agathodes designalis , the sky-pointing moth , is a moth in the crambidae family .

it was described by guenée in 1854 .

it is found from the southern united states , where it has been recorded from arizona , texas and florida , south to southern south america .

it is also found on the west indies .

the length of the forewings is 13-18.5 mm .

the forewings are marbled purple-and-green .

the hindwings are beige .

adults are on wing from may to september in four generations in florida .

the larvae feed on erythrina ( including erythrina flabelliformis ) , inga and citharexylum species .

the summer and fall generations feed on the leaves of their host plant , while larvae of the spring generation feed on the flowers .

young larvae are translucent and orange .

older larvae have an orange body with cream stripes and a bright red head .

agathodes designalis

there are nine other species of agathodes described . for example , agathodes ostentalis , similar to agathodes designalis , is found throughout asia and australia . larvae of agathodes ostentalis also feed on erythrina , but are different in appearance from those of agathodes designalis .
lateral view of pupa of agathodes designalis guenée . photographed in gainesville , florida .
an adult erythrina leaf - roller , agathodes designalis guenée . photographed in gainesville , florida .
prepupa , fall generation , of agathodes designalis guenée on erythrina herbacea . photographed in gainesville , florida .
eggs : eggs of agathodes designalis are laid singly on the ventral side of the leaf ( bourquin 1945 ) .
in florida , and throughout the north american populations , agathodes designalis is represented by the subspecies agathodes designalis monstralis , while agathodes designalis designalis was originally described from south america . additional work on the genus , including new methods such as the dna barcoding , is likely to change current classification of the genus ( dan janzen , pers . com . ) . for instance , larvae of agathodes designalis in costa rica are different from those in florida , perhaps emphasizing the underlying taxonomic differences ( janzen & hallwachs 2011 ; sourakov 2011 ) .
mature larva , spring generation , of agathodes designalis guenée feeding on the inflorescence of erythrina herbacea . photographed in gainesville , florida .
agathodes designalis has a marginal effect on erythrina herbacea , though it can destroy flowers , making coral bean less attractive as an ornamental .
there are nine other species of agathodes described . for example , agathodes ostentalis , similar to a . designalis , is found throughout asia and australia . larvae of a . ostentalis also feed on erythrina , but are different in appearance from those of a . designalis .
fourth instar larva , spring generation , of agathodes designalis guenée feeding on the inflorescence of erythrina herbacea . photographed in gainesville , florida .
larvae : young larvae of agathodes designalis are translucent and orange with six rows of short , black - sclerotized tubercles , and are cryptic .
prepupa , spring generation , of agathodes designalis guenée changes color after feeding on the inflorescence of erythrina herbacea . photographed in gainesville , florida .
coral bean , erythrina herbacea , a host plant of terastia meticulosalis guenée and agathodes designalis guenée . photographed in april at gainesville , florida .
coral bean , erythrina herbacea , a host plant of terastia meticulosalis guenée and agathodes designalis guenée . photographed during may at gainesville , florida .
agathodes designalis has a marginal effect on e . herbacea , though it can destroy flowers , making coral bean less attractive as an ornamental .
bourquin f . 1932 . observaciones biológicas sobre agathodes designalis ( fam . pyraustidae ) . revista de la sociedad entomológica argentina 5 : 13–14 .
figure 27 . lateral view of pupa of agathodes designalis guenée . photographed in gainesville , florida . photograph by andrei sourakov , university of florida .
dorsalk view of a mature larva , summer generation , of agathodes designalis guenée feeding on leaves of erythrina herbacea . photographed in gainesville , florida .
lateral view of a mature larva , summer generation , of agathodes designalis guenée feeding on leaves of erythrina herbacea . photographed in gainesville , florida .
frontal view of a mature larva , summer generation , of agathodes designalis guenée feeding on leaves of erythrina herbacea . photographed in gainesville , florida .
bourquin f . 1932 . observaciones biológicas sobre agathodes designalis ( fam . pyraustidae ) . revista de la sociedad entomológica argentina 5 : 13 - 14 .
mature larva , summer generation , of agathodes designalis guenée inside a shelter made from from the leaves of erythrina herbacea . photographed in gainesville , florida .
figure 2 . an adult erythrina leaf - roller , agathodes designalis guenée . photographed in gainesville , florida . photograph by andrei sourakov , university of florida .
figure 15 . an adult erythrina leaf - roller , agathodes designalis guenée . photographed in gainesville , florida . photograph by andrei sourakov , university of florida .
webbing of the first and second instar larvae , spring generation , of agathodes designalis guenée over the inflorescence of erythrina herbacea . photographed in gainesville , florida .
brou jr . , v . a . , 2008 . agathodes designalis guenée , 1854 ( lepidoptera : pyralidae ) in louisiana . southern lepidopterists ' society , pdf .
figure 26 . prepupa , fall generation , of agathodes designalis guenée on erythrina herbacea . photographed in gainesville , florida . photograph by andrei sourakov , university of florida .
mature larva , spring generation , of agathodes designalis guenée suspends itself by a silk thread afrer falling off the host plant , erythrina herbacea . photographed in gainesville , florida .
summer and fall generations of agathodes designalis feed on leaves and develop slower than the spring generation that feeds on flowers , which indicates the higher nutritional value of their spring diet .
figure 18 . mature larva , spring generation , of agathodes designalis guenée feeding on the inflorescence of erythrina herbacea . photographed in gainesville , florida . photograph by andrei sourakov , university of florida .
figure 20 . mature larva , spring generation , of agathodes designalis guenée feeding on the inflorescence of erythrina herbacea . photographed in gainesville , florida . photograph by andrei sourakov , university of florida .
sourakov a . 2011 . niche partitioning , co - evolution and life histories of erythrina moths , terastia meticulosalis and agathodes designalis ( lepidoptera : crambidae ) . tropical lepidoptera research 21 : 84–94 .
figure 17 . fourth instar larva , spring generation , of agathodes designalis guenée feeding on the inflorescence of erythrina herbacea . photographed in gainesville , florida . photograph by andrei sourakov , university of florida .
there are four generations of agathodes designalis between may and september in north - central florida . larvae of agathodes designalis collected in gainesville , florida in september completed their feeding by mid - october , and diapaused in a prepupal stage inside cocoons ( sourakov , 2011 ) . the diapause might not be present in the warmer parts of the moth ' s range , or might occur in different seasons .
agathodes designalis specimens reared in florida and costa rica are ~ 1 . 2 inches ( 3 . 0 cm ) in wing span , but specimens from south america found in collections can be larger .
niche partitioning , co - evolution and life histories of erythrina moths , terastia meticulosalis and agathodes designalis andrei sourakov . 2011 . trop . lepid . res . 21 ( 2 ) : 84 - 94 .
figure 21 . prepupa , spring generation , of agathodes designalis guenée changes color after feeding on the inflorescence of erythrina herbacea . photographed in gainesville , florida . photograph by andrei sourakov , university of florida .
figure 28 . coral bean , erythrina herbacea , a host plant of terastia meticulosalis guenée and agathodes designalis guenée . photographed in april at gainesville , florida . photograph by andrei sourakov , university of florida .
figure 29 . coral bean , erythrina herbacea , a host plant of terastia meticulosalis guenée and agathodes designalis guenée . photographed during may at gainesville , florida . photograph by andrei sourakov , university of florida .
sourakov a . 2011 . niche partitioning , co - evolution and life histories of erythrina moths , terastia meticulosalis and agathodes designalis ( lepidoptera : crambidae ) . tropical lepidoptera research 21 : 84 - 94 .
adult : agathodes designalis specimens reared in florida and costa rica are ~ 1 . 2 inches ( 3 . 0 cm ) in wing span , but specimens from south america found in collections can be larger .
figure 23 . dorsal view of a mature larva , summer generation , of agathodes designalis guenée feeding on leaves of erythrina herbacea . photographed in gainesville , florida . photograph by andrei sourakov , university of florida .
figure 24 . lateral view of a mature larva , summer generation , of agathodes designalis guenée feeding on leaves of erythrina herbacea . photographed in gainesville , florida . photograph by andrei sourakov , university of florida .
figure 25 . frontal view of a mature larva , summer generation , of agathodes designalis guenée feeding on leaves of erythrina herbacea . photographed in gainesville , florida . photograph by andrei sourakov , university of florida .
figure 22 . mature larva , summer generation , of agathodes designalis guenée inside a shelter made from from the leaves of erythrina herbacea . photographed in gainesville , florida . photograph by andrei sourakov , university of florida .
bourquin f . 1945 . observaciones biológicas sobre agathodes designalis . pp . 117 - 119 . in mariposas argentinas . vida , desarrollo , costumbres y hechos curiosos de algunos lepidópteros argentinos . buenos aires . 212 pp .
figure 16 . webbing of the first and second instar larvae , spring generation , of agathodes designalis guenée over the inflorescence of erythrina herbacea . photographed in gainesville , florida . photograph by andrei sourakov , university of florida .
pupae : agathodes designalis makes loose double - layered cocoons similar to these of terastia meticulosalis , and the pupa , though it appears more glossy and more uniformly colored brown , is otherwise similar to that of terastia meticulosalis .
agathodes designalis makes loose double - layered cocoons similar to these of t . meticulosalis , and the pupa , though it appears more glossy and more uniformly colored brown , is otherwise similar to that of t . meticulosalis .
eggs of a . designalis are laid singly on the ventral side of the leaf ( bourquin 1945 ) .
in florida , and throughout the north american populations , a . designalis is represented by the subspecies a . designalis monstralis , while a . designalis designalis was originally described from south america . additional work on the genus , including new methods such as the dna barcoding , is likely to change current classification of the genus ( dan janzen , pers . com . ) . for instance , larvae of a . designalis in costa rica are different from those in florida , perhaps emphasizing the underlying taxonomic differences ( janzen & hallwachs 2011 ; sourakov 2011 ) .
figure 19 . mature larva , spring generation , of agathodes designalis guenée suspends itself by a silk thread after falling off the host plant , erythrina herbacea . photographed in gainesville , florida . photograph by andrei sourakov , university of florida .
there are 17 known species in the genus agathodes but it is likely that the true number of species is considerably higher .
young larvae of a . designalis are translucent and orange with six rows of short , black - sclerotized tubercles , and are cryptic .
summer and fall generations of a . designalis feed on leaves and develop slower than the spring generation that feeds on flowers , which indicates the higher nutritional value of their spring diet .
the purple - and - green marbled forewings of agathodes designalis are colorful , while the hind wings are beige . the abdomen is held in an up - right position , which gave the moth an unofficial common name of
sky - pointing moth .
males have a pair of hair - pencils that are extended during
female calling ,
for the release of pheromones .
there are four generations of a . designalis between may and september in north - central florida . larvae of a . designalis collected in gainesville , florida , in september completed their feeding by mid - october and diapaused in a prepupal stage inside cocoons ( sourakov 2011 ) . the diapause might not be present in the warmer parts of the moth ' s range or might occur in different seasons .
habitually adopt a jet - fighter resting posture . after settling they slowly raise their abdomens . some species rest with the abdomen raised vertically or canted sideways , while others including designalis curve it forwards over the back , scorpion - style .
although little known , the erythrina moths terastia meticulosalis guenée and agathodes designalis guenée represent a remarkable case of niche partitioning ( sourakov 2011 ) . in florida , throughout the southern united states , and from mexico to argentina , these two species feed on plants of the genus erythrina ( fabaceae ) . this genus , while mostly known as attractive ornamentals , has medicinal and other uses ( powell and westley 1993 ) . the descriptions of the moths ' life histories below are based on north - central florida populations that use the coral bean , erythrina herbacea , as a hostplant . this plant is not only native to the region , but is also frequently used as an ornamental .
sourakov , a . , d . plotkin , a . y . kawahara , l . xiao , w . hallwachs , d . janzen . 2015 . on the taxonomy of the erythrina moths agathodes and terastia ( crambidae : spilomelinae ) : two different patterns of haplotype divergence and a new species of terastia . tropical lepidoptera research , 25 ( 2 ) : 80–98 .
the purple - and - green marbled forewings of a . designalis are colorful , while the hind wings are beige . the abdomen is held in an up - right position , which gave the moth an unofficial common name of
sky - pointing moth .
males have a pair of hair - pencils that are extended during
female calling ,
for the release of pheromones .
photographs are the copyrighted property of each photographer listed . contact individual photographers for permission to use for any purpose .
powell , j . a . & p . a . opler , moths of western north america , pl . 23 . 32m ; p . 179 . book review and ordering
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adult : purple - and - green marbled , hindwings are beige , abdomen held in an up - right position , males with pair of hair - pencils that are extended for the release of pheromones .
larva : early instars translucent and orange with six rows of short , black - sclerotized tubercles , and are cryptic . later instars with cream - colored longitudinal stripes , and the black tubercles become more prominent on the orange background , head is bright red .
mostly coastal localities from sc to e tx , plus se az / w . indies - map ( mpg )
eggs laid singly on the ventral side of the leaf ( bourquin 1945 ) . see complete life history by sourakov , 2011
boisduval & guenée , 1854 . histoire naturelle des insectes spécies général des lépidoptères 8 : 209 .
powell , j . a . & p . a . opler , 2009 . moths of western north america . university of california press . p . 179 , pl . 23 . 32
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figure 1 . an adult erythrina stem - borer , terastia meticulosalis guenée . photographed in gainesville , florida . photograph by andrei sourakov , university of florida .
there are four other species within the genus terastia . in the past , terastia subjectalis ( found in australia and throughout asia ) had been considered a synonym of terastia meticulosalis , but now is listed in modern literature as a separate species . the taxonomic status of terastia meticulosalis populations in africa , where it is listed from kenya , south africa , uganda and sierra leone , is unclear .
both species are found from south carolina to florida in the eastern united states , west to arizona , and then south to argentina .
in florida and throughout the southeastern united states , both species are quite distinct , but in collections they can be confused with members of the same genera , which are distributed world - wide .
adults : in terastia meticulosalis , size depends on the generation and locality , and therefore the diet , of the caterpillars . for instance , in florida , the spring generation that feeds on seeds was shown to be larger than the summer and fall generations that feed inside the stem . the wing span for spring generations was ~ 1 . 4 inches ( 3 . 7 cm ) versus ~ 1 . 2 inches ( 3 . 0 cm ) for summer generations , and as small as ~ 1 . 0 inches ( 2 . 5 cm ) in fall generations . ( sourakov , 2011 ) . collection specimens from south america are notably larger ( ~ 1 . 8 inches or 4 . 6 cm ) .
the marbled - brown forewings of terastia meticulosalis make this species cryptic when at rest . however , its hind wings are white , as seen above .
figure 3 . an adult erythrina stem - borer , terastia meticulosalis guenée . photographed in gainesville , florida . photograph by andrei sourakov , university of florida .
larvae : the young larvae of terastia meticulosalis are found inside the stems of erythrina herbacea , where their feeding produces a characteristic dying - off of the tip of the host plant . the infestation rate by terastia meticulosalis can be relatively high . as the availability of undamaged shoots of erythrina herbacea declines , neonate larvae bore and feed inside the leaf stalks and later make their way inside the stems .
figure 4 . erythrina herbacea in july , showing a characteristic dying off of the stem as the larva of terastia meticulosalis guenée burrows downwards . photographed in gainesville , florida . photograph by andrei sourakov , university of florida .
in younger larvae , the dorsal prothoracic plate and the head are black and heavily sclerotized . in older larvae , the prothoracic plate becomes cream - colored , slightly darker than the rest of the body , which is otherwise translucent and cream - colored .
figure 5 . second instar larva of terastia meticulosalis guenée . photographed in gainesville , florida . photograph by andrei sourakov , university of florida .
in the spring , young larvae of terastia meticulosalis infest the tips of the stem and gradually kill off the upper part of the plant , from where they move into the pods to feed on seeds .
figure 6 . third instar larva of terastia meticulosalis guenée , spring generation . photographed in gainesville , florida . photograph by andrei sourakov , university of florida .
prior to pupation , last instar larvae are able to consume large numbers of seeds and move from one pod to another . larvae of terastia meticulosalis sometimes accumulate red pigments prior to exiting pods to pupate on the forest floor . however , this was only observed in the spring generation that feeds on seeds ( sourakov 2011 ) .
figure 7 . prepupa of terastia meticulosalis guenée . photographed in gainesville , florida . photograph by andrei sourakov , university of florida .
the larvae of summer and fall generations of terastia meticulosalis feed inside the stems until pupation ( the seeds become hard and pods splits open during this time ) . hence , the summer / fall larvae look paler than spring ones .
figure 8 . mature larva of terastia meticulosalis guenée , hollowing the stem of erythrina herbacea , summer / fall generations . photographed in gainesville , florida . photograph by andrei sourakov , university of florida .
figure 9 . mature larva of terastia meticulosalis guenée , summer / fall generations . photographed in gainesville , florida . photograph by andrei sourakov , university of florida .
figure 10 . head and thorax of mature larva of terastia meticulosalis guenée , summer / fall generations . photographed in gainesville , florida . photograph by andrei sourakov , university of florida .
figure 11 . the larva of terastia meticulosalis guenée purges the hollow stem of erythrina herbacea from frass by crawling backwards to the entrance when defecating . photographed in gainesville , florida . photograph by andrei sourakov , university of florida .
pupae : terastia meticulosalis makes loose double - layered cocoons . the pupa is cream - colored at first , and then becomes light brown , with the proboscis extending far into the abdominal segments .
figure 12 . the pupa of terastia meticulosalis guenée inside its cocoon . photographed in gainesville , florida . photograph by andrei sourakov , university of florida .
figure 13 . the pupa of terastia meticulosalis guenée , lateral view . photographed in gainesville , florida . photograph by andrei sourakov , university of florida .
figure 14 . the pupa of terastia meticulosalis guenée , ventral view . photographed in gainesville , florida . photograph by andrei sourakov , university of florida .
the later instar larvae and prepupae appear aposematic . they develop cream - colored longitudinal stripes and the black tubercles become more prominent on the orange background , while the head is bright red .
in the spring generation that feeds on flowers , the prepupa can become red . the prepupae of summer and fall generations are cream - colored .
figure 30 . coral bean , erythrina herbacea , plant in late spring ( june ) following an infestation , with pods hollowed by larvae of terastia meticulosalis guenée . photographed in gainesville , florida . photograph by andrei sourakov , university of florida .
figure 31 . coral bean , erythrina herbacea , a hostplant of terastia meticulosalis guenée , as seen in august with pods open and seeds exposed . photographed in gainesville , florida . photograph by andrei sourakov , university of florida .
terastia meticulosalis is a serious pest of naturally - occurring and commercially grown erythrina , a genus valued for its many uses from agricultural and ornamental to medicinal and pest control ( powell & westley 1993 ) . raven ( 1974 ) suggested that terastia meticulosalis makes the cultivation of erythrina in southern florida almost impossible .
despite its small size , terastia meticulosalis requires a substantial amount of plant material to develop to the adult stage , including seeds . plants can be substantially damaged by larvae boring into the center of the stem , killing parts of the plant above . as a result , the health of host plants can be severely affected and the whole plant , or at least its reproductive organs , are often destroyed .
bhattacharya dp , mandal dk . 1976 . a new record of terastia meticulosalis guenée ( lepidoptera : pyralidae ) from the gt . nicobar island . newsletter zoological survey of india 2 : 23 - 24 .
brown sh . 2011 . erythrina herbacea , coral - bean , cardinal - spear , cherokee - bean . lee county - southwest florida ( 5 april 2013 ) .
dyar hg . 1901 . descriptions of some pyralid larvae from southern florida . journal of the new york entomological society 9 : 19 - 24 .
goff r . 2011 . terastia meticulosalis guenée , 1854 . african moths ( 5 april 2013 ) .
heppner jb . 2007 . lepidoptera of florida , part 1 : introduction and catalog . arthropods of florida and neighboring land areas . vol . 17 . florida department of agriculture , 670 pp .
janzen dh , hallwachs w . 2011 . area de conservación guanacaste ( acg ) , northwestern costa rica : caterpillars , pupae , butterflies & moths database . ( 5 april 2013 ) .
peck sb , heraty j , landry b , sinclair bj . 1998 . introduced insect fauna of an oceanic archipelago : the galápagos islands , ecuador . american entomologist 44 : 218 - 237 .
powell ja , opler pa . 2009 . moths of western north america . university of california press , berkeley , 369 pp .
powell mh , westley sb . ( editors ) . 1993 . erythrina production and use : a field manual . nitrogen fixing tree association . paia , hawaii . 55 pp .
raven ph . 1974 . erythrina ( fabaceae ) : achievements and opportunities . lloydia 37 : 321 - 331 .
swezey oh . 1923 . the erythrina twig - borer ( terastia meticulosalis ) in hawaii ( pyralidae , lepidoptera ) . proceedings of the hawaiian entomological society 5 : 297 - 298 .
taylor js . 1951 . notes on lepidoptera in the eastern cape province ( part ii ) . journal of the entomological society of southern africa 14 : 94 - 126 .
usgs . ( april 2006 ) . erythrina herbacea . geology and environmental change science center ( 5 april 2013 ) .
zimmerman ec . 1958 . insects of hawaii : a manual of the insects of the hawaiian islands , including an enumeration of the species and notes on their origin , distribution , hosts , parasites , etc . volume 8 ( lepidoptera : pyraloidea ) . university of hawaii press , honolulu . 456 pp .
photographs : andrei sourakov , florida museum of natural history , university of florida .
there are four other species within the genus terastia . in the past , t . subjectalis ( found in australia and throughout asia ) had been considered a synonym of t . meticulosalis , but now is listed in modern literature as a separate species . the taxonomic status of t . meticulosalis populations in africa , where it is listed from kenya , south africa , uganda , and sierra leone , is unclear .
an adult erythrina stem - borer , terastia meticulosalis guenée . photographed in gainesville , florida
in t . meticulosalis , size depends on the generation and locality and , therefore , the diet , of the caterpillars . for instance , in florida , the spring generation that feeds on seeds was shown to be larger than the summer and fall generations that feed inside the stem . the wing span for spring generations was ~ 1 . 4 inches ( 3 . 7 cm ) versus ~ 1 . 2 inches ( 3 . 0 cm ) for summer generations , and as small as ~ 1 . 0 inches ( 2 . 5 cm ) in fall generations ( sourakov 2011 ) . collection specimens from south america are notably larger ( ~ 1 . 8 inches or 4 . 6 cm ) .
the marbled - brown forewings of t . meticulosalis make this species cryptic when at rest . however , its hind wings are white , as seen above .
an adult erythrina stem - borer , terastia meticulosalis guenée . photographed in gainesville , florida .
the young larvae of t . meticulosalis are found inside the stems of erythrina herbacea , where their feeding produces a characteristic dying - off of the tip of the host plant . the infestation rate by t . meticulosalis can be relatively high . as the availability of undamaged shoots of e . herbacea declines , neonate larvae bore and feed inside the leaf stalks and later make their way inside the stems .
erythrina herbacea in july , showing a characteristic dying off of the stem asthe larva of terastia meticulosalis guenée burrows downwards . photographed in gainesville , florida .
in the spring , young larvae of t . meticulosalis infest the tips of the stem and gradually kill off the upper part of the plant , from where they move into the pods to feed on seeds .
third instar larva of terastia meticulosalis guenée , spring generation . photographed in gainesville , florida .
prior to pupation , last instar larvae are able to consume large numbers of seeds and move from one pod to another . larvae of t . meticulosalis sometimes accumulate red pigments prior to exiting pods to pupate on the forest floor . however , this was only observed in the spring generation that feeds on seeds ( sourakov 2011 ) .
the larvae of summer and fall generations of t . meticulosalis feed inside the stems until pupation ( the seeds become hard and pods splits open during this time ) . hence , the summer / fall larvae look paler than spring ones .
mature larva of terastia meticulosalis guenée , hollowing the stem of erythrina herbacea , summer / fallgenerations . photographed in gainesville , florida .
mature larva of terastia meticulosalis guenée , summer / fall generations . photographed in gainesville , florida .
head and thorax of mature larva of terastia meticulosalis guenée , summer / fall generations . photographed in gainesville , florida .
the larva of terastia meticulosalis guenée purges the hollow stem of erythrina herbacea from frass by crawling backwards to the entrance when defecating . photographed in gainesville , florida .
terastia meticulosalis makes loose double - layered cocoons . the pupa is cream - colored at first and then becomes light brown with the proboscis extending far into the abdominal segments .
the pupa of terastia meticulosalis guenée inside its cocoon . photographed in gainesville , florida .
the pupa of terastia meticulosalis guenée , lateral view . photographed in gainesville , florida .
the pupa of terastia meticulosalis guenée , ventral view . photographed in gainesville , florida .
the later instar larvae and prepupae appear aposematic . they develop cream - colored longitudinal stripes , and the black tubercles become more prominent on the orange background , while the head is bright red .
coral bean , erythrina herbacea , plant in late spring ( june ) following an infestation , with pods hollowed by larvae of terastia meticulosalis guenée . photographed in gainesville , florida .
coral bean , erythrina herbacea , a hostplant of terastia meticulosalis guenée , as seen in august with pods open and seeds exposed . photographed in gainesville , florida .
terastia meticulosalis is a serious pest of naturally - occurring and commercially grown erythrina , a genus valued for its many uses from agricultural and ornamental to medicinal and pest control ( powell & westley 1993 ) . raven ( 1974 ) suggested that t . meticulosalis makes the cultivation of erythrina in southern florida almost impossible .
despite its small size , t . meticulosalis requires a substantial amount of plant material to develop to the adult stage , including seeds . plants can be substantially damaged by larvae boring into the center of the stem , killing parts of the plant above . as a result , the health of host plants can be severely affected and the whole plant or at least its reproductive organs are often destroyed .
bhattacharya dp , mandal dk . 1976 . a new record of terastia meticulosalis guenée ( lepidoptera : pyralidae ) from the gt . nicobar island . newsletter zoological survey of india 2 : 23–24 .
brown sh . 2011 . erythrina herbacea , coral - bean , cardinal - spear , cherokee - bean . lee county - southwest florida . urltoken ( 5 january 2012 ) .
dyar hg . 1901 . descriptions of some pyralid larvae from southern florida . journal of the new york entomological society 9 : 19–24 .
janzen dh , hallwachs w . 2011 . area de conservación guanacaste ( acg ) , northwestern costa rica : caterpillars , pupae , butterflies & moths database . urltoken ( 5 january 2012 ) .
peck sb , heraty j , landry b , sinclair bj . 1998 . introduced insect fauna of an oceanic archipelago : the galápagos islands , ecuador . american entomologist 44 : 218–237 .
raven ph . 1974 . erythrina ( fabaceae ) : achievements and opportunities . lloydia 37 : 321–331 .
swezey oh . 1923 . the erythrina twig - borer ( terastia meticulosalis ) in hawaii ( pyralidae , lepidoptera ) . proceedings of the hawaiian entomological society 5 : 297–298 .
taylor js . 1951 . notes on lepidoptera in the eastern cape province ( part ii ) . journal of the entomological society of southern africa 14 : 94–126 .
usgs . ( april 2006 ) . erythrina herbacea . geology and environmental change science center . urltoken ( 5 january 2012 ) .
this host is a hybird : e . herbacea x e . crista - galli , and is a commercially sold ornamental
this document is eeny 516 , one of a series of the department of entomology and nematology , uf / ifas extension . original publication date february 2012 . revised april 2013 . reviewed june 2017 . visit the edis website at urltoken . this document is also available on the featured creatures website at urltoken .
the institute of food and agricultural sciences ( ifas ) is an equal opportunity institution authorized to provide research , educational information and other services only to individuals and institutions that function with non - discrimination with respect to race , creed , color , religion , age , disability , sex , sexual orientation , marital status , national origin , political opinions or affiliations . for more information on obtaining other uf / ifas extension publications , contact your county ' s uf / ifas extension office . u . s . department of agriculture , uf / ifas extension service , university of florida , ifas , florida a & m university cooperative extension program , and boards of county commissioners cooperating . nick t . place , dean for uf / ifas extension .
until recently the crambidae and pyralidae were grouped together as pyralidae , but the most recent revision by munroe & solis in 1999 classifies both as full families within the superfamily pyraloidea .
the crambidae comprises of about 11 , 630 known species worldwide , although the true number must be considerably higher , as these moths are generally small and a high percentage of them are dull and inconspicuous . the family is divided into 18 subfamilies , with the spilomelinae containing about 3 , 800 species worldwide . the taxonomy of the pyralidae and crambidae however are both still in disarray and the current classification is certain to change as more is learnt .
the illustrated species was photographed in rainforest at rio pindayo in eastern peru , at a altitude of about 200m . many other very similar species occur at higher altitudes , e . g . up to at least 2500m in the cloudforests of manu national park .
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diaphania costata ( f . ) ( lepidoptera : crambidae : spilomelinae ) , a commonly misidentified pest of ornamental apocynaceae . . .
by hayden , j . e . , e . r . hoebeke , m . a . bertone & v . a . brou jr .
proceedings of the entomological society of washington , 119 ( 2 ) : 173 - 190 . , 2017
hayden , j . e . , e . r . hoebeke , m . a . bertone & v . a . brou jr , 2017 . diaphania costata ( f . ) ( lepidoptera : crambidae : spilomelinae ) , a commonly misidentified pest of ornamental apocynaceae in the southern united states . proceedings of the entomological society of washington , 119 ( 2 ) : 173 - 190 . view and download at bioone here cite : 1415021
taxonomic revision of the spilomelinae ( lepidoptera , pyralidae s . l . ) of the galápagos islands , ecuador
kearfott , w . d . 1909 . descriptions of new species of north american crambid moths . proceedings of the united states national museum 35 : 367 - 393 .
a generic revision of the aquatic moths of north america : ( lepidoptera : pyralidae , nymphulinae ) .
solis m . a . , 2009 . transfer of all western hemisphere cybalomiinae to other subfamilies ( crambidae pyraloidea : lepidoptera ) : elusia schaus , dichochroma forbes , schacontia dyar , cybalomia extorris warren , and c . lojanalis ( dognin ) . proceedings of the entomological society of washington . 111 : 493–504 download at researchgate here
studies on the crambidae ( lepidoptera ) . part 41 . on some tropical crambidae with descriptions of new genera and species .
nacoleia charesalis ( walker , 1859 ) ( pyraloidea : crambidae : spilomelinae ) is a medium - sized brown moth with a native range from the southeast asian tropics to northern australia . it was first detected in southern florida , usa , in 2012 and known to be spreading rapidly northward . it resembles some native north american pyraustine species . this paper summarizes its current distribution , diagnostic characters and known behavior . download pdf here
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the arizona state university hasbrouck insect collection ( asuhic ) is one of the vital southwest arthropod collections in america north of mexico , providing important biological information . the principal objective of the catalog is to give a complete list of the lepidopterous insects held in the asuhic . furthermore , it will be an online catalog of the lepidoptera of arizona . the preliminary lepidoptera checklist is presented , consisting of 1983 species and 175 subspecies of 55 families in approximately 60 , 000 holdings at the asuhic . this article follows the recent classification and nomenclature ( hodges rw . 1983 . check list of the lepidoptera of america north of mexico . london , uk : e . w . classey ltd . and the wedge entomological research foundation ; moth photographers group ( mpg ) . 2014 . urltoken ) .
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illustrated in black and white by amsel ( 1954 ) pl . 80 , figs . 9 , 10 .


protostomia ( from greek πρωτο - proto - " first " and στόμα stoma " mouth " ) is a clade of animals .

together with the deuterostomes and xenacoelomorpha , its members make up the bilateria , mostly comprising animals with bilateral symmetry and three germ layers .

the major distinctions between deuterostomes and protostomes are found in embryonic development : most ( but , confusingly , not all ) protostomes undergo protostomy , whereas most deuterostomes undergo deuterostomy .

protostome

massive expansion and functional divergence of innate immune genes in a protostome . - pubmed - ncbi
what is a protostome ? .
grzimek ' s animal life encyclopedia . . retrieved july 09 , 2018 from urltoken urltoken
what made you want to look up protostome ? please tell us where you read or heard it ( including the quote , if possible ) .
what is a protostome ? .
grzimek ' s animal life encyclopedia . . encyclopedia . com . ( july 9 , 2018 ) . urltoken
‘if the ancestor of protostomes displayed a segmentation comparable to annelids and arthropods , this necessarily means that segmentation was lost independently in different protostome lineages . ’
► in the protostome p . caudatus , the blastopore forms the anus ( deuterostomy ) ► the hindgut markers brachyury and cdx are expressed in the blastopore and in the anus ► the foregut markers foxa and gsc are detected in the mouth , but not in the blastopore ► results favor deuterostomy in ecdysozoan , protostome , and bilaterian ancestors
citation : croset v , rytz r , cummins sf , budd a , brawand d , kaessmann h , et al . ( 2010 ) ancient protostome origin of chemosensory ionotropic glutamate receptors and the evolution of insect taste and olfaction . plos genet 6 ( 8 ) : e1001064 . urltoken
to handle and analyze large data sets , such as lists of morphological characters and nucleotide sequences . together with new paleontological discoveries in the fossil realm , these novel techniques and technologies provide modern biologists with a useful way to reexamine the traditional protostome relationships and to develop new hypotheses on animal relationships and evolution .
winnepenninckx , b . , t . backeljau , l . y . mackey , j . m . brooks , r . de wachter , s . kumar , and j . r . garey .
phylogeny of protostome worms derived from 18s rrna sequences .
molecular biology and evolution 12 ( 1995 ) : 641–649 .
to explore the evolutionary origin of the irs , we examined phylogenetic relationships of the identified protostome irs . reciprocal best - hit analysis using d . melanogaster sequences as queries revealed that a subset of this species ' irs was conserved in several distant lineages , allowing us to define putative orthologous groups . these include one group containing representatives of all protostome species ( ir25a ) , one represented by all arthropods ( ir93a ) , nine by most or all insects , and three by dipteran insects ( figure 2a and 2b ) . for most orthologous groups , a single gene for each species was identified . in a few cases , for example the ir75 group , certain species were represented by several closely related in - paralogues , some of which appeared to be pseudogenes ( figure 2a and 2b , table s3 , and datasets s1 and s2 ) .
the fate of the blastopore during development in the bilaterian ancestor is currently not well understood . in deuterostomes , the blastopore forms the anus , but its fate in protostome groups is variable [ 1 ] . this variability , combined with an absence of information from key taxa , hampers the reconstruction of the ancestral developmental mode of the protostomia and the bilateria . the blastopore fate of the bilaterian ancestor plays a crucial role in understanding the transition from radial to bilateral symmetric organisms [ 2 , 3 ] . priapulids have a conservative morphology , an abundant cambrian fossil record , and a phylogenetic position that make them a key group in understanding protostome evolution [ 4 , 5 ] . here , we characterize gastrulation and the embryonic expression of genes involved in bilaterian foregut and hindgut patterning in priapulus caudatus . we show that the blastopore gives rise to the anus at the vegetal pole and that the hindgut markers brachyury and caudal are expressed in the blastopore and anus , whereas the foregut markers foxa and goosecoid are expressed in the mouth in the animal hemisphere . thereby , gastrulation in the conservatively evolving protostome p . caudatus follows strictly a deuterostomic pattern . these results are more compatible with a deuterostomic rather than protostomic ( blastopore forms the mouth ) or amphistomic ( mouth and anus are formed simultaneously ) mode of development in the last common bilaterian ancestor .
additionally the point the authors are making in this paper is that a lot of the differences between
protostome
and
deuterostome
have to do with timing of turn - on of a - p axis genes ( e . g . six3 / 6 , goosecoid ) and mesoderm - specifying genes ( e . g . twist ) , so protostomy / deuterostomy is really sort of a heterochrony issue rather than diverging histories of body axis evolution .
several outstanding issues regarding ir evolution can now be addressed . first , we have shown that the irs were very likely to have been present in the last common ancestor of protostomia , an estimated 550–850 million years ago [ 20 ] . ir25a represents the probable oldest member of this repertoire and conservation of chemosensory organ expression of ir25a orthologues in molluscs , nematodes , crustaceans and insects strongly suggests that this receptor may have fulfilled a chemosensing function in the protostome ancestor .
well , not given what the authors here are saying . what they argue is that the underlying assumptions of what the blastopore is and isn ' t are probably wrong , and that
protostome
and
deuterostome
probably have more to do with blastopore closure than with a deep divide between animals where the first opening forms the mouth and those where the first opening forms the anus . there ' s a nuanced developmental argument here that the transformationist scenarios of yesteryear just don ' t have insight into .
contemporary research on protostome relationships utilizes a host of methods and technologies that were unavailable to biologists in the early twentieth century , such as grobben . modern biologists employ electron microscopy , fluorescent microscopy , biochemistry , and a collection of molecular techniques to sequence the genome , trace embryonic development , and gain insight into the origin of various genes and gene clusters , such as hox genes . other fields of research , including cladistic analysis and bioinformatics , continue to make important contributions . the latter fields are computer - based technologies that employ algorithms and statistics
over 100 years von groppen subdivided all bilateralsymmetric animals into protostomia and deuterostomia using embryological characters . recent molecular phylogenies support this grouping , but a closer examination of the embryology is drawing a different picture . all deuterostomes form the mouth separate from the blastopore , but protostomes show a high variability in the fate of the blastopore . i will provide new developmental data from different protostome taxa that suggest that deuterostomy is ancestral for the protostomes and thus for all bilateria . i will outline the impact of these finding for our understanding of animal body plan evolution .
well , not given what the authors here are saying . what they argue is that the underlying assumptions of what the blastopore is and isn ' t are probably wrong , and that
protostome
and
deuterostome
probably have more to do with blastopore closure than with a deep divide between animals where the first opening forms the mouth and those where the first opening forms the anus . there ' s a nuanced developmental argument here that the transformationist scenarios of yesteryear just don ' t have insight into . i had a bit of a conversation about this with one of the authors this summer , and it sounds like there ' s a lot more to come , but the conclusion should probably be that the ancestral bilaterian was a lot more complex than we give it credit for .
to investigate whether irs are likely to have an olfactory function beyond insects , we examined expression of the ir repertoire from a representative of a distantly related protostome lineage , aplysia molluscs , whose last common ancestor with d . melanogaster probably existed 550–850 million years ago [ 20 ] . we first used rt - pcr to analyse the expression of the ten aplysia ir genes in a variety of sensory , nervous and reproductive tissues ( figure 3a ) . notably , the aplysia ir25a orthologue is predominantly expressed in the olfactory organs , the rhinophore and oral tentacle [ 21 ] . two other aplysia - specific ir genes , ir214 and ir217 , are expressed in the rhinophore and oral tentacle , respectively , and not detected in other tissues , except for the large hermaphroditic duct ( ir214 ) and skin ( ir217 ) . five additional irs are also expressed in the oral tentacle , but displayed broader tissue expression in skin and the central nervous system ; both of these tissues are likely to contain other types of chemosensory cells [ 22 ] – [ 23 ] . expression of two ir genes , ir209 and ir213 , was not detected in this analysis ( data not shown ) .
the molecules that mediate innate immunity are encoded by relatively few genes and exhibit broad specificity . detailed annotation of the pacific oyster ( crassostrea gigas ) genome , a protostome invertebrate , reveals large - scale duplication and divergence of multigene families encoding molecules that effect innate immunity . transcriptome analyses indicate dynamic and orchestrated specific expression of numerous innate immune genes in response to experimental challenge with pathogens , including bacteria , and a pathogenic virus . variable expression of individual members of the multigene families encoding these genes also occurs during different types of abiotic stress ( environmentally - equivalent conditions of temperature , salinity and desiccation ) . multiple families of immune genes are responsive in concert to certain biotic and abiotic challenges . individual members of expanded families of immune genes are differentially expressed under both biotic challenge and abiotic stress conditions . members of the same families of innate immune molecules also are transcribed in developmental stage - and tissue - specific manners . an integrated , highly complex innate immune system that exhibits remarkable discriminatory properties and responses to different pathogens as well as environmental stress has arisen through the adaptive recruitment of tandem duplicated genes . the co - adaptive evolution of stress and innate immune responses appears to have an ancient origin in phylogeny .
( a ) table of antennal ir orthologous groups in the indicated protostome species . a shaded square signifies the presence of at least one representative gene in a species . figures within a square indicate the existence of multiple functional copies ; the “p” suffix indicates the number of pseudogene copies . we considered as pseudogenes only those with frameshift or internal stop codons inside conserved domains due to the difficulty in accurately annotating the termini of these sequences . ( b ) phylogenetic relationships of the genes shown in ( a ) . each colour represents a group of orthologous sequences . the sequences were aligned with probcons and the tree was built with raxml under the wag model of substitution with 1000 bootstrap replicates . the scale bar represents the expected number of substitutions per site . ( c ) phylogenetic relationships between iglurs and all ir25a orthologues , excluding low quality or short gene annotations . the tree was built with raxml under the wag model of substitution , with 1000 bootstrap replicates . bootstrap values for selected branches are shown as percentages . the scale bar represents the expected number of substitutions per site . ( d ) top : map of intron positions in an alignment of eight ir25a orthologues , 12 ampa / kainate receptors and 13 nmda receptors ( see dataset s3 for alignment file ) . coloured boxes illustrate introns whose positions and phases are conserved in at least one member of two different subgroups . empty coloured boxes indicate introns conserved in position , but not in phase . bottom : phylogram based on the position of introns in the same subset of sequences as above . the scale bar represents the number of non - conserved intron positions . ( e ) rt - pcr analysis of antennal ir gene expression of orthologous genes ( except dmelir75a and amelir75u , which are paralogous genes ) in d . melanogaster and a . mellifera tissues . control rt - pcr products for comparative analysis of gene expression correspond to the ribosomal genes rps7 ( d . melanogaster ) and rps8 ( a . mellifera ) . all rt - pcr products were sequenced to confirm their identity .
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the majority of coelomate invertebrates develop as protostomes (
first mouth
) in which the oral end of the animal develops from the first developmental opening , the blastopore . in the deuterostomes (
second mouth
: cf . deuteronomy ,
second book of the law
) , including echinodermata and the ancestors of the chordata , the oral end of the animal develops from the second opening on the dorsal surface of the animal ; the blastopore becomes the anus .
the deuterostomes also show indeterminate development , in which each of the cells of the eight - cell embryo if separated remain capable of developing as complete organisms . this contrasts to determinate development , in which the development fate of each cell in the adult organism has already been determined .
figure ©1999 campbell et al . ; text © 2017 by steven m . carr
any member of the lower invertebrate phyla in which the mouth appears before the anus during development , cleavage is spiral and determinate , and the coelom forms as a splitting of the mesoderm .
urltoken unabridged based on the random house unabridged dictionary , © random house , inc . 2018
any of a major group of animals defined by its embryonic development , in which the first opening in the embryo becomes the mouth . at this stage of development , the later specialization of any given embryonic cell has already been determined . protostomes are one of the two groups of animals having a true body cavity ( coelom ) and are believed to share a common ancestor . they include the mollusks , annelids , and arthropods .
the american heritage® science dictionary copyright © 2011 . published by houghton mifflin harcourt publishing company . all rights reserved .
the term protostomia ( from the greek
proto ,
meaning first , and
stoma ,
meaning mouth ) was coined by the biologist karl grobben in 1908 . it distinguishes a group of invertebrate animals based upon the fate of the blastopore ( the first opening of the early digestive tract ) during embryonic development . animals in which the blastopore becomes the mouth are called protostomes ; those in which the mouth develops after the anus are called deuterostomes ( from the greek
deutero ,
meaning second , and
stoma ,
meaning mouth ) .
protostomia and deuterostomia are considered super - phyletic taxa , each containing a variety of animal phyla . traditionally , the protostomes include the annelida , arthropoda , and mollusca , and the deuterostomes comprise the echinodermata and chordata . grobben was not the first biologist to recognize the distinction between these two groups , but he was the first to place importance on the fate of the blastopore as a major distinguishing criterion . historically , the two groups are distinguished by the following criteria :
cleavage pattern refers to the process of cell division from one fertilized cell , the zygote , into hundreds of cells , the embryo . in protostomes , the developing zygote undergoes spiral cleavage , a process in which the cells divide at a 45° angle to one another due to a realignment of the mitotic spindle . the realignment of the mitotic spindle causes each cell to divide unequally , resulting in a spiral displacement of small cells , the micromeres , that come to sit atop the border between larger cells , the macromeres . another superphyletic term used to describe animals with spiral cleavage is spiralia . spiral cleavage is also called determinate cleavage , because the function of the cells is determined early in the cleavage process . the removal of any cell from the developing embryo will result in abnormal development , and individually removed cells will not develop into complete larvae .
in deuterostomes , the zygote undergoes radial cleavage , a process in which the cells divide at right angles to one another . radial cleavage is also known as indeterminate cleavage , because the fate of the cells is not fixed early in development . the removal of a single cell from a developing embryo will not cause abnormal development , and individually removed cells can develop into complete larvae , producing identical twins , triplets , and so forth .
the fate of the blastopore has classically been used as the defining characteristic of protostomes and deuterostomes . in protostomes , the blastopore develops into the mouth , and the anus develops from an opening later in development . in deuterostomes , the blastopore develops into the anus , and the mouth develops secondarily .
mesoderm and coelom formation are intimately tied together during development . in protostomes , the mesoderm originates from a pair of cells called mesentoblasts ( also called
4d cells ) next to the blastopore , which then migrate into the blastocoel , the internal cavity of the embryo , to become various internal structures . in coelomates , the mesentoblasts hollow out to become coeloms , cavities lined by a contractile peritoneum , the myoepithelium . in protostomes , the process of coelom formation is called schizocoely . in deuterostomes , the mesoderm originates from the wall of the archenteron , an early digestive tract formed from endoderm . the archenteron pouches out to form coelomic cavities , in a process called enterocoely .
protostomia and deuterostomia are also characterized by different larvae . in most protostomes , the larval type is a trochophore , basically defined by the presence of two rings of multiciliated cells ( prototroch and metatroch ) surrounding a ciliated zone around the mouth . most deuterostomes have a dipleurula - type larva , defined by the presence of a field of cilia ( monociliated cells ) surrounding the mouth .
for more than a century , biologists have divided the bilateral animals into two main lineages ( the diphyletic origin of bilateria ) , the most well known of which is the protostomia / deuterostomia split . similar divisions include the zygoneura / ambulacralia - chordonia split proposed by the german invertebrate embryologist hatschek in 1888 , the hyponeuralia / epineuralia split proposed by the french zoologist cuenot in 1940 , and a gastroneuralia / notoneuralia split proposed by the german zoologist ulrich in 1951 , among others . these divisions often emphasized different developmental and adult features , thereby leading to different names and hypotheses about animal relationships . although none of these groups have been granted formal taxonomic rank ( for example , as a subkingdom or superphylum ) by the international code of zoological nomenclature , the names nevertheless remain active in the literature .
the evolutionary origin of the protostomia , and of the groups it includes , remains a major challenge to modern biologists . although proof of the monophyly of the protostomia is elusive , many of the phyla are clearly related , and make up clades that some biologists consider monophyletic . for example , in 1997 aguinaldo et al . proposed the establishment of two clades within the protostomia based on molecular sequence data : ecdysozoa ( the molting animals , including the arthropoda , nematoda , priapulida , and tardigrada ) , and lophotrochozoa ( the ciliated animals , including the annelida , echiura , and sipuncula ) . biologists continue to debate these hypotheses and test them with independent biochemical , developmental , molecular , and morphological data .
brusca , r . c . , and g . j . brusca . invertebrates , 2nd ed . new york : sinauer associates , 2003 .
nielsen , c . animal evolution : interrelationships of the living phyla , 2nd ed . new york : oxford university press , 2001 .
aguinaldo , a . m . a . , et al .
evidence for a clade of nematodes , arthropods and other moulting animals .
nature 387 ( 1997 ) : 483–491 .
løvtrup , s .
validity of the protostomia - deuterostomia theory .
systematic zoology 24 ( 1975 ) : 96–108 .
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a multicellular organism whose mouth develops from a primary embryonic opening , such as an annelid , mollusc , or arthropod .
‘in the complementary category , the protostomes , which includes annelids and arthropods , the blastopore develops into the mouth . ’
‘these animals are all protostomes - the mouth develops before the anus in the young embryo - and they have long been recognized as belonging together as a group . ’
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any of numerous invertebrate animals of the group protostomia , in which the mouth develops from the first opening in the embryo and the anus develops later , and including the mollusks , annelids , and arthropods .
any invertebrate in which the mouth appears before the anus during development , cleavage is spiral and determinate , and the coelom forms as a splitting of the mesoderm .
evolution ,
philosophical transactions of the royal society london b 347 ( 1995 ) : 305 - 58 .
invertebrate branch , including the phylum arthropoda ( jointed leg animals ) , annelida ( segmented worms ) , and mollusca ( bivalves , snails , and cephalopods ) .
and vertebrate genomes contain both types , a fact recognized by hibino et al .
lineages , help expand perspectives in gamete biology and add to our current knowledge of these topics , which is predominantly based on studies of mammals .
stereotypes , many species display modifications of the spiralian cleavage pattern , diverse modes of gastrulation , and of course varied larval forms .
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school of biological sciences , the university of queensland , st . lucia , queensland , australia
ionotropic glutamate receptors ( iglurs ) are a family of cell surface proteins best known for their role in allowing neurons to communicate with each other in the brain . we recently discovered a variant class of iglurs in the fruit fly ( drosophila melanogaster ) , named ionotropic receptors ( irs ) , which function as olfactory receptors in its “nose , ” prompting us to ask whether iglur / irs might have a more general function in detection of environmental chemicals . here , we have identified families of irs in olfactory and taste sensory organs throughout protostomes , one of the principal branches of animal life that includes snails , worms , crustaceans , and insects . our findings suggest that this receptor family has an evolutionary ancient function in detecting odors and tastants in the external world . by comparing the repertoires of these chemosensory irs among both closely - and distantly - related species , we have observed dynamic patterns of expansion and divergence of these receptor families in organisms occupying very different ecological niches . notably , many of the receptors we have identified are in insects that are of significant harm to human health , such as the malaria mosquito . these proteins represent attractive targets for novel types of insect repellents to control the host - seeking behaviors of such pest species .
editor : david l . stern , princeton university , howard hughes medical institute , united states of america
copyright : © 2010 croset et al . this is an open - access article distributed under the terms of the creative commons attribution license , which permits unrestricted use , distribution , and reproduction in any medium , provided the original author and source are credited .
funding : sfc was supported by a university of queensland fellowship and the australian research council , rr and db by roche research foundation fellowships , and vc by an embo short - term fellowship and a boehringer ingelheim foundation fellowship . research in rb ' s laboratory is funded by the university of lausanne , a european research council starting independent researcher grant , and the swiss national science foundation . the funders had no role in study design , data collection and analysis , decision to publish , or preparation of the manuscript .
ionotropic glutamate receptors ( iglurs ) are a conserved family of ligand - gated ion channels present in both eukaryotes and prokaryotes . by regulating cation flow across the plasma membrane in response to binding of extracellular glutamate and related ligands , iglurs represent an important signalling mechanism by which cells modify their internal physiology in response to external chemical signals .
iglurs have originated by combination of protein domains originally encoded by distinct genes ( figure 1a ) [ 1 ] – [ 2 ] . an extracellular amino - terminal domain ( atd ) is involved in assembly of iglur subunits into heteromeric complexes [ 3 ] . this precedes the ligand - binding domain ( lbd ) , whose two half - domains ( s1 and s2 ) form a “venus flytrap” structure that closes around glutamate and related agonists [ 4 ] . separating s1 and s2 in the primary structure is the ion channel pore , formed by two transmembrane segments and a re - entrant pore loop [ 5 ] . s2 is followed by a third transmembrane domain of unknown function and a cytosolic carboxy - terminal tail .
( a ) top : histogram showing the mean conservation index ( number of conserved physico - chemical properties ) [ 74 ] , [ 91 ] for 50 amino acid column - blocks of aligned d . melanogaster iglurs and irs , illustrating the higher conservation of the c - terminal region . the protein domain organisation of iglurs / irs is shown in cartoon form above the histogram and in linear form below it . bottom : illustration of the three pfam domains present in iglurs and irs . ir8a and ir25a contain the pfam domain corresponding to the iglur atd . ir21a , ir40a , ir64a and ir93a also contain long n - termini ( ∼400 amino acids ) but these have extremely low primary structural similarity to the atd . all other irs have much shorter n - terminal regions ( ∼200 amino acids ) that lack homology to the atd or other protein domains . ( b ) histogram of the number of non - nmda ( red ) , nmda ( yellow ) and ir ( blue ) sequences identified in the indicated eukaryotic species . an unscaled tree showing the phylogenetic relationships between these species is illustrated on the left .
animal iglurs have been best characterised for their essential roles in synaptic transmission as receptors for the excitatory neurotransmitter glutamate [ 1 ] , [ 6 ] . three pharmacologically and molecularly distinct subfamilies exist , named after their main agonist : α - amino - 3 - hydroxy - 5 - methyl - 4 - isoxazolepropionic acid ( ampa ) , kainate and n - methyl - d - aspartate ( nmda ) . ampa receptors mediate the vast majority of fast excitatory synaptic transmission in the vertebrate brain , while kainate receptors have a subtler modulatory role in this process . nmda receptors require two agonists for activation , glutamate and glycine , and function in synaptic and neuronal plasticity . representatives of these iglur subfamilies have been identified across vertebrates [ 7 ] , as well as invertebrates , such as the fruit fly drosophila melanogaster , the nematode worm caenorhabditis elegans and the sea slug aplysia californica [ 8 ] – [ 10 ] .
while most iglurs have exquisitely tuned synaptic functions , identification of iglur - related genes in prokaryotic and plant genomes provided initial indication of more diverse roles for this class of ion channel . a bacterial glutamate receptor , glur0 , was first characterised in the cyanobacterium , synechocystis pcc6803 [ 11 ] . glur0 conducts ions in response to binding of glutamate and other amino acids in vitro , suggesting a potential function in extracellular amino acid sensing in vivo . the flowering plant arabidopsis thaliana has 20 iglur - related genes , named glrs [ 12 ] – [ 13 ] . genetic analysis of one receptor , glr3 . 3 , has implicated it in mediating external amino acid - stimulated calcium increases in roots [ 14 ] .
we recently described a family of iglur - related proteins in d . melanogaster , named the ionotropic receptors ( irs ) [ 15 ] . several lines of evidence demonstrated that the irs define a new family of olfactory receptors . first , the ir lbds are highly divergent and lack one or more residues that directly contact the glutamate ligand in iglurs . second , several irs are expressed in sensory neurons in the principal d . melanogaster olfactory organ , the antenna , that do not express members of the other d . melanogaster chemosensory receptor families , the odorant receptors ( ors ) and gustatory receptors ( grs ) [ 16 ] . third , ir proteins localise to the ciliated endings of these sensory neurons and not to synapses [ 15 ] . finally , mis - expression of an ir in an ectopic neuron is sufficient to confer novel odour - evoked neuronal responses , providing direct genetic evidence for a role in odour sensing [ 15 ] .
the identification of the irs as a novel family of olfactory receptors in d . melanogaster provides a potential link between the well - characterised signalling activity of iglurs in glutamate neurotransmitter - evoked neuronal depolarisation and a potentially more ancient function of this family in environmental chemosensation . in this work , we have combined comparative genomics , molecular evolutionary analysis and expression studies to examine the evolution of the irs . four principal issues are addressed : first , when did olfactory irs first appear ? are they a recent acquisition as environmental chemosensors in d . melanogaster , or do they have earlier origins in insect or deeper animal lineages ? second , what is the most recent common ancestor of ir genes ? do they derive from ampa , kainate or nmda receptors , or do they represent a distinct subfamily that evolved from the ancestral animal iglur ? third , what mechanisms underlie the expansion and diversification of this multigene family ? finally , do irs function only as olfactory receptors or are they also involved in other sensory modalities ? through answers to these questions , we sought insights into ir evolution in the context of the origins of iglurs , the appearance and evolution of other chemosensory receptor repertoires and the changing selective pressures during animal diversification and exploitation of new ecological niches .
iglurs and irs are characterised by the presence of a conserved ligand - gated ion channel domain ( the combined pfam domains pf10613 and pf00060 [ 17 ] ) ( figure 1a ) . all iglurs additionally contain an atd ( pfam domain pf01094 ) , which is discernible , but more divergent , in only two d . melanogaster irs , ir8a and ir25a . most irs have only relatively short n - terminal regions preceding the lbd s1 domain ( figure 1a ) . to identify novel iglur / ir - related genes , we therefore constructed a hidden markov model ( hmm ) from an alignment of the conserved iglur / ir c - terminal region , which is specific to this protein family . in combination with exhaustive blast searches , we used this hmm to screen raw genomic sequences and available annotated protein databases of 32 diverse eukaryotic species and 971 prokaryotic genomes ( see materials and methods and table s2 in supporting information ) . these screens identified all previously described eukaryotic iglurs and all d . melanogaster irs , as well as 23 prokaryotic iglurs . novel sequences were manually reannotated and classified by sequence similarity , phylogenetic analysis and domain structure as either non - nmda ( i . e . ampa and kainate ) or nmda subfamily iglurs , or irs ( figure 1b , table s3 , and datasets s1 and s2 ) . like d . melanogaster irs , newly annotated irs have divergent lbds that lack some or all known glutamate - interacting residues , supporting their distinct classification from iglurs .
iglurs are widespread in eukaryotes , present in all analysed metazoa ( except the sponge , amphimedon queenslandica [ 18 ] ) and plantae , but absent in unicellular eukaryotes ( figure 1b , table s3 , and datasets s1 and s2 ) . analysis of iglur subfamilies on the eukaryotic phylogeny suggests that nmda receptors may have appeared after non - nmda receptors , as we identified them in eumetazoa but not in the placozoan trichoplax adhaerens . further support for this conclusion will require additional genome sequences . one member of the eumetazoa , the sea urchin strongylocentrotus purpuratus , may have secondarily lost nmda receptors . different species contain distinct numbers of each iglur subfamily : vertebrates , for example , have more nmda receptor subunits than invertebrates .
notably , irs were identified throughout protostomia , encompassing both ecdysozoa ( e . g . nematodes and arthropods ) and lophotrochozoa ( e . g . molluscs and annelids ) ( figure 1b , table s3 , and datasets s1 and s2 ) . there is substantial variation in the size of the ir repertoire , from three in c . elegans to eighty - five in the crustacean daphnia pulex . amongst insects , diptera ( i . e . flies and mosquitoes ) generally had a larger number of irs than other species . we did not identify irs in deuterostomia , cnidaria or placozoa .
consistent with its conservation in protostomia , ir25a is the ir with the most similar primary sequence to iglurs , suggesting that it is the ir gene most similar to the ancestral ir . analysis of the phylogenetic relationship of ir25a and eukaryotic iglurs locates it clearly together with the animal iglur family , in the non - nmda receptor clade ( figure 2c ) . to substantiate this conclusion , we asked whether the ir25a gene structure resembles more closely that of nmda or non - nmda receptors . intron positions and numbers are extremely variable across ir25a orthologues , with multiple cases of intron loss , gain and putative intron sliding events by a few nucleotides ( figure 2d ) . nevertheless , we identified eight intron positions that are conserved between at least subsets of ir25a orthologues and d . melanogaster non - nmda receptor genes , some of which may represent intron positions present in a common ancestral gene . by contrast , only a single intron that was conserved in position ( but not in phase ) was identified between dmelir25a ( but not other ir25a orthologues ) and dmelnmdar1 ( figure 2d ) . a phylogram of intron positions in ir25a , non - nmda and nmda sequences reveals greater similarity of ir25a intron positions to those of non - nmda receptors than nmda receptors ( figure 2d ) . together , these observations support a model in which ir25a evolved from a bilaterian non - nmda receptor gene .
the conserved d . melanogaster irs encompass the entire subset of its ir repertoire that is expressed in the antenna [ 15 ] . moreover , evidence for antennal expression of the three additional genes , dmelir41a , dmelir60a and dmelir68a , has been obtained by reverse transcription ( rt ) - pcr analysis , although we have not yet been able to corroborate this by rna in situ hybridisation ( data not shown ) . these combined phylogenetic and expression properties led us to designate this subfamily of receptors the “antennal irs” .
we examined whether antennal expression of this subfamily of irs is conserved outside d . melanogaster by performing a series of rt - pcr experiments on the honey bee , apis mellifera , for all six putative antennal ir orthologues : ir8a , ir25a , ir68a , ir75u , ir76b and ir93a ( see materials and methods for the nomenclature of newly - identified irs ) . as in d . melanogaster , we could reproducibly amplify all of these bee genes from antennal rna preparations but not in control brain rna , except for amelir68a and amelir75u , which are also detected in the brain ( figure 2e ) . thus , antennal expression of this subgroup of irs is conserved across the 350 million years separating dipteran and hymenopteran insect orders [ 19 ] , and therefore potentially in all insects .
( a ) top : schematic representation of aplysia , illustrating the location of selected sensory , neuronal and reproductive tissues used for rna isolation and rt - pcr ( adapted from [ 21 ] ) . the central nervous system samples comprised pooled cerebral , pleural , buccal , pedal and abdominal ganglia . the skin samples were taken from the side of the head . bottom : rt - pcr analysis of aplysia ir gene expression from the indicated species and tissues . only rhinophores from a . californica ( acal ) were tested due to limited availability of animals , while rhinophore and other tissues were examined for the closely related species a . dactylomela ( adac ) [ 92 ] . nucleotide sequence identity of ir orthologues between these species is > 85 % . control rt - pcr corresponds to β - actin . ( b ) schematic of aplysia rhinophore showing the approximate location of the field of views of the rhinophore groove olfactory tissue in ( c–e ) . ( c , d ) rna in situ hybridisation on a . dactylomela rhinophore sections using a digoxigenin - labelled antisense rna probe for adacir25a . micrographs reveal ir25a expression ( blue ) in small clusters of cells of a characteristic neuronal morphology close to the sensory epithelial surface . higher magnifications of specific cellular staining ( arrowhead ) are shown in the insets . the scale bars represent 100 µm . ( e ) control rna in situ hybridisation on an a . dactylomela rhinophore section with a digoxigenin - labelled sense riboprobe for adacir25a . no signal is apparent . the scale bar represents 100 µm .
to further characterise aplysia ir25a , we analysed its spatial expression in the mature a . dactylomela rhinophore by rna in situ hybridisation . an antisense probe for adacir25a labels a small number of cells in rhinophore cryosections . their size and morphology is typical of neurons , although we lack an unambiguous neuronal marker to confirm this identification ( figure 3b–3d ) . these cells are found either singly or in small clusters adjacent or close to the sensory epithelial surface in the rhinophore groove , in a similar position to cells expressing other types of chemosensory receptors [ 21 ] . a control sense riboprobe showed no specific staining ( figure 3e ) . together , these results are consistent with at least some of these molluscan irs having a chemosensory function .
the expression of putative ir25a orthologues has previously been reported in two other protostomia . an ir25a - related gene from the american lobster , homarus americanus , named oet - 07 , is specifically expressed in mature olfactory sensory neurons [ 24 ] – [ 25 ] . in c . elegans , a promoter reporter of the ir25a orthologue , glr - 7 , revealed expression in a number of pharyngeal neurons [ 9 ] , which might have a role in food sensing [ 26 ] . while both crustacean and nematode genes were classified in these studies as iglurs , there is no evidence that they act as canonical glutamate receptors , and we suggest that they fulfil instead a chemosensory function .
the antennal ir subfamily accounts for only a small fraction of the ir repertoire in most analysed insects and only 1–2 genes in other protostomia . the remaining majority of ir sequences are - amongst the genomes currently available - largely species - specific , with low amino acid sequence identity ( as little as 8 . 5 % ) with other ir genes in either the same or different species . we refer to this group of genes here as the “divergent irs” . dipteran insects have particularly large expansions of divergent irs ( figure 1b ) . phylogenetic analysis revealed no obvious orthologous relationships of these genes either between d . melanogaster and mosquitoes or amongst the three mosquito species ( aedes aegypti , culex quinquefasciatus and anopheles gambiae ) ( figure 4 ) . instead , this subfamily of irs displays a number of species - specific clades , perhaps reflective of the distinct ecological niches of these insects .
phylogenetic tree of all iglurs and irs from d . melanogaster ( blue ) , a . aegypti ( green ) , c . quinquefasciatus ( orange ) and a . gambiae ( red ) . sequences were aligned with probcons and the tree was built with raxml under the wag model of substitution , with 500 bootstrap replicates . the scale bar represents the expected number of substitutions per site . note that due to the high divergence and number of sequences analysed , bootstrap values in several of the most internal nodes are extremely low and the position of certain large clades of ir genes on the tree are distinct from trees in other figures .
by contrast to antennal irs , divergent ir expression has not been detected in d . melanogaster olfactory organs [ 15 ] , leading us to test whether these genes are expressed in other types of chemosensory tissue . as endogenous transcripts of non - olfactory chemosensory genes , such as grs , are difficult to detect [ 27 ] – [ 28 ] , we employed a sensitive transgenic approach to investigate divergent ir expression . we transformed flies with constructs containing putative promoter regions for these genes upstream of the yeast transcription factor gal4 and used these “driver” transgenes to induce expression of a gal4 - responsive uas - mcd8 : gfp fluorescent reporter [ 29 ] . we sampled divergent irs from several distinct clades , including ir7a , ir11a , ir52b , ir56a and ir100a ( figure 4 ) . all ir promoter - gal4 constructs were inserted in the same genomic location using the phic31 integrase system [ 30 ] , eliminating transgene - specific position effects on expression resulting from their site of integration .
expression of three of these divergent ir reporters was observed in highly selective populations of neurons in distinct gustatory organs ( figure 5a ) . in the adult , ir7a is expressed in at least eleven neurons in the labellum , a sense organ involved in peripheral taste detection ( figure 5b ) [ 31 ] . two reporters labelled neurons in internal sense organs in the pharynx : ir11a is expressed in one neuron in the ventral cibarial sense organ and ir100a is expressed in two neurons in the dorsal cibarial sense organ ( figure 5c and 5d ) . these internal pharyngeal neurons are thought to play a role in assessment of ingested food prior to entry into the main digestive system [ 16 ] . expression was not detected in any other neurons or other cell types in the adult head ( data not shown ) , although we cannot exclude expression in other regions of the body . ir52b and ir56a reporters were not detected in these experiments .
immunofluorescence with anti - gfp ( green ) and anti - ir25a ( magenta ) antibodies ( overlaid on bright - field images ) on whole - mount tissues from animals expressing a membrane targeted gfp reporter transgene ( uas - mcd8 : gfp ) under the control of the indicated ir promoter - gal4 driver transgenes . the scale bars represent 20 µm . ( a ) schematic of the adult d . melanogaster proboscis showing the location of the field of views in ( b–d ) . dcso : dorsal cibarial sense organ , vcso : ventral cibarial sense organ . ( b ) ir7a - gal4 drives expression of mcd8 : gfp in the labellum . ( c ) ir11a - gal4 drives expression of mcd8 : gfp in the vcso . ( d ) ir100a - gal4 drives expression of mcd8 : gfp in the dcso . ( e ) schematic of the d . melanogaster larval head showing the location of the field of views in ( f–h ) . to : terminal organ , dps : dorsal pharyngeal sense organ , pps : posterior pharyngeal sense organ . ( f ) ir7a - gal4 drives expression of mcd8 : gfp in the to . ( g ) ir11a - gal4 drives expression of mcd8 : gfp in the dps . ( h ) ir100a - gal4 drives expression of mcd8 : gfp in the pps .
we also examined expression of these reporters at an earlier stage in the d . melanogaster life cycle , third instar larvae , which display robust gustatory responses [ 16 ] . the same three ir reporters were exclusively detected in unique bilaterally - symmetric larval gustatory organs : ir7a was expressed in two neurons in the terminal organ at the periphery , ir11a in a single neuron in the ventral pharyngeal sense organ and ir100a in two neurons in the posterior pharyngeal sense organ ( figure 5e–5h ) . notably , all of these neurons in both adult and larval tissues ( except for a single ir7a - expressing cell in the terminal organ ) co - express ir25a , as revealed by a specific antibody against this receptor ( figure 5 ) [ 15 ] . ir25a is also expressed in several other cells in each of the gustatory organs , which may express other divergent irs not examined here . together these results support a role for divergent irs as taste receptors in distinct taste organs and stages of the d . melanogaster life cycle .
to obtain more detailed insights into the processes underlying the expansion and diversification of ir repertoires , we investigated their evolution over a shorter timescale by comparative analysis of d . melanogaster with 11 additional sequenced drosophilid species [ 32 ] – [ 33 ] . the last common ancestor of these drosophilids is estimated to have existed 40 million years ago [ 34 ] , by contrast to the ∼250 million years since the last common ancestor of d . melanogaster and the mosquito a . gambiae [ 35 ] . certain species may have diverged much more recently , such as d . simulans and d . sechellia , whose last common ancestor may have existed only 250 , 000 years ago [ 36 ] .
we used d . melanogaster sequences as queries in exhaustive blast searches of the drosophilid genomes . retrieved sequences were manually reannotated to unify gene structure predictions across species and , in some cases , genes were partially resequenced to close sequence gaps or verify them as pseudogenes ( see materials and methods , table s3 , and datasets s1 and s2 ) . although predicted full - length gene sequences could be annotated for most genes , 28 sequences remain incomplete - but assumed in further analysis to be functional - because of a lack of sequence data or difficulty in precise annotation of exons in divergent regions of these genes . of the 926 drosophilid sequences identified ( including those of d . melanogaster ) , 49 genes were classified as pseudogenes because they consisted of only short gene fragments or contained frameshift mutations and / or premature stop codons . we clustered all genes into orthologous groups by examining their sequence similarity , phylogenetic relationships and , in the case of ir47a , ir47b , ir47c , ir56e and ir60f , their micro - syntenic relationships ( table s1 and figure 6 ) . for drosophilid species that are most distant from d . melanogaster , definition of precise orthologous relationships was not always possible , particularly for groups of closely related ir genes ( e . g . ir52a–f , ir60b–f ) ( table s1 ) . orthologous groups were named after their d . melanogaster representatives or a logical variant in groups where no d . melanogaster gene was identified ( see materials and methods ) .


it is found in california .

the wingspan is 8 – 9 mm .

bucculatrix albaciliella

bucculatrix albaciliella is a moth in the bucculatricidae family , first described by braun in 1910 . it is found in california .
kari pihlaviita added the finnish common name
tarhatöyhtökoi
to
bucculatrix bechsteinella
.
kari pihlaviita added the finnish common name
koivutöyhtökoi
to
bucculatrix demaryella duponchel
.
kari pihlaviita added the finnish common name
lehmustöyhtökoi
to
bucculatrix thoracella thunberg 1794
.
kari pihlaviita added the finnish common name
leppätöyhtökoi
to
bucculatrix cidarella zeller 1839
.
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the wingspan is 8–9 mm . the forewings are snowy white , marked with pale ocherous spots and streaks . the hindwings are white , faintly tinged with grey . adults have been recorded on wing in april .