . 1 sec ( sd = 5 . 3 ) for resident males and 8 . 09 sec ( sd = 3 . 88 ) for substitute males .
fights between males always initiated when a male entered the site defended by a resident or by a substitute male . in both situations , the territory owners flew toward the intruders , colliding against them . after the collision , the territory owner ( for both resident and substitute males ) performed straight flights and / or circular chase or collision against the intruder . after expelling the invader , the winner returned to the perch of origin . the resident males were involved in 99 disputes and won 100 % of the contests , and the substitute males were involved in 81 disputes and also won 100 % of the contests . we observed two matings performed by resident males and eight matings by substitute males ( corresponding to the smaller proportion of the time budget of males ) . no individual without a territory was observed mating .
the models are ordered according to their aicc values ( aicc represents the value of the akaike information criterion corrected for small samples ; äi is the value difference between the most parsimonious model and model i , wi is the weight of akaike model i ) .
territorial males of m . imitans ( residents and substitutes ) remained most of the time perched . the fact that the time budget of substitute males was similar to that of the residents indicates that the marking manipulation did not affect their behavior . because the fights accounted for a small part of their time budget , it is possible that the energy costs accrued during continuous flight in the contests are low . the fact that there are collisions between males during the fights , however , indicates that injuries may represent costs . in this sense , it was expected that features related to size should determine male rhp ( lailvaux and irschick 2006 ) . contrary to expectations , the chance of a male to be the resident individual was unrelated to its morphological or physiological traits . perhaps the removal of resident males a short time after the beginning of territorial activity has reduced their chance to be the individuals that won the fights , which could obscure eventual differences between winners and losers in determining residency . nevertheless , it is common among territorial insects that the resident males are the first to arrive in the territory , and they stay defending the same place on consecutive days ( e . g . , forsyth and montgomerie 1987 , peixoto and benson 2009 ) . the fact that the resident male won 100 % of the disputes indicates that it was the individual with greater winning chances . consequently , it is improbable that the resident males were wrongly identified during the removal experiment .
because resident and substitute males in ownership of the territories won all the fights , it is possible that the asymmetry in territorial status determines the winner ( davies 1978 ) . although there is a lack of clear empirical demonstrations , theoretical models suggest that males may determine the contest winner through uncorrelated asymmetries , such as the simple distinction between territory owners and intruders ( maynard smith and parker 1976 , kokko et al . 2006 ) . it is also possible that disputes are settled on the basis of differences in territory value for each rival ( parker and rubenstein 1981 , enquist and leimar 1987 ) . in many species , individuals in ownership of territories present greater investment in fights than intruder rivals ( switzer 2004 , takeuchi and honda 2009 , bergman et al . 2010 , peixoto and benson 2011b ) . consequently , it is possible that even individuals with lower fighting ability remain as residents because asymmetries in motivation associated with the value that each rival attributes to the territory .
the lack of correspondence between male traits that are important in determining residency and the occurrence of physical contact between rivals during the fights is intriguing . it is possible that injuries are , in fact , an important fighting cost ( e . g . , junior and peixoto 2013 ) . in this situation , there may be alternative individual traits that affect contest settlement . perhaps the fighting ability of males is not determined solely by a particular trait , but by a combination of different characteristics that determine the overall male condition ( kemp et al . 2006 ) . on the other hand , vieira and peixoto ( 2013 ) have demonstrated that the functional relationship between fighting behaviors and how individual traits affect cost accrual during the fight is not universal . in this sense , it is possible that collisions do not generate any fighting cost . if this is true , disputes should be mainly determined by asymmetries in male motivation associated with differences in resource value , as occur in some butterfly species ( takeuchi and honda 2009 , bergman et al . 2010 ) .
we thank the researchers from the department of ecology at the university of brasilia ( unb ) , rosana tidon , and regina macedo for the laboratory access for the physiological extractions of males of m . imitans and for the theoretical content related to sexual selection , respectively . we also thank lorenzo zanette ( federal university of ceará ) and márcio cardoso ( federal university of rio grande do norte ) for comments on the manuscript . m . a . mourão received a scholarship provided by funcap in the early months of work .
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