the blackburnian warbler ( setophaga fusca [ formerly dendroica fusca ] ) is a small new world warbler .

they breed in eastern north america , from southern canada , westwards to the southern canadian prairies , the great lakes region and new england , to north carolina .

blackburnian warbler

the blackburnian warbler was named after anna blackburne , an english botanist . no other north american warbler has an orange throat .
lawrence , l . 1953 . notes on the nesting behavior of the blackburnian warbler .
we only shot a blackburnian warbler , a yellow - winged ditto , and a few finches .
this blackburnian warbler ( dendroica fusca ) perched among apple blossoms for subscriber merv cormier of st john , new brunswick .
morse , d . 1994 . blackburnian warbler ( dendroica fusca ) . the birds of north america , 102 : 583 .
ebird range map - blackburnian warbler . ebird . cornell lab of ornithology , n . d . web . 20 july 2012 .
displayed are dsl landscape capability ( lc ) data for the blackburnian warbler for 2010 ( dsl current ) and the . . .
currently , blackburnian warbler populations are not considered threatened . however , wintering habitats are rapidly declining and they are sensitive to forest fragmentation .
blackburnian warbler ( dendroica fusca ) . the internet bird collection . lynx edicions , n . d . web . 20 july 2012 .
in form the song is much like that of the blackburnian warbler , but the loudness , different quality , and lower pitch distinguish it .
lawrence , l . 1953 . notes on the nesting behavior of the blackburnian warbler . the wilson bulletin , 65 : 135 - 144 .
habitat : blackburnian warbler breeds in tall coniferous and mixed forests , but at the other times , it’s found in a variety of woodlands and tall bushes .
the blackburnian warbler is a forest bird that breeds along the eastern seaboard from the maritime provinces to massachusetts and new york and west to alberta . . . .
— blakˈbərnēən noun or blackburnian warbler ( ˈ ) ̷ ̷ | ̷ ̷ ̷ ̷ ̷ ̷ ( s ) usage : often capitalized b etymology : mrs . hugh blackburn , 18th century englishwoman + english ian : a no . american warbler ( …
warbler of the united states ( { dendroica blackburni [ ae ] } ) . the male is
small warbler with a long body . note triangular ear patch , pale eyebrow , and white wingbars .
blackburnian warblers inhabit boreal forests , but during migration can occur in a variety of woodlands .
the blackburnian warbler has gray to black upperparts with long , whitish streaks , dark wings with two broad , white wing bars , a dark line through the eye , and a dark patch behind the eye .
the blackburnian warbler’s nest is a cup of bark fibers , twigs , and grasses , and is lined with moss , lichens , and other materials . it is placed high on an outer branch of a conifer .
same place as previous blackburnian recording . in dense willow stand immediately adjacent to the ms river .
behaviour : blackburnian warbler is an active warbler of the tree canopy , conspicuously flitting , hovering and flycatching in search of insects among the outer leaves . it tends to keep high up in conifers on the breeding grounds . it gleans insects on small branches . it also hawks insects , flying from a perch to grab them in mid air .
the blackburnian warbler is a small songbird with a large breeding range of 1 , 910 , 000 square kilometers . it breeds in coniferous and mixed forests in southeastern canada and the northeastern united states south through the appalachians , and winters in montane tropical forests in southern central america , northern south america , and the andes . the blackburnian warbler has an estimated breeding population of 10 , 000 , 000 , and a conservation rating of least concern .
range : blackburnian warbler breeds from saskatchewan to western newfoundland , southward to northern minnesota and massachusetts , and in mountain southward to northern georgia . it winters in southern central america and northern south america , southward in the andes .
blackburnian warblers inhabit deciduous and coniferous forests during the summer . they prefer coniferous forests , but will be found in spruce trees or hemlock in deciduous forests . in any forest type in their summer range , blackburnian warblers are most active foraging and vocalizing in conifers . tree species preferred by blackburnian warblers are pines
morse , d . 1967 . the contexts of songs in black - throated green and blackburnian warblers .
diet : blackburnian warbler is mostly insectivorous . it eats primarily caterpillars and beetles . it feeds on insects and spiders . it gleans insects on small branches high in tree . it will supplement its diet with fruit in the winter .
protection / threats / status : blackburnian warbler is an uncommon , but known , cowbird host . population’s numbers of this species seems to be holding steady , despite loss of preferred habitat in some areas of breeding and wintering range .
— rusvasis krūminukas statusas t sritis zoologija | vardynas atitikmenys : lot . dendroica fusca angl . blackburnian warbler vok . fichtenwaldsänger , m rus . еловый лесной певун , m pranc . paruline à gorge orangée , f ryšiai : platesnis terminas –… …
stephenson , t . and s . whittle ( 2013 ) . the warbler guide . princeton university press , new jersey , usa .
small warbler with a long body . adult males have a brilliant orange throat and face with a black crown and triangular ear patch .
morse , douglass h . 2004 . blackburnian warbler ( setophaga fusca ) , the birds of north america online ( a . poole , ed . ) . ithaca : cornell lab of ornithology ; retrieved from the birds of north america online : urltoken
zerda lerner , s . , d . stauffer . 1998 . habitat selection by blackburnian warblers wintering in colombia .
morse , douglass h . 2004 . blackburnian warbler ( setophaga fusca ) , version 2 . 0 . in the birds of north america ( p . g . rodewald , editor ) . cornell lab of ornithology , ithaca , new york , usa .
swainson’s name is commemorated in three species : the widespread western raptor , swainson’s hawk ; the southeastern swainson ‘s warbler and our swainson’s thrush .
doepker , r . v . , r . d . earle , and j . j . ozoga . 1992 . characteristics of blackburnian warbler , dendroica fusca , breeding habitat in upper michigan . can . field - nat . 106 : 366 - 371 .
nice , m . 1926 . behavior of blackburnian , myrtle , and black - throated blue warblers , with young .
the blackburnian warbler does not appear to be well adapted to exploiting the periodic massive outbreaks of the spruce budworm caterpillar that occur in the north woods . some studies have indicated that breeding populations may actually fall in outbreak areas , perhaps due to competition with the cape may warbler , which along with the bay - breasted is known to actively seek out and exploit these areas , resulting in large broods and high population densities .
nesting blackburnian warblers are very difficult to study , in part because their nests can be 80 feet up in a tree .
blackburnian warbler : breeds from alberta east to newfoundland , south to the great lakes , southern new england , and along the appalachian mountains to northern georgia . spends winters in the tropics of south america . preferred habitats include mixed forests of hemlock , spruce , and various hardwoods .
chipley , r . m . 1980 . nonbreeding ecology of the blackburnian warbler . pp . 309 - 317 in migrant birds in the neotropics ( a . keast and e . s . morton , eds . ) . smithson . inst . press , washington , d . c .
8 . morse , d . h . 2004 . blackburnian warbler ( setophaga fusca ) , . the birds of north america online ( a poole , ed ) ithaca : cornell lab of ornithology ; retrieved from the birds of north america online : urltoken doi : 102173 / bna102 .
female cerulean warblers have a unique bluish color to the upperparts . the whitish streaks on the back of blackburnian warblers are distinctive .
although the blackburnian warbler is morphologically similar to many of its congeners that share these forests , it differs strikingly from them in coloration . breeding males are characterized by blazing orange plumage over much of the anterior part of their body , a color not shared by other members of this genus .
chipley , r . m . 1980 .
nonbreeding ecology of the blackburnian warbler .
in migrant birds in the neotropics . , edited by a . keast and e . s . morton , 309 - 317 . washington , d . c : smithson . inst . press . close
voice : sounds by xeno - canto blackburnian warbler’s call is a rich “chip” . most frequent song type is high and thin , ending in an ascending buzzy trill “sip - sip - sip - sip - titi - tzeeee” . a variant type sound like “zillup - zillup - zillup - zizizizizi” .
one evening i went walking along a winding lakeside path when i turned a corner and there , at eye level , was the most beautiful bird i have ever seen : a little warbler , beautiful as warblers generally are , but this one with a vivid flame - orange throat and breast that glows in memory still . such was my introduction to the blackburnian warbler ( dendroica fusca ) , the smallest and perhaps the most beautiful of the “spruce warblers . ”
the blackburnian warbler is a brilliantly colored neotropical migrant , one of a large group of dendroica wood warblers that coexist during the breeding season in the northeastern coniferous forests of north america . birds of this group are so similar structurally that early students of bird foraging and niche partitioning wondered how they coexisted . they are now known to separate ecologically by foraging areas ( macarthur 1958 , morse 1968 ) , the blackburnian exploiting a treetop niche .
blackburnian warblers are more tolerant of mixed woodlands than are those two budworm specialists , breeding as far south as the smoky mountains , often in association with hemlocks . neither do blackburnian populations appear to rise and fall as much according to the fortunes of the budworm . indeed , thus far the numbers of this warbler appear remarkably stable , despite ongoing habitat destruction on the wintering grounds and widespread losses of our hemlocks to the introduced wooly adelgid insect pest .
morse , d . h . 1994 . blackburnian warbler ( dendroica fusca ) . in the birds of north america , no . 102 ( a . poole and f . gill , eds . ) . the academy of natural sciences , philadelphia , and the american ornithologists ' union , washington , d . c .
what made you want to look up blackburnian ? please tell us where you read or heard it ( including the quote , if possible ) .
blackburnian warblers are insectivorous , diurnal foragers , usually gleaning insects from leaves . they also hover to capture insects from the bottom of leaves . they feed on
morse , d . 1967 . the contexts of songs in black - throated green and blackburnian warblers . the wilson bulletin , 79 : 64 - 74 .
macarthur also recorded details of the warblers ' foraging habits and discovered that they differed too . for example , the cape may warbler hawks flying insects much more often than does the blackburnian and tends to move vertically rather than horizontally ( matching its tendency to remain on the outside of the tree ) . the black - throated green hovers much more than the bay - breasted , and the more variable yellow - rumped has the most varied feeding habits . in addition , macarthur found evidence that food shortage limited the size of the warbler populations .
blackburnian warblers live in the wild for a average of 3 to 6 years . the primary cause of death is not surviving through migration . because of poor weather or insufficient energy stores , death is more common during spring migration , especially when flying over the gulf of mexico . blackburnian warblers are not kept in captivity .
blackburnian warblers breed in southeastern canada , the northeastern u . s . , and the appalachians . they winter mostly in south america . the population appears stable .
blackburnian warbler , wilson’s storm - petrel , swainson’s thrush , lincoln’s sparrow . the common names of all of these birds , common in maine , are based on a person’s name . i’ll bet most of these people aren’t familiar to you . in today’s column , i will give you a little background on the people whose names are commemorated in the bird names .
morse , d . h . 1967 . the contexts of songs in the black - throated green and blackburnian warblers . wilson bull . 79 : 62 - 72 .
young , l . , m . betts , a . diamond . 2005 . do blackburnian warblers select mixed forests ? the importance of spatial resolution in defining habitat .
the most distinctive characteristics of the blackburnian warbler are its treetop ecology and the breeding male ' s color . one of a large genus of morphologically and ecologically similar warblers , the blackburnian is the only one with orange in its plumage . the male ' s striking black - and - orange pattern makes some observers think of a miniature oriole . blackburnian warblers are usually found high in trees , even during migration , and are not readily noticed in the dense foliage unless their high - pitched song announces their presence . at times they may be detected at the ends of branches , picking among leaves for bugs or caterpillars . its canopy nesting habits have defied many researchers ' efforts to study its breeding ecology .
displayed are dsl landscape capability ( lc ) data for the blackburnian warbler for 2010 ( dsl current ) and the future ( dsl 2080 ) ; higher values shown in dark red . lc incorporates habitat , climate , and prevalence to estimate suitable and accessible conditions for the species . lc values can ' t be compared across species . courtesy of k . mcgarigal , umass .
zerda lerner , s . , d . stauffer . 1998 . habitat selection by blackburnian warblers wintering in colombia . journal of field ornithology , 69 : 457 - 465 .
nice , m . 1926 . behavior of blackburnian , myrtle , and black - throated blue warblers , with young . the wilson bulletin , 38 : 82 - 83 .
( ) a beautiful warbler of the united states ( dendroica blackburniae ) . the male is strongly marked with orange , yellow , and black on the head and neck , and has an orange - yellow breast .
the wood - warblers occur throughout north america except for the far northern tundra . the many species of this family have evolved to fill a wide variety of niches including marshes ( yellowthroats ) to tree trunks ( the black - and - white warbler ) , and spruce forests ( the cape may warbler ) . several species can reside in the same area , yet avoid competition by occupying slightly different habitats or feeding in different ways .
curson , j . ( 2018 ) . blackburnian warbler ( setophaga fusca ) . in : del hoyo , j . , elliott , a . , sargatal , j . , christie , d . a . & de juana , e . ( eds . ) . handbook of the birds of the world alive . lynx edicions , barcelona . ( retrieved from urltoken on 9 july 2018 ) .
morse , d . h . 1967c . the contexts of songs in the black - throated green and blackburnian warblers . wilson bull . no . 79 : 62 - 72 . close
blackburnian warblers typically live in a 0 . 4 to 1 . 1 hectare area . however , they live on small islands if there are tall , mature trees and suitable food sources .
left to right : cape may , yellow - rumped , black - throated green , blackburnian , and bay - breasted warblers . black areas in stylized conifers show where feeding is concentrated .
blackburnian warblers are migratory , wintering in southern central america and in south america , and are very rare vagrants to western europe . these birds were named after anna blackburne , an english botanist .
the wilson whose name is commemorated in wilson’s storm - petrel , wilson’s plover , wilson’s phalarope , wilson’s warbler and wilson’s snipe is alexander wilson ( 1766 - 1813 ) . wilson played an important role in the development of north american ornithology .
the wood - warblers are known for their colorful plumages – the blackburnian warbler being one of the most striking members of this family with its deep orange - red throat that contrasts with its handsome black and white plumage . however while many species are known for their beautiful breeding plumage colors , they are also known to bird watchers as being extremely challenging to identify when in the fall they revert to their drab tan , olive , and pale colored plumages .
lerner , s . de la zerda and d . f . stauffer . 1998 . habitat selection by blackburnian warblers wintering in colombia . j . field ornithol . no . 69 : 457 - 465 . close
male blackburnian warblers are easily identified by their orange throat , breast and forehead . they have a black crown and cheeks and are noted for their wing bars and tail spots . blackburnian warblers have white bars on a black back , wings , and tail . there are black streaks on a white background on the sides and flanks . the lower breast and stomach is slightly orange , fading to white towards the tail .
description : blackburnian warbler adult male summer is unmistakable with its flaming orange throat and chest , and lighter orange crown centre , supercilium and sides of head . upperparts are almost blackish , and it has a huge white wing patch . in other plumages , the blackish ear - covert patch surrounded by yellow or orangey supercilium , sides of neck and throat , dark greyish upperparts , double white wing bars , pale yellowish underparts with streaked flanks , and very dark legs , are the distinctive features of this warbler . tail shows extensive white in outer feathers when spread . male is brightly coloured , female presents similar pattern , but duller . immature is similar to adult female . immature male with more yellow throat , and immature female much paler .
habitat fragmentation effects are also a concern as these forests are reduced in response to climate change , particularly upslope . range reduction is expected for species that inhabit montane spruce - fir forests at the southern edge of their range 10 . models of future distribution considering habitat and climate changes predicted that landscape capability ( suitable climate and habitat ) for the blackburnian warbler would decrease by 71 % of their 2010 northeastern range by 2080 2 , a loss of ~ 1 % / year .
in summer , male blackburnian warblers display dark gray backs and double white wing bars , with yellowish rumps and dark brown crowns . the underparts of these birds are white , and are tinged with yellow and streaked black . the head is strongly patterned in yellow and black , with a flaming - orange throat . it is the only north american warbler with this striking plumage . other plumages , including the fall male and adult female , are washed - out versions of the summer male , and in particular lack the bright colors and strong head pattern . basic plumages show weaker yellows and gray in place of black in the breeding male . blackburnian warblers ' songs are a simple series of high
the origin of the name “blackburnian” will be obscure to most . in their day , though , the sibling team of ashton and anna blackburn were noted british naturalists . in the years prior to and during our revolution , austin traveled in the american colonies , collecting specimens and sending them to anna at her estate in lancashire , england . no less than seventeen type specimens of our native birds were later described from their collection , so perhaps we can forebear the blackburns their warbler .
a fiery gem of the treetops . in the northern forest in summer , the male blackburnian warbler may perch on the topmost twig of a spruce , showing off the flaming orange of his throat as he sings his thin , wiry song . the female also stays high in the conifers , and the nest is usually built far above the ground . long - distance migrants , most blackburnians spend the winter in south america , where they are often common in mountain forest in the andes .
young , l . , m . betts , a . diamond . 2005 . do blackburnian warblers select mixed forests ? the importance of spatial resolution in defining habitat . forest ecology and management , 214 : 358 - 372 .
blackburnian warbler populations are stable . partners in flight estimates a global breeding population of 10 million birds with 28 % spending some part of the year in the u . s , 72 % in canada , and all of them migrating through mexico on the way south to their wintering grounds . they are a u . s . - canada stewardship species and rate a 9 out of 20 on the continental concern score . they are not on the 2014 state of the birds watch list . back to top
male blackburnian warblers attract their mates by singing in tree tops . when a potential mate comes close , males flick their tail and peck at the branch . males defend their mate from other males by flicking their tail , pecking at branches , and occasionally fighting . blackburnian warblers , like other songbirds , are socially monogamous . males typically help raise and feed the young . in some cases they have been absent , likely due to occasional extra - pair copulations .
blackburnian warblers are typically preyed on by larger birds of prey and some mammals . most predation is on eggs , hatchlings , and fledglings , or on adults as they watch and defend nests . hawks are the most common predators .
blackburnian warblers are strong indicators of mixed deciduous - coniferous forests and undisturbed , mature coniferous forests . surveyors use this species to study the ecology and effect of forest fragmentation by timber harvesting . they are also important for controlling spruce budworm (
— / blak berr nee euhn / a black and white north american wood warbler , dendroica fusca , having an orange throat and an orange and black head . [ 1775 85 , amer . ; named after mrs . hugh blackburn , 18th century englishwoman ; see ian ] * * * …
students of ecology may recall the classic study by macarthur ( 1958 ) wherein the foraging habits of five warblers that breed in our northern coniferous forests were compared . the study demonstrated resource partitioning , wherein the canopy was roughly divided up between species , each exhibiting different foraging preferences . along with the cape may warbler , the blackburnian foraged in the top - most branches of the canopy . between these two , cape mays more frequently went after aerial prey and foraged at the very end of branches , while blackburnians traversed the high foliage in a more methodical fashion .
reproduction : blackburnian warbler’s nest is built by female , well out from the trunk , on a horizontal branch where it’s concealed by foliage or lichen . it’s located from 3 to over 80 feet above the ground , but placement is extremely variable . nest is cup - shaped , densely constructed , hidden in dense vegetation . spider silk secures nest to site . sides consist of twigs , barks , plant fibres and rootlets . it’s lined with lichens , mosses , fine grasses , hair , dead pine needles and sometimes , such exotic substances as string cotton , horsehair and cattail down .
blackburnian warblers usually fly in mixed - species flocks . typically there are 1 to 2 blackburnian warblers per flock , but there can be as many as 7 . they are not very social with each other or other species other than foraging near each other . they have learned to co - exist with other species and avoid direct competition by staying in the upper third of the tree canopy and foraging and nesting on the outside of the branches , a form of niche partitioning . even in the same habitats and with similar food sources , different species of the
blackburnian warblers are , on average , 13 cm in length with a wingspan of 21 cm . like most warblers , they have small , flattened , and short bills , and have thin , black legs with 3 toes in front and one in back ( sibley , 2001 ) .
in most parts of its breeding range , the blackburnian warbler nests in mature coniferous and mixed forest and often occurs in highest densities in old - growth forests . coniferous trees , whether spruce , balsam fir , pine , or hemlock , are an essential habitat component throughout the species range . at the end of the breeding season , young and adults often leave the conifers to forage in food - rich paper birches . as with other warblers , it may use virtually any woody habitat , and occasionally even herbaceous habitats , during migration . in winter it is found in high - altitude forests , woodlands , plantations , and treed pastures .
the kirtland’s warbler is an endangered species restricted to a very specific type of habitat mostly found in michigan ; jack pine forests . its habitat is managed for this species in a few national forests by ensuring that there are jack pine stands of the age and composition this species requires . brown - headed cowbird populations are also controlled on its breeding grounds .
— warbler war bler , n . 1 . one who , or that which , warbles ; a singer ; a songster ; applied chiefly to birds . [ 1913 webster ] in lulling strains the feathered warblers woo . tickell . [ 1913 webster ] 2 . ( zo [ o ] l . ) any one of numerous species of small old… …
cape mays and blackburnians also winter in very different locales , the cape may winters primarily in the west indies whereas the blackburnian is among those few warblers which commonly travel clear to south america . wintering blackburnians were recently reported by some of our traveling local birders as far south as ecuador .
blackburnian warblers sing at dusk and dawn in the frequency range of 4 to 12 khz . they typically sing the same song at both times . males duet during these times and sing at each other after hostile encounters with other males . males are usually stationary , perched higher in tree than females when singing . singing is used to advertise for breeding and to advertise territories . they have two distinct song types , occasionally alternating them during breeding season . they flick their tail and peck at branches while singing during breeding season . blackburnian warblers sing one song while stationary at the top of the tree and during encounters with other males . they chirp twice frequently throughout aggressive encounters with other males . the other song is primarily used while foraging and while in the presence of a female . blackburnian warblers who do not find mates sing the second song type for long periods of time .
members of the parulidae are not colonial nesters but often occur in mixed flocks with other species after the breeding season . they forage in a variety of ways for invertebrates , small fruits , and nectar . while the waterthrushes forage on the ground in streams and wetlands , and the black - and - white warbler creeps along tree trunks , most wood - warblers glean the vegetation of trees and bushes and make short sallies for their insect prey .
the bachman’s warbler is an enigmatic species considered to be extinct by most authorities although slim hopes for its continued existence are kept alive by a few possible sightings over the last thirty years . historically occurring in the southeastern united states , this little known species is thought to have been dependent upon canebrakes on its breeding and wintering grounds in cuba . although the reasons for its decline are unknown , destruction of these canebrakes is the most likely reason for its demise .
macarthur found that each warbler species divided its time differently among various parts of the tree . the cape may , for instance , stayed mostly toward the outside on the top , the bay - breasted fed mostly around the middle interior , while the yellow - rumped moved from part to part more than either of the other two . this is shown in the accompanying diagrams , in which the zones that contained 50 percent of the birds ' feeding activity are blackened .
blackburnian warblers are known for their increased presence during spruce budworm ( choristoneura species ) breakouts , in which their survival and reproduction are greater because of the abundance of prey . these warblers are also well - known for interspecific niche partitioning . which is how they avoid direct competition with species of the same genus ; they coexist by using different parts of the same tree for nesting and foraging .
populations of this species are vulnerable owing to the loss of preferred winter forest habitat in northern south america , although breeding bird survey data suggest that their numbers remain stable . blackburnian warblers are forest - interior species , and their numbers decline in forest fragments . southerly populations breeding in eastern hemlock and fraser fir are at risk as a result of wooly adelgids responsible for heavy mortality of these trees .
some aspects of the blackburnian warbler ' s breeding biology are relatively well known , including foraging ( macarthur 1958 , morse 1968 ) , population dynamics ( morse 1976a ) , interspecific interactions ( morse 1976b ) , habitat selection ( morse 1976a ) , and singing behavior ( morse 1967 ) . some ecological studies have been performed on their wintering grounds in colombia ( chipley 1980 ; lerner and stauffer 1998 ) . recent studies have focused on the effect of silvicultural practices on this and other species ( e . g . , hagan et al . 1996 , meiklejorn and hughes 1999 , hobson and bayne 2000a , cumming and diamond 2002 ) . however , information on many aspects of the species ' life cycle is rudimentary . little is known of its migratory ecology . many aspects of its breeding ecology remain largely or completely unknown , in part a consequence of its treetop existence in northern forests .
blackburnian warblers are around 11 to 13 cm ( 4 . 3 to 5 . 1 in ) long , with a 20 to 22 cm ( 7 . 9 to 8 . 7 in ) wingspan , and weigh 8 to 13 g ( 0 . 28 to 0 . 46 oz ) . the average mass of an adult bird is 9 . 7 g ( 0 . 34 oz ) . among standard measurements , the
blackburnian warblers build a nest consisting of an open cup of twigs , bark , plant fibers , and rootlets held to branch with spider web and lined with lichens , moss , hair , and dead pine needles that ' s placed near the end of a branch . three to five whitish eggs are laid its nest which is usually placed 2–38 m ( 6 . 6–124 . 7 ft ) above the ground , on a horizontal branch .
blackburnian warblers are commonly found throughout southeast canada , from alberta into the great lake areas to newfoundland and into the appalachians as far south as georgia during the summer season . however , during migration , this species is seen throughout the eastern coast of north america and as far west as oklahoma . for the most part , they migrate over the gulf of mexico ; however , some fly over the eastern coast of texas . during the winter they can be found in costa rica , panama , and into peru .
blackburnian warblers spend , on average , 22 minutes on the nest and 5 minutes off the nest after the eggs hatch . both males and females feed the young , females more so than males . males feed more than one hatchling at a time and females typically only feed one young at a time . they feed the young every 10 to 20 minutes . they take 2 to 4 weeks until the young fledge , both parents care for the young during that time . it takes several months before the young are independent of the parents .
blackburnian warblers are solitary during winter and highly territorial on their breeding grounds and do not mix with other passerine species outside of the migratory period . however , during migration , they often join local mixed foraging flocks of species such as chickadees , kinglets and nuthatches . these birds are basically insectivorous , but will include berries in their diets in wintertime . they usually forage by searching for insects or spiders in treetops . the breeding habitats of these birds are mature coniferous woodlands or mixed woodlands , especially ones containing spruce and hemlocks . it typically winters in tropical montane forests .
f ive species of insectivorous wood warblers - - cape may , yellow - rumped , black - throated green , blackburnian , and bay - breasted - - were the subject of a classic study of community ecology ( the science of interpreting species interactions ) . these species often share the same breeding grounds in mature coniferous forests . they had been thought by some ornithologists to occupy the same niche - - in other words , they appeared to assume identical roles in the same bird community . these five warblers would thus be an exception to the ecological rule of competitive exclusion . the rule states that two species with essentially the same niche cannot coexist because one will always out - compete and displace the other .
climate impacts climate - induced habitat loss may impact blackburnian warblers . for example , the eastern hemlock and fraser fir that the birds often breed in at the southern portion of their range are threatened by wooly adelgids ( adelges tsugae ) , a tree pest that appears to be increasing due to climate change 9 . the trees have already nearly disappeared from the hemlock forests of new jersey and southern new england as well as high - elevation forests in the southern appalachians because of wooly adelgids 8 . climate change may also threaten the montane spruce - fir forests that the birds use . boreal forests are expected to become less common because of climate change - possibly becoming locally extinct in the northeastern u . s . under severe climate projections 10 .
for his doctoral dissertation , the late robert macarthur , who became one of the nation ' s leading ecologists , set out to determine whether the five species of warblers actually did occupy the same niche . by measuring distances down from the top and outward from the trunk of individual spruce , fir , and pine trees , macarthur divided the trees into zones and recorded feeding positions of the different warblers within each . a record in zone
t3
indicated a bird feeding among the abundant new needles and buds of the tip of a branch , between 20 and 30 feet from the top of the tree . a record of
m3
signified feeding mostly among dead needles at the same height but in the middle zone of a branch . a record of
b2
represented a warbler feeding on the bare , lichen - covered base of a branch . in all , 16 different positions were distinguished ,
html public
- / / w3c / / dtd xhtml 1 . 0 strict / / en
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 .
cramp , s . and simmons , k . e . l . ( eds ) . 1977 - 1994 . handbook of the birds of europe , the middle east and africa . the birds of the western palearctic . oxford university press , oxford .
justification : this species has an extremely large range , and hence does not approach the thresholds for vulnerable under the range size criterion ( extent of occurrence < 20 , 000 km2 combined with a declining or fluctuating range size , habitat extent / quality , or population size and a small number of locations or severe fragmentation ) . the population trend appears to be increasing , and hence the species does not approach the thresholds for vulnerable under the population trend criterion ( > 30 % decline over ten years or three generations ) . the population size is extremely large , and hence does not approach the thresholds for vulnerable under the population size criterion ( < 10 , 000 mature individuals with a continuing decline estimated to be > 10 % in ten years or three generations , or with a specified population structure ) . for these reasons the species is evaluated as least concern .
greenland ; grenada ; iceland ; united kingdom ; virgin islands , u . s .
to make use of this information , please check the < terms of use > .
open cup of twigs , bark , plant fibers , and rootlets held to branch with spider web . lined with lichens , moss , hair , and dead pine needles . placed near tip of branch of conifer .
dunn , j . l . and k . l . garrett . 1997 . a field guide to the warblers of north america . boston , ma : houghton mifflin co .
lutmerding , j . a . and a . s . love . longevity records of north american birds . version 2015 . 2 . patuxent wildlife research center , bird banding laboratory 2015 .
north american bird conservation initiative . 2014 . the state of the birds 2014 report . us department of interior , washington , dc , usa .
sauer , j . r . , j . e . hines , j . e . fallon , k . l . pardieck , jr . ziolkowski , d . j . and w . a . link . the north american breeding bird survey , results and analysis 1966 - 2013 ( version 1 . 30 . 15 ) . usgs patuxtent wildlife research center 2014b . available from urltoken
sibley , d . a . ( 2014 ) . the sibley guide to birds , second edition . alfred a . knopf , new york , usa .
females / immatures are paler and yellower overall with 2 white wingbars . note distinctive triangular ear patch and yellow eyebrow .
hops around in vegetation and nervously looks around . this video has no audio .
females / immatures lack the orange coloration , showing tones of yellow instead . note triangular ear patch , pale eyebrow , and white wingbars .
the brilliant orange throat and head pattern is distinctive on adult males although the intensity of orange varies among individuals .
females / immatures have a distinctive dull black triangular ear patch and bold wingbars . this video has no audio .
males often sing from the tops of trees and branches . this video has no audio .
: throat and upper breast deep orange . sides of neck , eyestripe , line on forecrown , and eye arc yellow - orange . face patch , crown , and back black . lower breast yellowish with black streaks on sides . belly white or yellowish . wings black with broad white wingbars that run together into a white patch . back black with two creamy white lines . tail black with large white patches in outer tail feathers .
: similar to breeding , but oranges less intense and more yellow . olive edges to black back feathers . wingbars more distinct and less of a continuous white patch .
: throat , upper breast , eyestripe , forecrown stripe , and sides of neck orange yellow . crown , face patch , and flank streaks greenish gray . back gray with dark and light streaking . two broad white wingbars .
: similar to breeding , but more olive - brown above and with less white in the wing .
immature similar to adult female . immature male with more yellow throat , some black in eyeline , and more yellow onto flanks . immature female much paler , with yellowish throat and eyeline , blurry streaking on sides , grayer face patch and crown , and narrower wingbars .
may be especially vulnerable to loss of wintering habitat , with cutting of forest at mid - levels in mountains in the tropics .
woodlands ; conifers in summer . breeds in boreal coniferous and mixed forests , especially spruce and hemlock . in southern part of breeding range in appalachians , can inhabit completely deciduous forests . when migrating , occurs in all kinds of trees and brush . during winter in the tropics , usually in humid mountain forest .
feeds mostly in treetops , searching along small branches and twigs . also hovers to take insects from undersides and tips of foliage . will search dead leaf clumps ; occasionally flies out to catch flying insects . in spruce forests , males tend to forage higher than females . in winter in the andes , forages in mixed flocks with various tropical birds .
4 , sometimes 3 - 5 . white to greenish white , with blotches of reddish brown concentrated near the larger end . only females incubate , probably 12 - 13 days . male feeds female during incubation . young : both parents feed nestlings . when the young leave the nest , the parents separate , each caring for part of the brood .
both parents feed nestlings . when the young leave the nest , the parents separate , each caring for part of the brood .
mostly insects , especially caterpillars . in summer , feeds on many caterpillars , particularly those of spruce budworm ; also eats beetles , ants , flies , and many other insects , also spiders . especially during winter , will take some berries as well .
details of nesting behavior not well known , partly because nests are high and hard to observe . male defends nesting territory by singing , sometimes by attacking intruding males . in courtship , male sings , and performs displays with gliding flight and fluttering wings and tail . nest : almost always placed in dense vegetation near tips of branches of conifers , and usually high , sometimes up to 80 ' above ground . nest ( probably built by female ) is cup - shaped and made of twigs , bark , and fibers ; lined with lichens , moss , grass , hair , and conifer needles .
from wintering areas ( mostly in andes of south america ) , many apparently move north through central america , then fly north across gulf of mexico . fall migration may be spread out over a broader front .
very thin and wiry , increasing in speed and rising to the limit of hearing , sleet - sleet - sleet - sleet - sleetee - sleeeee . also tiddly - tiddly - tiddly - tiddly at same speed and pitch .
audio © lang elliott , bob mcguire , kevin colver , martyn stewart and others .
in the broadest and most detailed study of its kind , audubon scientists have used hundreds of thousands of citizen - science observations and sophisticated climate models to predict how birds in the u . s . and canada will react to climate change .
the darker the color , the more favorable the climate conditions are for survival . the outlined areas represent approximate current range for each season .
each map is a visual guide to where a particular bird species may find the climate conditions it needs to survive in the future . we call this the bird’s “climatic range . ”
the darker the shaded area , the more likely it is the bird species will find suitable climate conditions to survive there .
the outline of the approximate current range for each season remains fixed in each frame , allowing you to compare how the range will expand , contract , or shift in the future .
the first frame of the animation shows where the bird can find a suitable climate today ( based on data from 2000 ) . the next three frames predict where this bird’s suitable climate may shift in the future—one frame each for 2020 , 2050 , and 2080 .
you can play or pause the animation with the orange button in the lower left , or select an individual frame to study by clicking on its year .
the darker the color , the more favorable the climate conditions are for survival . the outlined areas represent approximate current range for each season . more on reading these maps .
building collisions kill hundreds of millions of birds each year . senator booker introduced a bill to reduce bird deaths caused by federal buildings .
animal services was able to rescue just three birds after a massive number of spring migrants hit a glass - sided tower in galveston .
tell congress to oppose a harmful rider that threatens sage - grouse and other wildlife .
tell congress and the department of the interior to uphold the country ' s most important bird protection law .
html public
- / / w3c / / dtd html + rdfa 1 . 1 / / en
/ * > < ! - - * / . alpha - grid - debug . container - 12 { background - image : none ; } / *
an estimated 55 % of the species ' north american population breeds within the boreal forest .
this species is one of more than 30 birds selected for in - depth profiles . find out why and see all selected boreal birds »
5
( 13 cm ) . breeding male black and white with vivid orange throat , crown patch , and eyebrow ; and large white wing patch ; female similar , but has yellow throat . back of both sexes boldly striped . immature male similar to female .
4 brown - spotted white eggs in a twig nest lined with lichens , mosses , and hair , usually placed high in a large conifer .
blancher , p . 2003 . the importance of canada ' s boreal forest to landbirds . canadian boreal initiative .
macarthur , r . h . 1958 . population ecology of some warblers of northeastern coniferous forests . ecology 39 : 599 - 619 .
morse , d . h . 1970 . ecological aspects of some mixed - species foraging flocks of birds . ecol . monogr . 40 : 119 - 168 .
sauer , j . r . , j . e . hines , and j . fallon . 2004 . the north american breeding bird survey , results and analysis 1966 - 2003 . version 2004 . 1 . usgs patuxent wildlife research center , laurel , md .
birding content provided by national wildlife federation / enature , with support from ducks unlimited / the pew charitable trusts .
be the first to receive news & important conservation alerts about the boreal forest .
affinities within genus uncertain ; some suggestions that closest relative may be s . cerulea , based mainly on similarity of songs , but a molecular phylogeny recovered this species as sister to s . castanea . has hybridized with s . castanea and at least once with mniotilta varia , also possibly with s . pensylvanica and possibly with s . kirtlandii . monotypic .
s canada ( c saskatchewan , locally from alberta , e to s newfoundland and nova scotia ) s in ne usa to c wisconsin and new england , and in appalachians to extreme n georgia . migrates to central america and nw south america , s in andes to c peru and , less commonly , c bolivia .
13 cm ; 9·7 g . male breeding has crown and ear - coverts black , orange lower eye - crescent , fiery orange forecrown patch , supercilium , neck side , throat and upper breast ; . . .
two song types . type 1 song a series of thin , very high - pitched , whistled
swee
notes , . . .
feeds mainly on insects and other arthropods ; occasionally takes berries in winter . forages mainly by gleaning in canopy , also sometimes . . .
long - distance migrant . leaves breeding grounds mostly during aug , moving s chiefly through . . .
not globally threatened ( least concern ) . fairly common throughout range . breeding densities vary with habitat , with 0·7–1·1 pairs / ha in mature red spruce . . .
only subscribers are able to see the bibliography . login or subscribe to get access to a lot of extra features !
only members are able to post public comments . to make the most of all of hbw ' s features , discover our subscriptions now !
previously considered to include species now separated in peucedramidae , zeledoniidae and teretistridae , as well as two genera now in phaenicophilidae . the most comprehensive molecular phylogeny of present family to date # r recovered nine principal clades ; almost all genera correspond to well - supported units , while two confined to west indies are maintained based on other characters . several traditionally recognized genera prove to be paraphyletic or otherwise unsustainable : dendroica is abandoned because setophaga is nested within it and has priority ; ergaticus is embedded within the clade occupied by cardellina ; euthlypis is subsumed within basileuterus ; oporornis is restricted to one species while its former congeners are removed to geothlypis ; parula proves to be widely paraphyletic ( two species move to setophaga , the rest to a resurrected oreothlypis ) ; phaeothlypis is indistinguishable from myiothlypis ; and the three wilsonia are more correctly placed in cardellina ( two species ) and setophaga ( one ) .
previously restricted to a single species ( s . ruticilla ) ; but comprehensive molecular phylogeny reveals that this species is deeply embedded within large clade that includes two species formerly in parula ( s . americana and s . pitiayumi ) and another ( s . citrina ) traditionally in wilsonia , as well as all species previously treated in dendroica # r ; catharopeza might also be subsumed into this genus # r , but better maintained as a separate monospecific genus , in view of unusual morphology and behaviour , as well as its basal position in the clade . as type species of parula , wilsonia and dendroica are all included in present grouping , these names become synonyms of oldest available name for this clade , setophaga .
get access to the contents of the hbw including all species accounts , family texts , plates , audiovisual links , updates and related resources .
also available : 2 - year subscription package : 55 . 90 € ( instead of 59 . 90 € ) 3 - year subscription package : 82 € ( instead of 89 . 85 € )
supporting members help us to develop and update the project more quickly and to reach more people by keeping prices down .


parahepialus nebulosus is a species of moth of the family hepialidae .

it was described by alphéraky in 1889 , and is known from the tibet autonomous region in china .

parahepialus

parahepialus zou & zhang , 2010 is synonymized with thitarodes viette , 1968 . thitarodes quadrata jiang , li , li , li & han , sp . n . is described from sichuan , china . illustrations of the adults and the genitalia are given for the type species of parahepialus and the new species .
there are different characteristics for genus identification found in the genitalia morphology of the genus hepialus insects reported in china , and most species of hepialus described by chinese scientists were transferred to thitarodes by foreign scholars . the taxonomy of the genus hepialus currently adopted in china was revised and newly divided into four genera , parahepialus gen . nov . , ahamus gen . nov . , hepialus and thitarodes viette , 1968 on the basis of the particular structure of the valve on male genitalia . 62 species belonging to the original hepialus described in china were rearranged into the four genus , 1 species of parahepialus , 1 species of hepialus , 18 species of ahamus , 40 species of thitarodes . its rationality was supported by the systemic phylogenetic tree constructed by the cytb gene of some original hepialus species collected from genbank .
there are other species that share similar hosts as ophiocordyceps sinensis . for example , o . crassispora and c . kurijimeansis are recorded on thitarodes baimaensis and t . armoricanus , respectively ( table 2 ) . ophiocordyceps gracilis , o . lanpingensis , and o . laojunshanensis are recorded on ahamus altaicola , a . jianchuanensis , and a . yunnanensis , respectively ( table 2 ) . ophiocordyceps gracilis is also recorded on three more hosts hepialus humuli , korscheltellus lupulina , and parahepialus nebulosus . similarly , cordyceps hepialidicola is reported on endoclita excrescens and o . ramosissimum and o . xuefengensis on endoclita nodus ( table 2 ) .
ophiocordyceps sinensis has the widest host range recorded on hepialidae , covering 55 spp . [ 95 , 96 ] . among them , thitarodes and ahamus are the two major genera ( table 2 ) . thirty - two spp . of thitarodes ( t . albipictus , t . armoricanus , t . baimaensis , t . baqingensis , t . bibelteus , t . biruensis , t . callinivalis , t . cingulatus , t . damxungensis , t . deqinensis , t . dongyuensis , t . ferrugineus , t . gonggaensis , t . jialangensis , t . jinshaensis , t . kangdingensis , t . kangdingroides , t . latitegumenus , t . litangensis , t . markamensis , t . meiliensis , t . namensis , t . namlinensis , t . oblifurcus , t . pratensis , t . pui , t . renzhiensis , t . varians , t . xunhuaensis , t . yeriensis , t . zaliensis , and t . zhongzhiensis ) and 13 spp . of ahamus ( a . anomopterus , a . gangcaensis , a . jianchuanensis , a . lijiangensis , a . luquensis , a . maquensis , a . sichuanus , a . yulongensis , a . yunlongensis , a . yunnanensis , a . yushuensis , a . zadoiensis , and a . zhayuensis ) are the host species of ophiocordyceps sinensis ( table 2 ) . the other host species are pharmacis carna , p . fusconebulosa , p . pyrenaicus , magnificus jiuzhiensis , m . zhiduoensis , bipectilus yunnanensis , endoclita davidi , gazoryctra ganna , parahepialus nebulosus , and hepialus xiaojinensis ( table 2 ) .
for full functionality of researchgate it is necessary to enable javascript . here are the instructions how to enable javascript in your web browser .
shilap revista de lepidopterología , vol . 44 , núm . 175 , septiembre , 2016 , pp . 373 - 378
: 411 , pl . 25 , fig . 135 , by original designation .
chu & wang ( 2004 ) ( figs 1 , 4 ) . however , we found the figure
was used in chu & wang ( 2004 ) . we found that the male genitalia were broken during preparation ,
jiang , li , li , li & han , sp . n . (
brown ventrally . female ( fig . 3 ) . forewing length : 16 - 17 mm . essentially as described for male , except
( figs 8 - 12 ) : uncus short and acute , strongly sclerotized . tegumen
broad , rounded terminally . a quadrate lobe protruding from base of vinculum with a broad and rounded
of the zoological systematics and evolution of the chinese academy of sciences ( no . o529yx5105 ) .
dhendup , k . , cannon , p . , hywel - jones , n . & rai , t . b . , 2010 . –
liang , x . c . , yang , d . r . , chen , f . r . , long , y . c . & yang , y . x , 1988 . – four new species of the genus hepialus ( ghost moth ) from yunnan , china . – zoological research , 9 ( 4 ) : 419 - 425 .
ueda , k . , 1996 . – a new species of thitarodes viette ( lepidoptera , hepialidae ) from japan . – bulletin of the kitakyushu museum of the natural history , 15 : 35 - 41 .
viette , p . e . l . , 1968 . – contribution à l ' études des hepialidae ( 36ème note ) : lepidoptera hepialidae du népal . – khumbu himal , 3 : 128 - 133 .
one new species and one new record for the genus ninodes warren from china ( lepidoptera , geometridae . . .
a new species of the genus ninodes warren , n . quadratus sp . n . , is described from china and compared with related species , based on numerous museum specimens . n . albarius beljaev & park , 1998 , described from korea , is newly recorded for china . illustrations of external features and genitalia for each species of ninodes are presented .
the subfamily cyclidiinae from china is reviewed : two genera and seven species are reported from china . one new subspecies , cyclidia fractifasciata indistincta subsp . n . , is described . two new synonyms are established : cyclidia substigmaria ( hübner , 1831 ) ( = cyclidia substigmaria brunna chu & wang , 1987 , syn . n . = cyclidia tetraspota chu & wang , 1987 , syn . n . ) . one misidentification in chu & . . . [ show full abstract ]
allopatric divergence and secondary contact without genetic admixture for arichanna perimelaina ( lep . . .
heterophleps inusitata , an extremely rare new moth species from western yunnan , china ( lepidoptera , . . .
the genus synegiodes swinhoe , 1892 is reviewed and redescribed , s . sanguinaria ( moore , 1868 ) is formally fixed as type - species according to the provisions of the international code of zoological nomenclature . seven species , including s . brunnearia ( leech , 1897 ) , are presently recognized as members of this genus and two new species from china and n . vietnam , s . expansus sp . nov . and s . . . . [ show full abstract ]
= ; [ nhm card ] ; nielsen , robinson & wagner , 2000 , j . nat . hist . 34 : 847 ( list ) ; jiang , li , li , li & han , 2016 , shilap revta . lepid . 44 ( 175 ) : 374
= ; jiang , li , li , li & han , 2016 , shilap revta . lepid . 44 ( 175 ) : 374
ahamus zou & zhang , 2010 ; j . hun . univ . sci . tech . 25 ( 1 ) : 116 ; ts : hepialus jianchuanensis yang
thitarodes ( hepialidae ) ; [ nhm card ] ; nielsen , robinson & wagner , 2000 , j . nat . hist . 34 : 847 ( list ) ; jiang , li , li , li & han , 2016 , shilap revta . lepid . 44 ( 175 ) : 374
arizanus ( matsumura , 1931 ) ( hepialus ) ; 6000 illust . insects japan . - empire : 1022 ; tl : japan
thitarodes arizanus ; nielsen , robinson & wagner , 2000 , j . nat . hist . 34 : 847 ( list )
= ; [ nhm card ] ; nielsen , robinson & wagner , 2000 , j . nat . hist . 34 : 847 ( list )
thitarodes armoricanus ; [ nhm card ] ; nielsen , robinson & wagner , 2000 , j . nat . hist . 34 : 847 ( list ) ; zou , liu & zhang , 2010 , j . hun . univ . sci . tech . 25 ( 1 ) : 117
thitarodes damxungensis ; nielsen , robinson & wagner , 2000 , j . nat . hist . 34 : 847 ( list ) ; zou , liu & zhang , 2010 , j . hun . univ . sci . tech . 25 ( 1 ) : 118
hepialus albipictus yang , 1993 ; acta zootax . sinica 18 : 184 ; tl : yunnan , deqin co . , renzhi snow mtn ( 28°52 ' n , 99°14 ' e , 4600 - 4780m )
thitarodes albipictus ; nielsen , robinson & wagner , 2000 , j . nat . hist . 34 : 847 ( list ) ; zou , liu & zhang , 2010 , j . hun . univ . sci . tech . 25 ( 1 ) : 117
hepialus jinshaensis yang , 1993 ; acta zootax . sinica 18 : 185 ; tl : yunnan , deqin co . , baima , ( 28°34 ' n , 99°18 ' e , 4600m )
thitarodes jinshaensis ; nielsen , robinson & wagner , 2000 , j . nat . hist . 34 : 847 ( list ) ; zou , liu & zhang , 2010 , j . hun . univ . sci . tech . 25 ( 1 ) : 117
hepialus deqinensis liang , 1988 ; zool . res . 9 ( 4 ) : 419 , 424 ; tl : yunnan , deqin county , jiawu snow mtn
thitarodes deqingensis [ sic , recte deqinensis ? ] ; zou , liu & zhang , 2010 , j . hun . univ . sci . tech . 25 ( 1 ) : 118
hepialus litangensis liang , 1995 ; zool . res . 16 : 210 , 212 ; tl : litang , sichuan
thitarodes litangensis ; nielsen , robinson & wagner , 2000 , j . nat . hist . 34 : 847 ( list ) ; zou , liu & zhang , 2010 , j . hun . univ . sci . tech . 25 ( 1 ) : 118
hepialus baimaensis liang , 1988 ; zool . res . 9 ( 4 ) : 419 , 424 ; tl : yunnan , deqin county , baimae
thitarodes baimaensis ; [ nhm card ] ; nielsen , robinson & wagner , 2000 , j . nat . hist . 34 : 847 ( list ) ; zou , liu & zhang , 2010 , j . hun . univ . sci . tech . 25 ( 1 ) : 118
hepialus meiliensis liang , 1988 ; zool . res . 9 ( 4 ) : 420 , 425 ; tl : yunnan , deqin county , meili snow mtn
thitarodes meiliensis ; [ nhm card ] ; nielsen , robinson & wagner , 2000 , j . nat . hist . 34 : 847 ( list ) ; zou , liu & zhang , 2010 , j . hun . univ . sci . tech . 25 ( 1 ) : 118
hepialus callinivalis liang , 1995 ; zool . res . 16 : 209 , 212 ; tl : meili snow mtn , deqin county , yunnan
thitarodes callinivalis ; nielsen , robinson & wagner , 2000 , j . nat . hist . 34 : 847 ( list ) ; zou , liu & zhang , 2010 , j . hun . univ . sci . tech . 25 ( 1 ) : 118
hepialus jialangensis yang , 1994 ; zool . res . 15 ( 3 ) : 6 , 10 ; tl : xizang , zogang county , meili snow mtn ( 4000 - 4600m )
thitarodes jialangensis ; nielsen , robinson & wagner , 2000 , j . nat . hist . 34 : 847 ( list ) ; zou , liu & zhang , 2010 , j . hun . univ . sci . tech . 25 ( 1 ) : 118
hepialus xiaojinensis tu , ma & zhang , 2009 ; entomotaxonomia 31 : 123 , 126 ; tl : xiaojin co . ( 30°54 ' n , 102°18 ' e , 4300 - 4800m , sichuan
thitarodes richthofeni ; nielsen , robinson & wagner , 2000 , j . nat . hist . 34 : 847 ( list )
dierli viette , 1968 ; ergeb . forsch . nepal himalaya 3 : 132 ; tl : nepal
thitarodes dierli ; [ nhm card ] ; ueda , 2000 , tinea 16 ( suppl . 1 ) : 74 ; nielsen , robinson & wagner , 2000 , j . nat . hist . 34 : 847 ( list )
eberti viette , 1968 ; ergeb . forsch . nepal himalaya 3 : 130 ; tl : thodung , nepal
thitarodes eberti ; [ nhm card ] ; nielsen , robinson & wagner , 2000 , j . nat . hist . 34 : 847 ( list ) ; ueda , 2000 , tinea 16 ( suppl . 1 ) : 74
danieli viette , 1968 ; ergeb . forsch . nepal himalaya 3 : 128 ; tl : nepal
thitarodes danieli ; [ nhm card ] ; ueda , 2000 , tinea 16 ( suppl . 1 ) : 71 ; nielsen , robinson & wagner , 2000 , j . nat . hist . 34 : 847 ( list )
ahamus yushuensis ; zou , liu & zhang , 2010 , j . hun . univ . sci . tech . 25 ( 1 ) : 116
ahamus altaicola ; zou , liu & zhang , 2010 , j . hun . univ . sci . tech . 25 ( 1 ) : 116
ahamus zhayuensis ; zou , liu & zhang , 2010 , j . hun . univ . sci . tech . 25 ( 1 ) : 116
ahamus lijiangensis ; zou , liu & zhang , 2010 , j . hun . univ . sci . tech . 25 ( 1 ) : 116
hepialus jianchuanensis yang , 1994 ; zool . res . 15 ( 3 ) : 5 , 10 ; tl : yunnan , jianchan county , laojun mtn , 2900 - 3100m
ahamus jianchuanensis ; zou , liu & zhang , 2010 , j . hun . univ . sci . tech . 25 ( 1 ) : 116
hepialus anomopterus yang , 1994 ; zool . res . 15 ( 3 ) : 7 , 10 ; tl : yunnan , jianchan county , laojun mtn , ( 2800 - 3100m )
ahamus anomopterus ; zou , liu & zhang , 2010 , j . hun . univ . sci . tech . 25 ( 1 ) : 117
hepialus yunnanensis yang , li & shen , 1992 ; zool . res . 13 : 245 , 249 ; tl : yunnan , laojun mtn , 26°45 ' n , 99°51 ' e
ahamus yunnanensis ; zou , liu & zhang , 2010 , j . hun . univ . sci . tech . 25 ( 1 ) : 116
ahamus yunlongensis ; zou , liu & zhang , 2010 , j . hun . univ . sci . tech . 25 ( 1 ) : 116
hepialus yulongensis liang , 1988 ; zool . res . 9 ( 4 ) : 421 , 425 ; tl : yunnan , lijiang , yulong snow mtn
thitarodes yulongensis ; [ nhm card ] ; nielsen , robinson & wagner , 2000 , j . nat . hist . 34 : 847 ( list )
thitarodes quadrata jiang , li , li , li & han , 2016 ; shilap revta . lepid . 44 ( 175 ) : 374 ; tl : china , sichuan , xiaojin
thitarodes sichuanus ; [ nhm card ] ; nielsen , robinson & wagner , 2000 , j . nat . hist . 34 : 847 ( list )
? ahamus menyuanensis [ sic , recte menyuanicus ] ; zou , liu & zhang , 2010 , j . hun . univ . sci . tech . 25 ( 1 ) : 116
thitarodes xizangensis ; [ nhm card ] ; nielsen , robinson & wagner , 2000 , j . nat . hist . 34 : 847 ( list )
thitarodes kangdingensis ; [ nhm card ] ; nielsen , robinson & wagner , 2000 , j . nat . hist . 34 : 847 ( list ) ; zou , liu & zhang , 2010 , j . hun . univ . sci . tech . 25 ( 1 ) : 118
thitarodes oblifurcus ; [ nhm card ] ; nielsen , robinson & wagner , 2000 , j . nat . hist . 34 : 847 ( list ) ; zou , liu & zhang , 2010 , j . hun . univ . sci . tech . 25 ( 1 ) : 118
thitarodes baqingensis ; nielsen , robinson & wagner , 2000 , j . nat . hist . 34 : 847 ( list ) ; zou , liu & zhang , 2010 , j . hun . univ . sci . tech . 25 ( 1 ) : 118
hepialus ferrugineus li , yang & shen , 1993 ; acta ent . sinica 36 ( 4 ) : 495 , 496 ; tl : yunnan , baima snow mountain , 4200 - 4500m
thitarodes ferrugineus ; nielsen , robinson & wagner , 2000 , j . nat . hist . 34 : 847 ( list ) ; zou , liu & zhang , 2010 , j . hun . univ . sci . tech . 25 ( 1 ) : 118
hepialus gonggaensis fu & huang , 1991 ; acta ent . sinica 34 ( 3 ) : 362 ; tl : sichuan
thitarodes gonggaensis ; [ nhm card ] ; nielsen , robinson & wagner , 2000 , j . nat . hist . 34 : 847 ( list ) ; zou , liu & zhang , 2010 , j . hun . univ . sci . tech . 25 ( 1 ) : 118
thitarodes zhangmoensis ; [ nhm card ] ; nielsen , robinson & wagner , 2000 , j . nat . hist . 34 : 847 ( list ) ; zou , liu & zhang , 2010 , j . hun . univ . sci . tech . 25 ( 1 ) : 118
thitarodes kangdingroides ; [ nhm card ] ; nielsen , robinson & wagner , 2000 , j . nat . hist . 34 : 847 ( list ) ; zou , liu & zhang , 2010 , j . hun . univ . sci . tech . 25 ( 1 ) : 118
hepialus markamensis yang , li & shen , 1992 ; zool . res . 13 : 246 , 249 ; tl : xizang , markam county , nimasha snow mtn , 28°59 ' n 98°46 ' e
thitarodes markamensis ; nielsen , robinson & wagner , 2000 , j . nat . hist . 34 : 847 ( list ) ; zou , liu & zhang , 2010 , j . hun . univ . sci . tech . 25 ( 1 ) : 118
hepialus zaliensis yang , 1994 ; zool . res . 15 ( 3 ) : 7 , 10 ; tl : xizang , markam county , zhali snow mtn , 28°58 ' n 98°48 ' e , 4600 - 4900m
thitarodes zaliensis ; nielsen , robinson & wagner , 2000 , j . nat . hist . 34 : 847 ( list ) ; zou , liu & zhang , 2010 , j . hun . univ . sci . tech . 25 ( 1 ) : 118
= ; nielsen , robinson & wagner , 2000 , j . nat . hist . 34 : 847 ( list )
thitarodes malaisei ; nielsen , robinson & wagner , 2000 , j . nat . hist . 34 : 847 ( list )
sinarabesca ( bryk , 1942 ) ( hepialus ) ; ent . tidskr . 63 : 153 ; tl : kansu
thitarodes sinarabesca ; nielsen , robinson & wagner , 2000 , j . nat . hist . 34 : 847 ( list )
hepialus nebulosus alphéraky , 1889 ; in romanoff , mém . lép . 5 : 85 ; tl : ne . tibet
thitarodes nebulosus ; nielsen , robinson & wagner , 2000 , j . nat . hist . 34 : 848 ( list )
thitarodes varius ; nielsen , robinson & wagner , 2000 , j . nat . hist . 34 : 848 ( list )
thitarodes varians ; nielsen , robinson & wagner , 2000 , j . nat . hist . 34 : 848 ( list )
hepialus variabilis bremer , 1861 ; bull . acad . imp . sci . st . petersb . 3 : 478
thitarodes variabilis ; [ nhm card ] ; nielsen , robinson & wagner , 2000 , j . nat . hist . 34 : 848 ( list )
thitarodes nipponensis ueda , 1996 ; bull . kitakyushu mus . nat . hist . 15 : 35 ; tl : japan , shirakawadani , izumimura , kumanoto pref .
thitarodes nipponensis ; nielsen , robinson & wagner , 2000 , j . nat . hist . 34 : 848 ( list )
thitarodes luteus ; nielsen , robinson & wagner , 2000 , j . nat . hist . 34 : 848 ( list )
hepialus renzhiensis yang , shen , yang , liang , dong , chun , lu & sinaduji , 1991 ; acta ent . sinica 34 ( 2 ) : 218 , 224 ; tl : yunnan
thitarodes renzhiensis ; [ nhm card ] ; nielsen , robinson & wagner , 2000 , j . nat . hist . 34 : 848 ( list ) ; zou , liu & zhang , 2010 , j . hun . univ . sci . tech . 25 ( 1 ) : 118
hepialus zhongzhiensis liang , 1995 ; zool . res . 16 : 207 , 211 ; tl : renzhi snow mtn , deqin county , yunnan
thitarodes zhongzhiensis ; nielsen , robinson & wagner , 2000 , j . nat . hist . 34 : 848 ( list ) ; zou , liu & zhang , 2010 , j . hun . univ . sci . tech . 25 ( 1 ) : 118
hepialus yeriensis liang , 1995 ; zool . res . 16 : 207 , 211 ; tl : yeri snow mtn , deqin county , yunnan
thitarodes yeriensis ; nielsen , robinson & wagner , 2000 , j . nat . hist . 34 : 848 ( list ) ; zou , liu & zhang , 2010 , j . hun . univ . sci . tech . 25 ( 1 ) : 118
hepialus pratensis yang , li & shen , 1992 ; zool . res . 13 : 247 , 250 ; tl : yunnan , deqin county , baima snow mtn , 28°23 ' n 99°01 ' e
thitarodes pratensis ; nielsen , robinson & wagner , 2000 , j . nat . hist . 34 : 848 ( list ) ; zou , liu & zhang , 2010 , j . hun . univ . sci . tech . 25 ( 1 ) : 118
hepialus cingulatus yang & zhang , 1995 ; acta ent . sinica 38 ( 3 ) : 360 , 362 ; tl : gansu , wenxian county , 3200m
thitarodes cingulatus ; nielsen , robinson & wagner , 2000 , j . nat . hist . 34 : 848 ( list ) ; zou , liu & zhang , 2010 , j . hun . univ . sci . tech . 25 ( 1 ) : 118
hepialus luquensis yang , yang & zhang , 1995 ; acta ent . sinica 38 ( 3 ) : 360 , 362 ; tl : gansu , luqu county , 34°13 ' n , 102°24 ' e , 4276m
ahamus luquensis ; zou , liu & zhang , 2010 , j . hun . univ . sci . tech . 25 ( 1 ) : 117
hepialus xunhuaensis yang & yang , 1995 ; acta ent . sinica 38 ( 3 ) : 359 , 362 ; tl : qinghai , xunhua county , ( 35°38 ' n , 102°42 ' e ) , 3800m
thitarodes xunhuaensis ; nielsen , robinson & wagner , 2000 , j . nat . hist . 34 : 848 ( list ) ; zou , liu & zhang , 2010 , j . hun . univ . sci . tech . 25 ( 1 ) : 118
hepialus bibelteus shen & zhou , 1997 ; acta ent . sinica 40 ( 2 ) : 198 , 200 ; tl : meidu ( 28°22 ' n , 90°01 ' e ) , baima snow mountain , deqing county , 4500m , yunnan
thitarodes bibelteus ; zou , liu & zhang , 2010 , j . hun . univ . sci . tech . 25 ( 1 ) : 118
hepialus biruensis fu , 2002 ; acta ent . sinica 45 ( suppl . ) : 56 ; tl : xizang , biru county , 4400 - 4700m
hepialus biruens [ sic , recte biruensis ] ; zou , liu & zhang , 2010 , j . hun . univ . sci . tech . 25 ( 1 ) : 118
thitarodes sejilaensis zou , liu & zhang , 2011 ; pan - pacific ent . 87 ( 2 ) : 107 ; tl : mt sejila , linzhi county , tibet , 4500m , 29°36 ' n , 94°35 ' e
thitarodes harutai ueda , 2000 ; tinea 16 ( suppl . 1 ) : 85 ; tl : nepal , mechi , laam pokhari , 2850m
thitarodes jiachaensis zou , liu & zhang , 2011 ; pan - pacific ent . 87 ( 2 ) : 110 ; tl : jiancha county , tibet , 535m , 29°26 . 714 ' n , 94°42 . 888 ' e
thitarodes kingdonwardi ueda , 2000 ; tinea 16 ( suppl . 1 ) : 84 ; tl : se . tibet , tsangpo valley , nyima la ( 14000ft )
thitarodes kishidai ueda , 2000 ; tinea 16 ( suppl . 1 ) : 81 ; tl : nepal , lete nr nilgiri ( 2400m )
thitarodes limbui ueda , 2000 ; tinea 16 ( suppl . 1 ) : 85 ; tl : nepal , mechi , khambachen ( 3950m )
thitarodes maculatum ueda , 2000 ; tinea 16 ( suppl . 1 ) : 72 ; tl : nepal , chungbu khola ( 14500ft )
thitarodes namnai maczey , dhendup , cannon , hywl - jones & rai , 2010 ; zootaxa 2412 : 43 ; tl : bhutan , namna , n 27°44 ' 02 . 3
e 89°23 ' 32 . 2
, 4750m
thitarodes caligophilus maczey , dhendup , cannon , hywl - jones & rai , 2010 ; zootaxa 2412 : 47 ; tl : bhutan , namna , n 27°44 ' 02 . 3
e 89°23 ' 32 . 2
, 4750m
thitarodes hainanensis ; zou , liu & zhang , 2010 , j . hun . univ . sci . tech . 25 ( 1 ) : 118
thitarodes namensis ; zou , liu & zhang , 2010 , j . hun . univ . sci . tech . 25 ( 1 ) : 118
thitarodes xigazeensis ; zou , liu & zhang , 2010 , j . hun . univ . sci . tech . 25 ( 1 ) : 118
thitarodes yongshengensis ; zou , liu & zhang , 2010 , j . hun . univ . sci . tech . 25 ( 1 ) : 118
thitarodes dinggyeensis ; zou , liu & zhang , 2010 , j . hun . univ . sci . tech . 25 ( 1 ) : 118
thitarodes nanmlinensis ; zou , liu & zhang , 2010 , j . hun . univ . sci . tech . 25 ( 1 ) : 118
thitarodes pui ; zou , liu & zhang , 2010 , j . hun . univ . sci . tech . 25 ( 1 ) : 118
ahamus zadoiensis ; zou , liu & zhang , 2010 , j . hun . univ . sci . tech . 25 ( 1 ) : 116
ahamus gangcaensis ; zou , liu & zhang , 2010 , j . hun . univ . sci . tech . 25 ( 1 ) : 116
ahamus maquensis ; zou , liu & zhang , 2010 , j . hun . univ . sci . tech . 25 ( 1 ) : 116
thitarodes yadongensis ; zou , liu & zhang , 2010 , j . hun . univ . sci . tech . 25 ( 1 ) : 118
dongyuensis ( yang et al . , 1996 ) ( hepialus ) ; ( nom . nud . )
dongyuensis ; nielsen , robinson & wagner , 2000 , j . nat . hist . 34 : 848 ( nom . nud . )
[ maps ] warning ! the maps are automatically generated from the textual information , and the process does not always produce acceptable result ; see about maps for more info .
lépidoptères rapportés du thibet par le général n . m . przewalsky de son voyage de 1884 - 1885 in romanoff ,
neue schmetterlinge aus den reichsmuseum in stockholm . nachtrag zur lepidopteren - ausbeute der sven hedinschen expedition ( 1927 - 1930 )
entomological results from the swedish expedition 1934 to burma and british india . lepidoptera : fam . notodontidae stephens , cossidae newman und hepialidae stephens gesammelt von rené malaise
zou , liu & zhang , 2011 two new species of thitarodes ( lepidoptera : hepialidae ) from tibet in china pan - pacific ent . 87 ( 2 ) : 106 - 113
if you have corrections , comments or information to add into these pages , just send mail to markku savela keep in mind that the taxonomic information is copied from various sources , and may include many inaccuracies . expert help is welcome .
html public
- / / w3c / / dtd xhtml 1 . 0 transitional / / en
urltoken
cajviewer7 . 0 supports all the cnki file formats ; adobereader only supports the pdf format .
yang darong ; yang yuexiong ( kunming institute of zoology , academia sinica kunming 650107 ) zhang sanyuan ( institute of diversified economy of lunnan district . gansu province wudu 746000 ) ; three new species of the genus hepialus from qinghai and gansu , china ( lepidoptera : hepialidae ) [ j ] ; acta entomologica sinica ; 1995 - 03
jiang shuai - shuai * , zou zhi - wen * , liu xin , zhang gu - ren * * ( state key laboratory for biological control of sun yat - sen university , guangzhou 510275 , china ) ; morphology and antennal sensilla of meteorus sp . , a parasitoid of hepialus pui larvae [ j ] ; journal of environmental entomology ; 2009 - 03
liu xi - jin guo ying - lan ( institute of microbiology , academia sinica , beijing ) yu yong - xin ( zhejiang institute of chinese materia medico , hangzhou ) zeng wei ( sichuan institute of chinese materia medica , chongqing ) ; isolation and identification of the anamorphic state of cordyceps sinensis ( berk . ) sacc . [ j ] ; mycosystema ; 1989 - 01
zhang - wu ; chen yong - jiu ; shen fa - rong ; yang yue - xiong ; yang da - rong ; zhang ya - ping ( laboratory of cellular and molecular evolution , kunming institute of apology , cab kunming 650223 ) ( laboratory of insect resources , kunming institute of zoology . cas kunming 6 ; study of genetic divergence in cordyceps sinensis and c . crassispora from northwest of yunnan by using rapd [ j ] ; mycosystema ; 1999 - 02
kang ji - chuan , liang zong - qi , liu ai - ying , richard y . c . kong ( department of plant pathology , university of stellenbosch , p . bag x1 , matieland 7602 , south africa ) ( laboratory of fungus resources , guizhou university , guiyang 550025 ) ( department of biology ; molecular evidence of polymorphism in cordyceps based on 5 . 8s rdna and its2 sequences [ j ] ; mycosystema ; 2000 - 04
li zeng - zhi ; huang bo ; fan met - zhen ( college of forest utilizatin , anhui agricultural university , hefei , anhui 230036 ) ; molecular evidence for anamorph determination ofcordyceps sinensis ( berk . ) sacc . i . relation between hirsutella sinensis and c . sinensis [ j ] ; mycosystema ; 2000 - 01
dai jin - xia ~ ( 1 * * ) , zheng zhe - min ~ 2 ( 1 . college of life science , ningxia university , yinchuan 750021 , china ; 2 . college of life science , shaanxi normal university , xi ' an 710062 , china ) ; phylogenetic relationships of eleven species of pentatominae based on sequences of cytochrome b gene [ j ] ; entomological knowledge ; 2005 - 04
©2006 tsinghua tongfang knowledge network technology co . , ltd . ( beijing ) ( ttkn ) all rights reserved
ning ye , shuquan rao , tingfu du , huiling hu , . . . qi xu
nozomi mihara , tadashige chiba , kosuke yamaguchi , haruka sudo , . . . kazushi imai
global transcriptome analysis of halolamina sp . to decipher the salt tolerance in extremely halophilic archaea
copyright © 2018 elsevier b . v . or its licensors or contributors . sciencedirect ® is a registered trademark of elsevier b . v .
1 institute for bio - medical convergence , international st . mary’s hospital and college of medicine , catholic kwandong university , incheon 404 - 834 , republic of korea 2 environmental science , ishikawa prefectural university , 1 - 308 suematsu , nonoichi , ishikawa 921 - 8836 , japan 3 mushroom research division , national institute of horticultural and herbal science , rural development administration , eumseong 369 - 873 , republic of korea 4 forest biodiversity division , korea national arboretum , pocheon 487 - 820 , republic of korea 5 college of pharmacy , chung - ang university , seoul 156 - 756 , republic of korea
copyright © 2016 bhushan shrestha et al . this is an open access article distributed under the creative commons attribution license , which permits unrestricted use , distribution , and reproduction in any medium , provided the original work is properly cited .
cordyceps fr . is one of the hypocrealean genera , comprising more than 400 spp . that parasitize a wide range of insects and few fungal genera . externally , they produce clavate , cylindrical or thin filamentous , filiform stroma on the hosts . the history of exploring and identifying cordyceps fungi is long and stretches back to the 17th century [ 8 ] . one of the most popular fungi is ophiocordyceps sinensis ( syn . cordyceps sinensis ) that grows on hepialid larvae in the alpine grasslands of the himalayas and the tibetan plateau [ 9 ] . this host - fungus complex is traditionally highly revered for enhancing immunity , protecting lungs , enriching kidneys , restorative and tonic effects , treating impotence , and so forth [ 10 – 13 ] .
until the phylogenetic classification by sung et al . [ 14 ] , cordyceps remained a big genus placed in family clavicipitaceae of order hypocreales . however , phylogenetic studies showed that neither cordyceps nor clavicipitaceae was monophyletic [ 14 – 18 ] and hence cordyceps was segregated into several phylogenetic genera within three families of hypocreales [ 14 ] . according to the phylogenetic classification , cordyceps is now restricted to the clade containing the type species c . militaris , circumscribed to cordycipitaceae [ 14 ] . newly segregated genera ophiocordyceps and elaphocordyceps were placed under another family ophiocordycipitaceae ; the other two genera metacordyceps and tyrannicordyceps remained in clavicipitaceae sensu stricto [ 14 , 19 ] .
following the recent revision in the international code of nomenclature for algae , fungi , and plants ( icn ) , elaphocordyceps is now synonymized with tolypocladium [ 20 ] and metacordyceps with metarhizium [ 21 ] . drechmeria , podocrella , polycephalomyces , and sphaerocordyceps are other genera that consist of members that were formerly described under cordyceps . cordyceps along with the newly segregated genera is linked to many asexual insect - pathogenic genera distributed in all three families clavicipitaceae s . s . , cordycipitaceae , and ophiocordycipitaceae [ 4 , 14 , 20 ] . further synonymizations of genera are recently proposed within ophiocordycipitaceae following icn [ 22 ] .
the host range of cordyceps in classical sense is very broad and includes several orders ( coleoptera , lepidoptera , hymenoptera , hemiptera , orthoptera , araneae , diptera , blattodea , mantodea , dermaptera , odonata , phasmatodea , etc . ) . based on the published literature , nearly 60 % of the species of cordyceps sensu lato are recorded on two orders coleoptera and lepidoptera . the other major host orders are hymenoptera , hemiptera , orthoptera , and araneae . the majority of hosts ( more than 95 % ) in lepidoptera ( moths , butterflies ) and coleoptera ( beetle ) are larvae ; the rest are adults or pupae , making the host identification difficult . in contrast , majority of hosts in other orders are adults such as spiders ( araneae ) , fly ( diptera ) , ant , bee , and wasp ( hymenoptera ) , grasshopper , locust , and cricket ( orthoptera ) , cicada , bug , scale - insect , and coccid ( hemiptera ) , cockroach and termite ( blattodea ) , mantis ( mantodea ) , earwig ( dermaptera ) , dragonfly ( odonata ) , and stick - insect ( phasmatodea ) . why lepidopterans and coleopterans are mostly susceptible at larval stage while other orders are more susceptible at adult stage is not well understood .
cordyceps fungi range in habitat from aerial to epigeal to subterranean ones [ 74 ] , based on their respective host species . when the host stage is adult , they grow on exposed environments such as leaf litter or are attached to some plant parts such as leaf and branch and in contrast to the larval or pupal stages that are buried in soil ( hypogeal ) or on wood up to nearly 50 cm , for example , o . longissima [ 75 ] and o . xuefengensis [ 76 ] . ophiocordyceps caloceroides that is buried in the soil more than 50 cm is exceptional which grows on adult spiders [ 77 ] .
the pathogenic fungi sometimes produce parasite manipulated behaviors on hosts , such as death grip or biting behavior [ 78 – 80 ] . some of the parasitized ants always bite lower side veins but never the laminar blade or the upper surface [ 79 , 80 ] . similarly , some species of ants are attached to lower side of twigs whereas others are found attached to leaves [ 81 ] . in other cases , aphids , ants , grasshoppers , planthoppers , flies , and so forth shift to elevated position at the time of dying , known as summit disease [ 82 , 83 ] .
in this review , we present brief introductions of coleoptera and lepidoptera and their major subgroups parasitized by cordyceps fungi . we hope the present review will serve as a reference for researchers and scholars to get a quick look at coleopteran and lepidopteran hosts of cordyceps spp .
we accumulated host information from published literature and analysed the hosts that belonged to coleoptera and lepidoptera . we found that hosts of only 60 % of cordyceps fungi are known at family or genus / species ranks ; the rest are known only at the order rank and hence no further discussions of the hosts could be done for them . hence , in this review , we dealt with only those hosts that are known at family or infrafamily ranks within coleoptera and lepidoptera and tried to analyse the major taxonomic subgroups that are associated with cordyceps fungi . we also analysed the host stage among coleopteran and lepidopteran hosts based on published literature .
coleoptera is currently the most species - rich group on this planet [ 84 ] . the order is classified into four suborders ( polyphaga , adephaga , myxophaga , and archostemata ) [ 85 ] . polyphaga is the largest suborder covering 90 % of total beetle species and is classified into more than 170 families . adephaga is the second largest suborder , followed by myxophaga and archostemata . all the known coleopteran hosts belong to suborders polyphaga and adephaga under 8 superfamilies and 11 families . short descriptions of host families are given below , followed by cordyceps spp . recorded on them .
this is one of the largest families in coleoptera and its members are commonly called scarabs or scarab beetles . scarabaeid larvae ( commonly called grubs ) are short , thick , pale yellow or white , mostly live underground or under debris , and feed on dead organic matter ( scavengers ) . twenty - seven cordyceps spp . are recorded in this family ( table 1 ) . among them , c . brittlebankii is recorded on heteronyx sp . , c . brongniartii on anomala cuprea , c . coxii on lepidiota sp . , c . pseudoinsignis and c . velutipes on melolontha sp . , ophiocordyceps aphodii on aphodius howitti and a . tasmaniae , o . melolonthae on ancyloncha puncticollis , lachnosterna fusca , and melolontha sp . , o . michiganensis on scarabaeus sp . , o . ravenelii on l . fusca , phyllophaga sp . , and rhizotrogus sp . , and o . stylophora on costelytra zealandica ( table 1 ) . other species recorded in this family are c . obliquiordinata , c . scarabaeicola , metarhizium brittlebankisoides , ophiocordyceps arbuscula , o . barnesii , o . elongatiperitheciata , o . geniculata , o . gracillima , o . highlandensis , o . konnoana , o . larvicola , o . macularis , o . neovolkiana , o . nigrella , o . superficialis , o . variabilis , and sphaerocordyceps palustris ( table 1 ) . out of 27 cordyceps spp . , 26 are recorded on larvae except c . scarabaeicola that is recorded on adult ( table 1 ) .
table 1 : an aggregate list of coleopteran hosts ( family / genus / species ) of cordyceps species .
members of this family are commonly called earth - boring dung beetles , which mostly excavate burrows to lay their eggs . they are normally detritivores but occasionally behave as coprophagous . cordyceps geotrupis is recorded on an adult of geotrupes sp . in this family ( table 1 ) .
members of this family are commonly called stag beetles . the name stag beetle is derived from the distinctive mandibles found on the males that resemble the antlers of stags . the larvae feed on rotting deciduous wood for several years before pupating . ophiocordyceps scottiana is recorded on rhyssonotus nebulosus in this family ( table 1 ) .
members of this family are commonly called click beetles , elaters , snapping beetles , spring beetles , skipjacks , and so forth . the adults are typically nocturnal and phytophagous . larvae are slender , elongate , cylindrical , or somewhat flattened , with relatively hard bodies , somewhat resembling mealworms . fifteen spp . of cordyceps have been recorded from this family ( table 1 ) , all of which grow on larvae except ophiocordyceps salebrosa that grows on an adult ( table 1 ) . six spp . are recorded on larvae of the following hosts : cordyceps nirtolii on melanotus communis , c . shanxiensis on m . caudex and pleonomus canaliculatus , metarhizium campsosterni on campsosternus auratus , m . martiale on hemirhipus sp . , ophiocordyceps jiangxiensis on c . auratus and c . fruhstorferi , and o . stylophora on denticollis linearis ( table 1 ) . other spp . recorded in this family are cordyceps aurantiaca , c . huntii , c . rubra , c . velutipes , ophiocordyceps brunneipunctata , o . elateridicola , o . gracilioides , o . purpureostromata , and o . salebrosa ( table 1 ) .
the long - horned beetles , also known as longicorns , are cosmopolitan , typically characterized by extremely long antennae , which are often as long as or longer than the beetle’s body . the larvae , called roundheaded borers , bore into wood , where they can cause extensive damage to either living trees or untreated lumber . four spp . are known from this family , all growing on larvae ( table 1 ) . they are ophiocordyceps dovei on oemona hirta , o . entomorrhiza on leptura sp . , o . larvicola on callidium sp . , and o . stylophora on phoracantha semipunctata .
the family , commonly known as leaf beetle , is one of the largest beetle families . adults and larvae feed on all sorts of plant tissues . most chrysomelids are conspicuously colored , typically in glossy yellow to red or metallic blue - green hues . ophiocordyceps entomorrhiza is recorded on adult of diabrotica sp . and o . superficialis on a chrysomelid larva ( table 1 ) .
it is a family of the pleasing fungus beetles . they feed on plant and fungal matter , a few of them being notable pests . cordyceps erotyli is recorded on adult of erotylus sp . in this family ( table 1 ) .
it is the family of the true weevils or snout beetles . weevils are almost entirely plant feeders . cordyceps militaris is recorded on larva of ips sexdentatus , ophiocordyceps curculionum on adult of heilipus celsus , o . entomorrhiza on adult of apion flavipes , and podocrella peltata on larva of cryptorhynchus corticicolus ( table 1 ) .
tenebrionids are often referred to as darkling beetles as they have usually black or brown elytra and are nocturnal in habit . they are found worldwide though they are more diverse in semiarid and arid ecosystems . adults of most species are saprophagous while larvae are mostly detritivores ( litter feeders ) or xylophagous . five spp . are recorded on larvae of this family ( table 1 ) : c . militaris on tenebrio molitor , o . acicularis on nictobates sp . , o . entomorrhiza on meneristes laticollis , and o . larvicola on cylindronotus sp . , helops caraboides , and h . lanipes . ophiocordyceps formosana is the fifth species recorded on larva of this family .
the rove beetles belong to family staphylinidae , primarily distinguished by their short elytra . the group is currently recognized as the largest family of beetles . they are an ecologically and morphologically diverse group of beetles and commonly encountered in terrestrial ecosystems . cordyceps memorabilis is recorded on adult of staphylinus sp . whereas ophiocordyceps entomorrhiza is recorded on larva of ocypus sp . cordyceps staphylinidicola and o . superficialis are other spp . recorded on larvae of this family ( table 1 ) .
it is a large , cosmopolitan family . its members are commonly known as ground beetles . they are mostly shiny black or metallic and have ridged wing covers ( elytra ) . common habitats are under the bark of trees , under logs , or among rocks or sand by the edge of ponds and rivers . six spp . are known from this family ( table 1 ) . among them , o . entomorrhiza is recorded on adults and larvae of several species such as carabus auronitens , c . coriaceus , c . glabratus , c . hortensis , c . intricatus , c . nemoralis , c . nemorensis , c . violaceus , calathus sp . , calosoma sp . , coptolabrus sp . , hadrocarabus problematicus , and pterostichus sp . ophiocordyceps volkiana is recorded on larva of eripus heterogaster . cordyceps nikkoensis , ophiocordyceps carabidicola , o . michiganensis , and o . stylophora are other species recorded on larvae of this family .
among the coleopteran families , scarabaeidae is parasitized by the highest number of cordyceps fungi ( 27 spp . ) , followed by elateridae ( 15 spp . ) ( table 1 ) . among 55 parasitizing fungi ( table 1 ) , we found that only few parasitize more than one family . for example , ophiocordyceps entomorrhiza parasitizes 6 families , followed by o . stylophora ( 4 families ) , o . larvicola and o . superficialis ( 3 families each ) , and c . militaris , c . velutipes , and o . michiganensis ( 2 families each ) ( table 1 ) . from host range point of view , ophiocordyceps entomorrhiza shows the widest range , infecting 13 spp . in carabidae and one sp . in each cerambycidae , chrysomelidae , curculionidae , staphylinidae , and tenebrionidae ( table 1 ) . it is also exceptional in that it infects both larval and adult stages .
as mentioned earlier , larva is the most dominant host stage susceptible to cordyceps pathogens in coleoptera , with fewer adult stages being parasitized ( table 1 ) . besides o . entomorrhiza , six other spp . are recorded on coleopteran adults . they are c . erotyli , c . geotrupis , c . memorabilis , o . curculionum , o . salebrosa , and c . scarabaeicola ; the remaining ones are recorded on larvae .
lepidoptera is among the large orders of insects [ 121 ] . it has around 160 , 000 spp . that are classified into 4 suborders , 45 superfamilies , and 139 families [ 122 ] . out of four suborders ( aglossata , glossata , heterobathmiina , and zeugloptera ) , cordyceps and allied genera are known only from glossata . it is the largest suborder consisting of almost 99 . 9 % of all described lepidopterans [ 122 , 123 ] . it is further classified into six infraorders ( dacnonypha , acanthoctesia , lophocoronina , neopseustina , exoporia , and heteroneura ) [ 122 ] . among them , cordyceps hosts are known from two infraorders exoporia and heteroneura .
lepidopterans have from three to more than a dozen larval instars ( caterpillars ) , often five [ 123 , 124 ] , inhabiting as root - or stem - borers or foliage eaters and leaf - miners of angiosperms , usually within narrow range of host plants . lepidopterans have distinct feeding habits , basically plant - feeding during larval stage and nectar - feeding during adult stage . exoporia and heteroneura are discussed below in short , followed by cordyceps pathogens parasitizing them .
it is a small infraorder consisting of 636 spp . that are classified into 2 superfamilies [ 122 ] . cordyceps and allied genera are recorded only from the superfamily hepialoidea . the superfamily comprises five families that are distributed in diverse vegetation such as forest , shrubland , grassland , tundra , swamp , and bog with the most varied diet habit among moth families [ 123 , 125 ] . among the families , cordyceps spp . are recorded from family hepialidae alone ( table 2 ) .
table 2 : cordyceps species recorded on hepialid hosts ( lepidoptera , glossata , exoporia , and hepialidae ) .
many species are recorded in australia and new zealand on hepialid hosts . for instance , cordyceps cranstounii , c . hawkesii , d . gunnii , o . robertsii , and o . taylorii are recorded on abantiades labyrinthicus and other abantiades spp . , aenetus virescens , aoraia dinodes , a . ensyii , dasypodia selenophora , oxycanus dirempta , and other oxycanus spp . , trictena atripalpis and other trictena spp . , wiseana spp . , and so forth ( table 2 ) . another cordyceps sp . rarely described on hepialus is c . militaris ( table 2 ) . besides that , cordyceps cuncunae and o . emeiensis are also recorded on hepialid hosts ( table 2 ) .
this infraorder consists of more than 98 % of lepidopteran species [ 122 ] . nine superfamilies among more than 30 in the infraorder are recorded as hosts of cordyceps spp . among the host families , tineidae is the only microlepidopteran family ; the rest are macrolepidopterans . papilionidae and pieridae are the two butterfly families , the rest being moths . the host families are briefly described below followed by cordyceps spp . recorded on them .
( 1 ) erebidae . it is the largest family in lepidoptera with 24 , 569 described spp . [ 122 ] . despite the high species richness , only two cordyceps spp . are recorded on this family . of them , c . militaris is recorded on calliteara pudibunda and leucoma salicis whereas c . nikkoensis is reported on a larva of the family ( table 3 ) .
table 3 : cordyceps species recorded on heteroneuran hosts ( lepidoptera , glossata , and heteroneura ) .
( 2 ) noctuidae . it is commonly known as owlet moths . cordyceps militaris is recorded on 4 spp . of this family : arcte coerula , colocasia coryli , euxoa ochrogaster , and panolis flammea ( table 3 ) . ophiocordyceps elongata is recorded on acronicta americana . other species recorded in this family are cordyceps alpicola , c . bifusispora , c . bulolensis , c . cristata , and c . tuberculata ( table 3 ) .
( 3 ) notodontidae . its members are mainly concentrated in the tropical areas . some examples of this family are puss moths . cordyceps militaris is recorded on 5 spp . of this family . they are fentonia ocypete , lampronadata cristata , phalera assimilis , p . bucephala , and syntypistis punctatella ( table 3 ) .
( 1 ) limacodidae ( cochlididae ) . the members of this family are known as slug moths and are mostly tropical . ophiocordyceps cochlidiicola is recorded on larva and pupa of this family ( table 3 ) .
( 1 ) cossidae . the members are known as carpenter millers and are found worldwide . four spp . are recorded in this family . of them , metarhizium indigoticum is recorded on yakudza vicarius . two recently described spp . , ophiocordyceps arborescens and o . macroacicularis , are recorded on cossus sp . ( table 3 ) . the other species recorded on this family is cordyceps bassiana .
( 1 ) drepanidae . the members of this family have worldwide distribution . cordyceps militaris is recorded on 4 spp . of this family : achlya flavicornis , ochropacha duplaris , tethea ocularis , and tetheella fluctuosa ( table 3 ) .
( 1 ) tineidae . the members are small to medium - sized moths . they are worldwide in distribution but are particularly common in the palaeartic ecozone . cordyceps cardinalis is recorded on larva of this family ( table 3 ) .
( 1 ) papilionidae . it is a family of colorful swallowtail butterflies . some of the members are the largest butterflies in the world . the majority are distributed in the tropical region . cordyceps tuberculata is reported on adult of this family ( table 3 ) .
( 2 ) pieridae . it is a large family of butterfly , mostly distributed in the tropical parts of the world . its members are mostly white , orange , or yellow in pigmentation . metarhizium taii is reported on pieris rapae in this family ( table 3 ) .
( 1 ) geometridae . it is the second largest family in lepidoptera with 23 , 002 spp . [ 122 ] . it is commonly known as inch worms . some of the members are notorious pests . cordyceps militaris and c . riverae are recorded from this family parasitizing biston panterinaria , lycia hirtaria , and triphosa sp . ( table 3 ) .
( 1 ) bombycidae . it is known as silkworm family . the most well - known member is bombyx mori , native to northern china . cordyceps militaris is recorded on b . mori and c . michaelisii on a species of bombyx ( table 3 ) .
( 2 ) endromidae . this is a small family . cordyceps militaris is recorded on andraca bipunctata in this family ( table 3 ) .
( 3 ) saturniidae . the members include giant silk moths , royal moths , and emperor moths . they are described worldwide but are particularly known from tropical and subtropical regions . some of the members are the largest moth species . cordyceps longdongensis parasitizes actias artemis whereas c . militaris parasitizes anisota senatoria ( table 3 ) .
( 1 ) lasiocampidae . its members are known as snout moths or lappet moths . they are large in size with broad wings and are known worldwide . cordyceps militaris is reported on dendrolimus pini , d . superans , and macrothylacia rubi ( table 3 ) .
out of 22 spp . recorded on heteroneuran hosts ( table 3 ) , c . militaris has the widest host range , extending to 2 infraorders , 6 superfamilies , 10 families , and 29 spp . ( table 3 ) . it is a cosmopolitan species , distributed from sea level to more than 2000 m above sea level [ 126 ] . probably due to its wide host range and adaptability to wider habitats , this species demonstrates rapid in vitro growth and fructifications [ 127 ] . cordyceps tuberculata is another species recorded on multiple families . larva is the most suitable host for cordyceps spp . compared to pupa and adult ( table 3 ) .
in total , 16 families in lepidoptera ( 1 exoporian and 15 heteroneuran ) are identified as host families of cordyceps and allied genera . hepialidae , though a small family , hosts 6 cordyceps spp . , 1 drechmeria sp . , and 10 ophiocordyceps spp . ( table 2 ) . they are mostly distributed in either asia or australia / new zealand . after hepialidae , noctuidae and sphingidae are the families mostly infected by cordyceps spp . ( table 3 ) .