woolly aphids ( subfamily : eriosomatinae ) are sucking insects that live on plant fluids and produce a filamentous waxy white covering which resembles cotton or wool .

the adults are winged and move to new locations where they lay egg masses .

the nymphs often form large cottony masses on twigs , for protection from predators .

they occur throughout the northern hemisphere .

many of the numerous species of woolly aphids have only one host plant species , or alternating generations on two specific hosts .

the woolly apple aphid , eriosoma lanigerum is a widespread pest of fruit trees , feeding principally on apple , but also , pears , hawthorn , ash , alders , elms and oaks .

in flight they have been described as looking like " flying mice " , and are given nicknames like " angel flies " , " fluff bugs " , " fairy flies " , and " ash bugs " " snow bugs " , " fluffer fairy " , " poodle fly " , " fluffy gnats " .

eriosomatinae

morphological phylogeny of gall - forming aphids of the tribe eriosomatinae ( aphididae : eriosomatinae ) .
another significant difference with other species of eriosomatinae is the shape and position of rectal bladder . in
les according to the 3 tribes of eriosomatinae and the out - groups . e genbank accession n
evolutionary relationships of pemphigus and allied genera ( hemiptera : aphididae : eriosomatinae ) and . . .
look at wing venation for further clues in distinguishing among these . below is a pemphigus ( eriosomatinae ) for comparison .
honeydew from root aphids ( aphidae : eriosomatinae ) is an important part of the diet of the subterranean ant lasius nearcticus . new york , usa
type genus : eriosoma leach , 1818 ; origin of priority for family - group names not recorded in database . first use as eriosomatinae by baker , a . c . , 1920 .
many woolly aphids are in subfamily eriosomatinae . however , there are several exceptions ( and according to andrew jensen ,
there are examples of wax - covered aphids in almost all subfamilies .
) :
. . . in addition , in all higher - level phylogenetic studies of aphididae that have been conducted , neither aphid data ( vondohlen & moran 2000 ; ortiz - rivas et al . 2004 ; ortizrivas & mart ınez - torres 2010 ) nor buchnera data ( nov akov a et al . 2013 ) supported the eriosomatinae as a monophyletic clade . its monophyly was also not corroborated in the morphological cladistic study ( zhang & chen 1999 ) and molecular phylogenetic analyses of eriosomatinae ( zhang & qiao 2008 ; li et al . 2014 ) . the subfamily eriosomatinae has been traditionally recognized as a monophylum based on the synapomorphy of sexuales apterous , dwarfish and lacking rostrum ( heie 1980 ; zhang et al . 1999a ) . . . .
. . . in addition , in all higher - level phylogenetic studies of aphididae that have been conducted , neither aphid data ( von dohlen & moran 2000 ; ortiz - rivas et al . 2004 ; ortizrivas & mart ınez - torres 2010 ) nor buchnera data ( nov akov a et al . 2013 ) supported the eriosomatinae as a monophyletic clade . its monophyly was also not corroborated in the morphological cladistic study ( zhang & chen 1999 ) and molecular phylogenetic analyses of eriosomatinae ( zhang & qiao 2008 ; li et al . 2014 ) . the subfamily eriosomatinae has been traditionally recognized as a monophylum based on the synapomorphy of sexuales apterous , dwarfish and lacking rostrum ( heie 1980 ; zhang et al . 1999a ) . . . .
) , except of eriosomatinae , which were not studied in this respect . the existing differences concern mainly the lengths of particular segments of the tract . due to the lack of filter chamber , the adhering of the loops of crenated intestine to the ventral wall of the stomach may play the role of osmoregulation , as hypothesized for the pea aphid ( rhodes et al .
eriosomatinae , the gall - forming aphid subfamily , traditionally consists of 3 tribes , eriosomatini , pemphigini , and fordini . however , the phylogenetic relationships among these tribes remain controversial , which has made it difficult to conduct further investigation regarding the evolution of galls and host alternations in this group . we analyzed the molecular phylogeny of the subfamily eriosomatinae , combining sequences from 2 mitochondrial genes ( coi and coii ) and 2 nuclear genes ( ef - 1 alpha and lwo ) . the reconstructions were implemented based on single - gene and multigene datasets through 3 different reconstructing algorithms , respectively ; analyses with 5 different out - groups were also conducted . results revealed a large paraphyletic clade , in which there were 4 out - groups clustering between eriosomatini and the other 2 tribes . however , the monophyly of the 3 tribes was well supported by the obtained trees , respectively .
. . . but the relationships among genera were not well supported with the position of the north american melaphis poorly resolved . using the mitochondrial coi and coii and the nuclear ef - 1a and lwo , li et al . ( 2014 ) constructed the relationships of subfamily eriosomatinae , sampling six genera of the melaphidina aphids . their results supported the monophyly of melaphidina , but the relationships among the melaphidina genera were only well supported by the bayesian posterior probabilities ( pp ) without strong bootstrap support ( bs ) . . . .
as a conclusion , it must be stated that the alimentary tract of apterous viviparous female of anholocyclic population of g . utricualria consists of all anatomical organs typical of aphid subfamily eriosomatinae and constitutes a continuous tube . it is unlikely that g . setulosa may have differently built alimentary tract , since existing data on its foregut and hindgut fully agree with that of presented study on g . utricularia . however , taking into account complicated life cycle and heteroecy of geoica spp , . there is essential need for further studies to confirm whether the situation of geoica spp . ( and also various morphs of species in this genus ) is similar to that of g . utricularia or to that described by ponsen .
the analysis of b . aphidicola genes that follow an evolutionary pattern that agrees with the molecular clock hypothesis [ 12 , 13 ] can be used to estimate the divergence time between pairs of aphids . this is possible because two aphid species , acyrthosiphon pisum and schizaphis graminum belonging to two tribes of the subfamily aphidinae , have an estimated divergence time calibrated from their fossil record of 50 to 70 my [ 14 ] . in addition , using molecular data from complete b . aphidicola genomes available , pérez - brocal and coworkers calculated the divergence time of aphids belonging to subfamilies eriosomatinae ( baizongia pistaciae ) and lachninae ( cinara cedri ) [ 15 ] . based on morphological traits , the subfamilies eriosomatinae and lachninae have traditionally been considered very divergent . in fact , most phylogenetic hypotheses based both on morphological and molecular data consider the lachninae as a sister group of the aphidinae [ 11 , 16 ] . however , the position of this subfamily remains controversial , as recent phylogenies based on molecular sequences located the subfamily in a basal position [ 17 – 19 ] . here , we follow a genomic approach to deepen the evolutionary analyses and propose a phylogeny of the three subfamilies of aphids based on the genome sequence of their primary endosymbionts b . aphidicola . in addition , in order to detect if there is any selective effect related to the specific role of the genes , we also gave a closer look to the acceleration pattern of each functional category .
this species is widely distributed and are quite often encountered in yards . yet details of its life history are , as far as my literature searches can tell me , mostly undescribed : when does the switch to sexual reproduction happen ? how does the foundress feed or is she fed by her kids ? how and where do they overwinter ? how many asexual generations are completed within a year ? what kinds of predators and parasites plague the colonies ? some the species’ close kin ( other wooly and gall - forming insects in the subfamily eriosomatinae of the aphididae ) have ant - like life histories , forming solider castes within the colony . grylloprociphilus seems less specialized , but without detailed life history studies , we can’t say more .
since the establishment of the symbiosis between the ancestor of modern aphids and their primary endosymbiont , buchnera aphidicola , insects and bacteria have coevolved . due to this parallel evolution , the analysis of bacterial genomic features constitutes a useful tool to understand their evolutionary history . here we report , based on data from b . aphidicola , the molecular evolutionary analysis , the phylogenetic relationships among lineages and a comparison of sequence evolutionary rates of symbionts of four aphid species from three subfamilies . our results support previous hypotheses of divergence of b . aphidicola and their host lineages during the early cretaceous and indicate a closer relationship between subfamilies eriosomatinae and lachninae than with the aphidinae . they also reveal a general evolutionary pattern among strains at the functional level . we also point out the effect of lifecycle and generation time as a possible explanation for the accelerated rate in b . aphidicola from the lachninae .
existing literature data report the lack of stomach and crenated intestine in the aphid species geoica setulosa ( passerini , 1860 ) , a representative of subfamily eriosomatinae . this odd anatomical feature seemed remarkable , due to the presence of fully developed intestine in closely related genera and mutualistic relationship with ants of this genus . the study aimed at repeated anatomical research of geoica utricularia ( passerini 1856 ) , in order to confirm what seemed to be a generic feature . standard histological methods were applied , with addition of oblique light microscopy , fluorescence microscopy and confocal laser scanning microscopy . the results indicated the existence of a fully developed intestine , with broad sac - shaped stomach and loops of the crenated intestine . the general anatomy of the alimentary tract of g . utricularia resembles that of other representatives of the tribe fordini . also well - developed rectal gland is present , most probably playing a role in modifying the carbohydrate composition of excreted honeydew .
our phylogenetic analysis supports the presence of one clade clustering b . aphidicola bbp and bcc , and another clade consisting of b . aphidicola bap and bsg . this result is consistent with a panorama of a rapid evolutionary radiation of the main subfamilies of aphids , during the early cretaceous ( 144 - 100 mya ) , which seems concordant with previous proposals [ 3 ] . in addition , our evolutionary molecular data from b . aphidicola point out that aphids belonging to subfamilies eriosomatinae and lachninae share a common ancestor more closely related than compared to the members of subfamily aphidinae . if true , our data refute the traditional phylogenetic reconstructions that placed aphidinae and lachninae as a monophyletic group [ 11 ] . however , we do not have evidence to conclude whether , within the subfamily lachninae , tribes feeding on conifers are ancestral or more recent than those living on herbaceous angiosperms , since our analysis does not resolve which strain ( and thus which host aphid ) is basal compared to the others . to solve this point , it would be necessary to sequence the genome of a greater number of b . aphidicola strains , including members of the different tribes from the subfamily lachninae ( work in progress ) . this would allow us to establish the date of divergence between those tribes and , thus , try to relate this fact to the change of vegetal host in either direction .
regarding synonymous substitutions , when pairs of strains of b . aphidicola were compared based on the average number of synonymous substitutions per site ( ) , a greater accumulation was observed in the b . aphidicola bbp strain compared to bacteria from aphids of the subfamily aphidinae ( b . aphidicola bap and bsg ) , while the smallest value is found between the b . aphidicola bap and bsg strains [ 15 ] . however , if the temporary factor is considered , the rates of synonymous nucleotide substitutions per site and million years are greater in the endosymbionts from the aphidinae ( b . aphidicola bap and bsg strains ) , registering the b . aphidicola bcc strain the smallest values . these results demonstrate that the synonymous substitution rate in b . aphidicola is a variable character , yet the explanation for these divergent patterns is not obvious . as stated elsewhere [ 7 , 14 , 44 ] , these differences can be attributed to differences in the host’s life cycle , as well as ecological factors such as host - alternation and variations in the effective population size showed by the two members of the aphidinae subfamily compared to the other two aphid lineages . additionally a differential mutation rate per generation cannot be ruled out . for example , endosymbionts from aphids with short generation times can accumulate more synonymous mutations per million years ( case of the aphidinae ) than those with longer generation times , such as the eriosomatinae and the lachninae . future studies are required to understand the evolutionary processes driving these patterns .
in our efforts to understand the biology of soldier - producing aphids , we attempted to maintain them in the laboratory using a chemically defined artificial diet . the ability of 16 species from the subfamilies eriosomatinae and hormaphidinae , most of which are soldier - producing species , to survive on the artificial diet was examined . some species neither fed nor grew on the diet , whereas other species accepted the diet , grew to some extent , and managed to produce a small number of short - lived offspring . although they performed poorly on the diet in general , aphid performance was correlated with the stage in the life cycle and the developmental stadium in that aphids of the gall generation tended to accept the diet and survive on it , whereas aphids of the non - gall generation did not . also , old insects tended to perform better on the diet than young nymphs . notably , only one species , tuberaphis styraci , a gall - forming aphid that produces 2nd instar sterile soldier , showed good performance on the diet . insects collected from galls ( generation g0 ) survived on the diet , grew well , and produced many progeny . three successive generations ( g1 , g2 and g3 ) were produced on the diet . developmental period , adult body size , and age of first reproduction were almost constant through g0 , g1 and g2 whereas fecundity , adult longevity and daily offspring production declined as the generations proceeded . these results are comparable to previous studies in which pest aphids have been maintained on similar artificial diets for several generations . therefore , it is suggested that the artificial - diet rearing system will provide a useful tool to investigate various biological aspects of the soldier - producing eusocial aphid , t . styraci .
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nomenclatural details : original name . with precedence over myzoxilinae amyot & audient - serville from which it takes the date of priority , 1843 ( iczn article 40 . 2 )
börner . 1944 . in brohmer . fauna von deutschland : ein bestimmungsbuch unserer heimischen tierwelt . fifth edition 218
eastop & van emden . 1972 . in van emden [ ed . ] . aphid technology 13 > > note : = pemphiginae
nieto nafría , mier durante & g . remaudière . 1997 . revue française d’entomologie nouvelle série 19 ( 3 - 4 ) : 82 , 88 > > note : kirkaldy 1905 : 418 ( 1843 )
zhang , guangxue , qiao & xiaolin chen . 1999 . in zhang , guangxue [ ed . ] . fauna of agricultural and forestry aphids of northwest , china : insecta homoptera aphidinea 121
nieto nafría , mier durante & pérez hidalgo . 2011 . in nieto nafría & favret [ ed . ] . registers of family - group and genus - group taxa of aphidoidea ( hemiptera sternorrhyncha ) 56 > > note : kirkaldy 1905 ( 1843 )
li , xingyi , jiang & qiao . 2014 . turkish journal of zoology 38 : 285 - 297
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formerly a separate family , eriosomatidae . tree of life places many ( all ? ) of the included genera in family pemphigidae .
many , but by no means all , root aphids belong to this subfamily . as of 4 / 12 / 10 , the only non - eriosomatine root aphids on bugguide are these aphis ( aphidinae ) :
subfamily hormaphidinae includes two species that cause galls on witch hazel . the manzanita leaf gall aphid is in subfamily tamaliinae .
( the following information is based on bugguide ' s images and may be modified based on future observations . )
the asian woolly hackberry aphid ( calaphidinae : shivaphis celti ) has red eyes , banded antennae , and distinctively patterned wings .
the woolly beech aphid ( phyllaphidinae : phyllaphis fagi ) forms woolly masses on the undersides of beech leaves ( primarily european varieties ) . we do not have adult images for this species yet .
the balsam twig aphid ( mindarinae : mindarus abietinus ) produces white fluff on fir twigs . adults have only a thin , light coating of this material .
alleyne , e . h . and f . o . morrison . 1977 . some canadian poplar aphid galls . the canadian entomologist 109 ( 3 ) : 321 - 328 . [ should help sort out some of the unidentified poplar galls ]
borror and delong ' s introduction to the study of insects norman f . johnson , charles a . triplehorn . 2004 . brooks cole .
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in all aphid species , the ectodermal part of the anterior region of the alimentary tract consists of the stylet bundle , pharynx , foregut and oesophageal valve . the pharynx consists of the pharyngeal duct , valve and pump ( ponsen
) . the midgut is the endodermal part of the alimentary tract consisting of the stomach , crenated intestine and descending intestine . the midgut is responsible for production and excretion of enzymes and the absorption of nutritional substances . in this part of the alimentary tract , often the so - called filter chamber develops ( ponsen
) . it is an adaptation to feeding on the phloem sap , and it takes part in regulation of the osmotic pressure due to the high concentration of sugars in the sap ( rhodes et al .
) . in various groups of aphids , where the filter chamber is present , it is developed variously , e . g . in the subfamily lachninae , the proximal part of the midgut is coiled inside the chamber , creating loops ( klimaszewski and wojciechowski
) . despite the high diversity of the structure of the filter chamber , it is known that foregut and hindgut are connected , so showing the continuity of the tract . in species without the filter chamber , the coiled segments of hindgut adhere the stomach , e . g .
) . the wall of the descending intestine is internally made up of epithelium , whose cells are adapted to resorption of water , amino - acids and minerals and ends with rectum .
) . there is no filter chamber . the very short ectodermal part of the posterior region of the alimentary tract consists of a rectum , epidermal invagination and an anal opening .
and secondary host being the roots of various grasses . their life cycle is two years , and on their primary host , they live in galls ( heie
) . in central europe , the anholocyclic population occurs , with parthenogenetic way of reproduction throughout the year . there are , however , data concerning
consists of a foregut , which is closed at its posterior end , and a blindly starting descending intestine ( fig .
) . it seemed to be odd , taking into account the necessity for constant resorption of vast quantity of protein - poor food ( cristofoletti et al .
) . these species have even the anal plate developed into a trophobiotic organ ( modified anal plate ) , enabling proper positioning of the honeydew droplet ( figs .
alimentary tract of g . utricularia : a extracted from the body ; b schematic reconstruction ; c location within the body ; f foregut , ov oesophageal valve , st stomach , al abdominal loop , di descending intestine
alimentary tract of g . utricularia : a , b longitudinal section through head and thorax ; c transverse section through salivary glands ; d location of stomach within the body ; f foregut , sg salivary glands , ov oesophageal valve , st stomach , di descending intestine
alimentary tract of g . utricularia : a cleared specimen in oblique light microscopy ; b view in clsm , rhodamine - b stained ; c , d cleared specimens in fluorescent microscopy , rhodamine - b stained ; sg salivary glands , ov oesophageal valve , st stomach , al abdominal loop
alimentary tract of g . utricularia : a frontal section through the body ; b longitudinal section through thorax and abdomen ; c , d longitudinal section through posterior part of the abdomen ; sg salivary gland , st stomach , al abdominal loop , di descending intestine , hg hindgut , rb rectal bladder , rg rectal gland , ei epidermal invagination , r rectum , ap anal plate
) , to verify data on the reported lack of the stomach and crenated intestine .
was used because it is more common and abundant , and both species differ only slightly in their morphology and biology . in
, where there are two rows of long setae . the purpose of these setae ( in both species ) is to keep the droplet of honeydew until it is collected by worker ant . their host plants , life mode and relations with ants are in their anholocyclic range very similar and significantly overlap .
the material was collected in october and november 2014 on moist meadows in the vicinity of piekary śląskie , in southern poland . in total , 32 individuals , apterous viviparous females of anholocyclic population on secondary host of
in order to analyse the anatomical structure of the digestive system of g . utricularia , the paraffin method was applied . the material was collected into eppendorf microtubes containing 70 % ethanol for keeping and preserving collected specimens . next , insects were dehydrated in increasing concentration of ethanol ( 90 , 96 , 100 % ) . in order to ensure transparency , specimens were kept in methyl benzoate for one night . then , material was consecutively transferred to benzene , benzene with paraffin ( in proportions 2 : 3 and 1 : 2 ) , paraffin i ( melting point : 56 °c ) and finally to paraffin ii ( melting point : 60 °c ) , where it was kept over night . after this process , material was immersed in paraffin ii . the bars obtained were sectioned into 5 µm strips , which were stuck on slides in a 0 . 5 % gelatine solution at temperature 50–52 °c . then , the slides were dried in 37 °c .
slides were next deparaffined in xylene and treated with a series of ethanol solutions ( 100–60 % ) . they were rinsed in distilled water , stained with ehrlich’s acid hematoxylin for about 20 min , rinsed again and differentiated with xylidine ponceau . after this process , preparations were treated with series of ethanol solutions ( 60–100 % ) , rinsed twice in xylene and embedded in canadian balm or dpx .
this process was applied to 15 individuals of g . urticularia . histological preparations were prepared : cross - sections from six individuals and longitudinal sections from nine individuals . in total , 49 microscopic slides were made , including 26 preparations of cross - section and 23 of longitudinal section . the cross - section series were also used to reconstruct the course of alimentary duct and in some cases the number and shape of cells .
the specimens of g . utricularia were put into a droplet of 30 % ethanol , on the microscopic glass , and with the mounting needles , the whole alimentary tract was extracted from the body . the extracted organs were preserved in glycerol and mounted , to make all the anatomical structures visible at the stereomicroscope . a total number of 13 preparations were made using this technique .
the documentation was prepared using nikon eclipse e6000 , with measurements made by lucia net program . the pictures of translucent specimens were taken under stereomicroscope equipped with monochromatic camera axio cam programmed with axio vision .
in order to investigate localization and structure of gastric tract in g . utricularia , specimens were examined with oblique light microscopy , fluorescence microscopy and confocal laser scanning microscopy ( clsm ) . before the examination , collected specimens were fixated overnight in cold 4 % pfa ( paraformaldehyde ) and subsequently underwent dehydration in graduated ethyl alcohol series , then optical clearing in methyl salicylate .
for fluorescent and clsm imaging , specimens were prior to dehydration treated with 30 % hydrogen peroxide for 24 h and thereafter stained with rhodamine - b before clearing in methyl salicylate . the staining with rhodamine - b was applied for visualizing elastic tissues in clsm ( shelley
) . all reagents used in procedures had analytical grade purity . fluorescent and oblique light imaging was performed with carl zeiss stereo lumar . v12 stereomicroscope and clsm with olympus fv1000 confocal system . acquired images were processed in fiji open source image processing package ( smolla et al .
begins with the external mouthparts and runs through the body cavity without any break or separately located organs . it ends at the anal orifice located above the dorsal edge of the anal plate ( figs .
a ) is a thin straight tube , about 27 μm long , with the diameter of 0 . 01–0 . 02 μm . in the prothorax , it runs between the two main salivary glands ( fig .
c ) , which are built of ca . 20 cells . they are in shape of fan , bottle or other and comprise one or two nuclei located in central or in basal position in the cell , which is typical of exocrine glands ( fig .
a–c ) . their diameter is 0 . 08–0 . 12 μm . the wall of the foregut is made up of one layer of small flat epithelial cells , without any folds into its lumen . in the thorax , the foregut joins the stomach , by invagination into the cavity of stomach ( figs .
the midgut is the longest part of the alimentary tract and consists of the stomach , crenated and descending intestine .
the stomach begins in the second segment of thorax ( mesothorax ) ( figs .
d ) and extends towards the abdomen , slightly dorsally within the body . in longitudinal section , it has a shape of ventro - dorsally flattened bag ( figs .
b ) , constricting gradually along its length . in its widest part , it has a diameter of 0 . 24–0 . 30 μm , and its length varies from 0 . 43 to 0 . 49 μm . the stomach is made up of two types of cells . the anterior part of stomach is built up of elongated , cone - shaped or finger - shaped cells , adhering to each other with lateral membranes at the cell bases ( figs .
a , b ) . their free tips , directed towards the cavity of stomach , broaden the internal surface of the stomach . the cytoplasm of these cells comprises visible granules , which altogether indicates their secretive function .
b ) are flat and hexagonal in shape and possess oval nuclei . the stomach narrows and opens into the crenated intestine , which runs distally ; then , it loops towards the head ( figs .
a ) , adhering the ventral surface of the stomach , and again loops towards the end of abdomen and opens into the descending intestine .
the descending intestine is a straight tube with thin walls , running through abdomen , and its wall is made up of flattened epithelial cells throughout its length ( b , c ) . at its distal end , the descending intestine is broadened , and a well - developed rectal bladder is formed ( fig .
c , d ) . it is positioned dorsally and has a roundish bubble shape , with a few secretive cells in its cavity . these cells are fan - shaped and secrete into the bladder ( fig .
d ) . the rectal bladder opens into the epidermal invagination . it is a straight tube opening with the anal orifice .
( von heyden 1837 ) , another representative of erisomatinae . the presence of doubled nuclei in some of the cells in salivary glands is not surprising , since it was proved to occur also in other aphid families ( ponsen
) , the rectal bladder consists of a ring of cells of polygonal shape , surrounding the hindgut near the rectum . it concerns only females of
spp . , because the presence of rectal bladder was not confirmed in males of this genus .
the role of rectal bladder is not sufficiently recognized . it is probably homologous with malphigian tubules ( ponsen
) , which have disappeared during the evolution of aphids . its possible role is that it takes part in regulation of the honeydew composition ( dixon
) , which contains carbohydrates , especially disaccharides : raffinose and melecitose , synthesized in aphid intestine , and their presence may play role in mutualistic relation of aphids and ants . in particular , the most attractive of ants is the honeydew rich in melecitose ( fisher and shingleton 2001 ) . aphids producing honeydew with 30–70 % of melecitose ( e . g .
stroyan 1950 ) are more often ant attended than those not having melecitose in their honeydew ( e . g .
( koch 1854 ) , with lower amount of melecitose in its honeydew . taking into account the high level of myrmecophily of both
spp . , the presence of well - developed rectal bladder may serve the purpose of regulating the composition of honeydew .
we wish to thank m . b . ponsen ( agricultural university wageningen ) for his kind response and all valuable suggestions and corrections made on the first version of the manuscript . we also thank two anonymous referees for their valuable comments and suggestions on the manuscript .
all applicable international , national and / or institutional guidelines for the care and use of animals were followed .
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three distantly related arthropod groups have converged on a most remarkable social lifestyle . these are gall thrips ( thysanoptera : phlaeothripidae ) , gall aphids ( sternorrhyncha : hormaphididae , pemphigidae , and a few aphididae ) , and snapping shrimp ( decapoda : alpheidae ) , groups in which specialized soldier or defender morphs have evolved in the defense of a static and critical resource , often a nest site that doubles as food source . all three taxa exhibit a range of social structures , the most striking of which involve a morphological division of labor into soldiers and nonsoldier forms . thrips and aphid soldiers have appendages that are enlarged and thickened relative to those of nonsoldiers in the same colony ; these are often armed with spines . similarly , snapping shrimp soldiers bear enlarged chelae . these social groups do not exhibit parental care or much general social interaction ; rather , sociality centers on defense of a valuable and largely irreplaceable resource : the colony domicile . for this reason , this form of sociality is termed
are evident . some social aphids , for example , engage in nest hygiene , expelling honeydew globules that accumulate as a result of the way these insects feed on sap and excrete excess water .
) is produced by a foundress that induces development of a hollow gall on young stem or leaf tissue of the host plant . her brood colony will develop within the safety of the hollow gall , feeding on the plant tissue from within . if the gall is breached , however , the defender morphs will emerge and patrol the surface , aggressively attacking any intruder they encounter . these species are beset by a host of predacious insects such as syrphid fly larvae ; gall thrips also face severe pressure by usurping kleptoparasitic thrips species that kill the foundress and make the gall their own . the domicile of snapping shrimp consists of crevices in sponges , a resource that is less ephemeral than galls but also highly susceptible to usurpation . all of these domiciles represent a valuable , if not critical , commodity that is not easily replaced if lost .
mortality stemming from predation and nest usurpation is likely the most important ecological pressure in the evolution of defender morphs , though it is important to point out that kin selection , in the case of thrips and aphids , may play a role as well . soldier - producing thrips species , which like all members of the order thysanoptera are haplodiploid , often exhibit high levels of inbreeding . aphid colony mates are clonally produced by the foundress , a mode of asexual reproduction called ‘parthenogenesis’ that is found in virtually all aphids and related groups . in social species of both thrips and aphids , there is , accordingly , a high degree of genetic relatedness between colony mates : an average of 0 . 75 in social thrips colonies , and 1 . 0 in colonies of social aphids . this may favor kin selection in these groups , in which indirect fitness – realized through the successful reproduction of relatives – more than compensates the loss of direct fitness that results from altruistic sacrifice ( soldier morphs reproduce less and have a greater chance of dying as a result of colony defense ) .
aphids with morphologically different life stages that exploit taxonomically different host plants are typically difficult to assign to the correct species .
( elm ) species , as the primary host , to identify species and associate the different life stages , thereby using them as a diagnostic tool . using sequence data already available from adult specimens , a phylogeny was produced and the unknown morph sequences clustered with
, confirming the species identity . the authors then described in detail the morphology of the secondary morph , allowing inclusion in future identification keys .
) communities in order to associate adult and larval life stages from tropical lowlands of nepal . although adult taxonomy is well developed and identification is possible ( albeit by taxonomic experts ) , new species are being discovered and the relation of the root - feeding larvae , which have limited morphologically characterising features , to the adults aboveground is unknown . these authors used more in - depth phylogenetic analyses to align sequences to species since intraspecific variation can make this difficult when using absolute methods of sequence divergence and especially for unknown , large assemblages of species . combined mitochondrial and nuclear gene trees ( using maximum likelihood ) and statistical parsimony , whereby populations are subdivided into subgroups , and population aggregation analysis , which combines populations that are uniform for a particular character state , percentage differences (
- distance ) and amova and species delimitation methods for estimating species boundaries from the tree were used to assign species membership . this resulted in 24 species of which 19 could be associated with adults and identified using linnaean names ( nearly 93 % of unknown larvae were identified to species ) . the
sequences , which has implications for gene marker choice in such studies . as for the aphid study , the use of these techniques for determining species and then searching for diagnostic morphological characters of both larvae and adults is advocated . in terms of relating this to species biology , the species assemblages found above - and belowground can depend upon larval mortality , which varies both spatially and temporally ; in particular , larvae and adults were often not found in the same season , which can confound attempts to associate larvae with adults . with a sampling scheme that takes these factors into account , it may be possible to use such dna - based taxonomy to further investigate spatiotemporal patterns and geographic variation .
host - plant resistance is not new to agriculture , but has been applied for at least 200 years . among the earliest successes were resistance to hessian fly ,
, and references therein ) . host - plant resistance is likely to become increasingly important given the continuing homogenization of agricultural landscapes . until the advent of the green revolution , crops were diverse and often hidden in heterogeneous landscapes that consisted of trees , woodland , or other natural and derived habitats . monocultures have eroded landscape diversity and resulted in a loss of habitat for natural enemies , which often increases the vulnerability of crops (
) . agricultural expansion , declining agricultural diversity , and declining crop genetic diversity have demanded a greater focus on intrinsic plant protection . this has coincided with an increasing emphasis on plant molecular biology since the 1990s , largely driven by advances in the molecular tools available to science , including molecular breeding and transgenic technologies .
indicates that for cpb , potato aphids , and tuber moths , the number and proportion of scientific publications related to hpr and insect - plant interactions have generally increased over the past 35 years . during the 2000s , research into conventional hpr and transgenic approaches to plant protection dominated all other management approaches ( cpb 31 % , tuber moths 38 % , and aphids / viruses 24 % of all papers published from 2000 to 2011 ) , including all research on pest natural enemies and biological control . for tuber moths , significant advances have been made in the use of pheromones and granulosis viruses for protection of stored potatoes (
f ) , but for field crops hpr remains the main focus of research . interestingly ,
suggests that research into conventional and transgenic resistance may be mutually competitive in terms of attention or funding opportunities . in the early 2000s , research into transgenics reached its peak ; however , more recently there has been a shift to research of natural induced responses to insect attack and plant biochemical defenses .
it is worth noting that aphids , the masters of phase polyphenism , exhibit yet another form of polyphenism . some aphid species facultatively produce a behaviorally and morphologically distinct soldier caste . since its original description (
abbot et al . , 2001 ; carlin et al . , 1994 ; pike and foster , 2008
foster and rhoden , 1998 ; fukatsu et al . , 2005 ; rhoden and foster , 2002 ; schutze and maschwitz , 1991
) . little is known about proximate mechanisms that induce soldier formation . positive correlations have been found between soldier proportion and colony size (
) , inferring proximate cues that already act during embryonic stages and gradually separating developmental pathways in first instar nymphs .
jhas can be broadly categorized into two groups : the terpenoid compounds , such as methoprene , hydroprene and kinoprene ; and phenoxy jhas , like fenoxycarb and pyriproxyfen . jhas have been found effective especially against adult stages of dipteran and homopteran insects but not against lepidopteran insect pests . as jhas are especially effective against adult stages of the insects , their poor efficacy against lepidopteran crop pests may be because most of these pests cause the damage as larvae (
terpenoid jhas , such as methoprene and kinoprene , have been used successfully against mosquito larvae , scales , and mealybugs . phenoxy jhas cause morphogenetic effects , ovicidal action , and sterility on treated insects . fenoxycarb offers effective control against fire ants , fleas , and sucking insect pests of crops . it is highly effective against sucking pests , such as apple wooly aphid ,
) . pyriproxyfen has been used for controlling aphids , whiteflies , and psyllids causing damage to crops . it has been used for control of locusts ,
most of the bpu compounds have been found very effective against lepidopteran insect pests . diflubenzuron has been found effective against a wide range of lepidopteran pests and locusts (
) . flucycloxuron is effectively used as an acaricide against spider mites . flufenoxuron and novaluron are particularly effective against apple codling moth ,
kanno et al . , 1981 ; yarom et al . , 1988 ; patel et al . , 2010
) . the basis of classical biocontrol is the “enemy release hypothesis . ” according to this hypothesis , the exotic species become pests in new environments by escaping the influence of those natural enemies that suppress their populations in their native range . thus , classical biocontrol reestablishes the top - down control by reintroducing the natural enemies of the pest into its new range (
van driesche and bellows , 1995 ; crawley , 1997 ; keane and crawley , 2002 ; hajek , 2004 ; naranjo et al . , 2015
thus , classical biocontrol primarily describes the releases of insect predators , parasitoids , and pathogens to control other insect pests and insect herbivores to control weeds . this form of biocontrol is appropriate when insects that spread or are introduced ( usually accidentally ) to areas outside of their natural range become pests mainly because of the absence of their natural enemies . the same strategy has also been called “importation” by
( homoptera : mgarodidae ) in california in the late 1800s . likewise , ash whitefly (
, which was introduced in june 1898 . although , the predator did not control soft green scale ,
, which was its specific target , in july 1951 , it controlled various mealybugs infesting fruit crops , coffee , ornamental plants , etc . , in south india . the predator is now effective in suppressing mealybug infestations on citrus , guava , grapes , mulberry , coffee , mango , pomegranate , custard apple , ber , etc . , and green shield scale on sapota , mango , guava , brinjal , and crotons in karnataka (
( native of eastern united states ) , was accidentally introduced to india from england . it soon spread to all of the apple - growing areas of the country and started causing severe damage . for the control of woolly aphid , exotic aphelinid parasitoid ,
( native of north america ) , was introduced from the united kingdom to saharanpur ( uttar pradesh , india ) . however , the parasitoid failed to establish itself because of the intense activity of a ladybird beetle ,
, a native of the caribbean region and central america , was introduced into india in 1995 . first reported from kerala , it soon spread to all of the southern states , causing serious damage to several plants . it started attacking more than 253 plant species belonging to 176 genera and 60 families . however , serious damage was caused to avocado , banana , cassava , guava , papaya , and mango in addition to several ornamental and avenue trees . as a result , exotic aphelinid parasitoids ,
causing a perceptible reduction in the pest population . although the parasitism levels due to both parasitoids vary from 29 % to 70 % and exceed 90 % during some parts of the year , the former is performing better than the latter and has established itself in kerala , karnataka , and several parts of andhra pradesh , where it was previously absent (
despite having many beneficial aspects , classical biocontrol is currently not being encouraged because negative environmental effects may arise through ill - considered introductions of exotic natural enemies . many introduced agents have failed to control pests ; for example , more than 60 predators and parasitoids have been introduced into the northeastern part of north america with little effect on the target gypsy moth ,
( lymantriidae ) . some introductions have strengthened the pest problems whereas others have become pests themselves . exotic introductions generally are irreversible , and nontarget species can suffer worse consequences from efficient natural enemies than from chemical insecticides , which are unlikely to cause total extinctions of native insect species . there are documented cases of introduced biocontrol agents wiping out native invertebrates . several endemic hawaiian insects ( target and nontarget ) have become extinct largely as a result of biocontrol introductions (
although initial classical biocontrol was predominantly focused on control of introduced pests with the goal of reestablishing host / natural enemy associations that keep pests in check in their areas of origin , the strategy has now also been applied against native pests ( e . g . , see
is used when an exotic natural enemy is introduced against a native pest . however , the introduction ( importation , augmentation , and release ) of exotic natural enemies against exotic pests with which they did not coevolve is termed
such new associations are supposed to be very effective at controlling pests because the pest has not coevolved with the introduced enemies . unfortunately , the exotic species that are most likely to be effective biocontrol agents because of their ability to utilize new hosts are also those most likely to be a threat to nontarget species . an example of the possible dangers of neoclassical control is provided by the work of jeffrey lockwood , who campaigned against the introduction of a parasitic wasp and an entomophagous fungus from australia as control agents of native rangeland grasshoppers in the western united states . the potential adverse environmental effects of such introductions include the suppression or extinction of many nontarget grasshopper species with probable concomitant losses of biodiversity and existing weed control and disruptions to food chains and plant community structure . the inability to predict the ecological outcomes of neoclassical introductions means that they are highly risky , especially in systems where the exotic agent is free to expand its range over large geographical areas .


thiotricha fusca is a moth of the gelechiidae family .

it was described by omelko in 1993 .

it is found in japan , the russian far east ( primorye ) and china ( jilin ) .

thiotricha fusca

pyla fusca ( = matilella fusca ) brown knot - horn - norfolk micro moths - the micro moths of norfolk .
thiotricha fusca omelko , 1993 ; biol . issl . est . kul ' t . ekosyst . prim . kr . : 204
polyhumno fusca ; ponomarenko , park & bae , 2006 , j . asia - pacif . ent . 9 ( 2 ) : 108
thiotricha pontifera meyrick , 1932 ; exotic microlep . 4 ( 7 ) : 196
thiotricha synodonta meyrick , 1936 ; exotic microlep . 5 ( 2 ) : 45
thiotricha fridaella legrand , 1958 ; bull . soc . ent . fr . 63 : 142
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thiotricha atractodes meyrick , 1922 ; exotic microlep . 2 ( 16 ) : 502 ; tl : queensland , cairns
thiotricha complicata meyrick , 1918 ; exotic microlep . 2 ( 4 ) : 122 ; tl : coorg , dibidi
thiotricha nephodesma meyrick , 1918 ; exotic microlep . 2 ( 4 ) : 125 ; tl : assam , khasis
thiotricha obvoluta meyrick , 1918 ; exotic microlep . 2 ( 4 ) : 126 ; tl : assam , khasis
thiotricha oxygramma meyrick , 1918 ; exotic microlep . 2 ( 4 ) : 123 ; tl : assam , khasis
thiotricha polyaula meyrick , 1918 ; exotic microlep . 2 ( 4 ) : 125 ; tl : assam , khasis
thiotricha rhodomicta meyrick , 1918 ; exotic microlep . 2 ( 4 ) : 126 ; tl : assam , khasis
thiotricha synacma meyrick , 1918 ; exotic microlep . 2 ( 4 ) : 124 ; tl : assam , khasis
thiotricha xanthaspis meyrick , 1918 ; exotic microlep . 2 ( 4 ) : 122 ; tl : assam , khasis
thiotricha ( thiotrichinae ) ; karsholt , mutanen , lee & kaila , 2013 , syst . ent . 38 : 343
thiotricha amphixysta meyrick , 1929 ; exot . microlep . 3 ( 16 ) : 500 ; tl : assam , khasis
thiotricha attenuata ; ueda & fujiwara , 2005 , trans . lepid . soc . japan 56 ( 1 ) : 75
thiotricha aucupatrix meyrick , 1929 ; exot . microlep . 3 ( 16 ) : 499 ; tl : peru , iquitos
thiotricha celata ; ueda & fujiwara , 2005 , trans . lepid . soc . japan 56 ( 1 ) : 75
thiotricha clepsidoxa meyrick , 1929 ; exot . microlep . 3 ( 16 ) : 499 ; tl : ceylon , maskeliya
thiotricha coleella ; [ nhm card ] ; huemer , 1993 , nota lepid . 16 : 55 ; [ fe ]
thiotricha hamulata meyrick , 1921 ; zool . meded . leyden 6 : 162 ; tl : java , preangor , 5000ft
thiotricha hexanesa meyrick , 1929 ; exot . microlep . 3 ( 16 ) : 501 ; tl : ceylon , maskeliya
thiotricha janitrix meyrick , 1912 ; exot . microlep . 1 ( 2 ) : 64 ; tl : bengal , pusa
thiotricha obliquata ; ueda & fujiwara , 2005 , trans . lepid . soc . japan 56 ( 1 ) : 75
thiotricha rabida meyrick , 1929 ; exot . microlep . 3 ( 16 ) : 500 ; tl : assam , khasis
thiotricha cuneiformis meyrick , 1918 ; exotic microlep . 2 ( 4 ) : 124 ; tl : coorg , dibidi , 3500ft
thiotricha pancratiastis meyrick , 1921 ; exotic microlep . 2 ( 14 ) : 426 ; tl : assam , shillong , 5000ft
thiotricha pyrphora meyrick , 1918 ; exotic microlep . 2 ( 4 ) : 122 ; tl : coorg , dibidi , 3500ft
thiotricha scioplecta meyrick , 1918 ; exotic microlep . 2 ( 4 ) : 123 ; tl : assam , shilleong , 5000ft
thiotricha corylella omelko , 1993 ; biol . issl . est . kul ' t . ekosyst . prim . kr . : 209
thiotricha flagellatrix meyrick , 1929 ; exot . microlep . 3 ( 16 ) : 499 ; tl : assam , shillong , 5000ft
thiotricha indistincta omelko , 1993 ; biol . issl . est . kul ' t . ekosyst . prim . kr . : 205
thiotricha termanthes meyrick , 1929 ; exot . microlep . 3 ( 16 ) : 500 ; tl : assam , shillong , 5000ft
thiotricha tetraphala meyrick , 1886 ; trans . n . z . inst . 18 : 164 ; tl : dunedin , new zealand
thiotricha thorybodes meyrick , 1886 ; trans . n . z . inst . 18 : 164 ; tl : christchurch , new zealand
thiotricha xanthodora meyrick , 1923 ; exot . microlep . 3 ( 1 - 2 ) : 24 ; tl : burma , pyinmana
thiotricha balanopa meyrick , 1918 ; exotic microlep . 2 ( 4 ) : 123 ; tl : assam , khasis ; borneo , kuching
thiotricha hemiphaea turner , 1919 ; proc . r . soc . qd 31 ( 10 ) : 118 ; tl : queensland , toowoomba
thiotricha acrantha meyrick , 1908 ; j . bombay nat . hist . soc . 18 ( 2 ) : 440 ; tl : khasi hills
thiotricha acronipha turner , 1919 ; proc . r . soc . qd 31 ( 10 ) : 118 ; tl : queensland , stradbroke island
thiotricha characias meyrick , 1908 ; j . bombay nat . hist . soc . 18 ( 2 ) : 440 ; tl : palni hills
thiotricha chrysantha meyrick , 1908 ; j . bombay nat . hist . soc . 18 ( 2 ) : 443 ; tl : khasi hills
thiotricha embolarcha meyrick , 1929 ; exot . microlep . 3 ( 16 ) : 500 ; tl : java , mt . salak , 4500ft
thiotricha epiclista meyrick , 1908 ; j . bombay nat . hist . soc . 18 ( 2 ) : 440 ; tl : khasi hills
thiotricha grammitis meyrick , 1908 ; j . bombay nat . hist . soc . 18 ( 2 ) : 442 ; tl : khasi hills
thiotricha hoplomacha meyrick , 1908 ; j . bombay nat . hist . soc . 18 ( 2 ) : 441 ; tl : khasi hills
thiotricha rhodopa meyrick , 1908 ; j . bombay nat . hist . soc . 18 ( 2 ) : 442 ; tl : khasi hills
thiotricha argyrea turner , 1919 ; proc . r . soc . qd 31 ( 10 ) : 118 ; tl : n . queensland , atherton
thiotricha clinopeda meyrick , 1918 ; exotic microlep . 2 ( 4 ) : 124 ; tl : coorg , dibidi , 3500ft ; ceylon , maskeliya
thiotricha galenaea meyrick , 1908 ; j . bombay nat . hist . soc . 18 ( 2 ) : 443 ; tl : maskeliya , ceylon
thiotricha glenias meyrick , 1908 ; j . bombay nat . hist . soc . 18 ( 2 ) : 439 ; tl : maskeliya , ceylon
thiotricha nephelodesma meyrick , 1926 ; exot . microlep . 3 ( 9 ) : 280 ; tl : new ireland , st . matthias i .
thiotricha orthiastis meyrick , 1905 ; j . bombay nat . hist . soc . 16 ( 4 ) : 591 ; tl : rawalpindi , punjab
thiotricha pancratiastis ; [ nhm card ] ; ueda & fujiwara , 2005 , trans . lepid . soc . japan 56 ( 1 ) : 75
thiotricha saulotis meyrick , 1906 ; j . bombay nat . hist . soc . 17 ( 1 ) : 138 ; tl : maskeliya , ceylon
thiotricha scotaea meyrick , 1908 ; j . bombay nat . hist . soc . 18 ( 2 ) : 442 ; tl : maskeliya , ceylon
thiotricha trapezoidella ; [ nhm card ] ; ueda & fujiwara , 2005 , trans . lepid . soc . japan 56 ( 1 ) : 75
thiotricha tylephora ; [ nhm card ] ; ueda & fujiwara , 2005 , trans . lepid . soc . japan 56 ( 1 ) : 75
thiotricha centritis meyrick , 1908 ; j . bombay nat . hist . soc . 18 ( 2 ) : 439 ; tl : palni hills , 6000ft
thiotricha galactaea meyrick , 1908 ; j . bombay nat . hist . soc . 18 ( 2 ) : 441 ; tl : palni hills , 6000ft
thiotricha panglycera turner , 1919 ; proc . r . soc . qd 31 ( 10 ) : 117 ; tl : n . queensland , cairns and kuranda
thiotricha prosoestea turner , 1919 ; proc . r . soc . qd 31 ( 10 ) : 116 ; tl : n . queensland , kuranda near cairns
thiotricha anticentra meyrick , 1904 ; proc . linn . soc . n . s . w . 29 ( 2 ) : 296 ; tl : brisbane , queensland
thiotricha balanopa ; [ nhm card ] ; ueda & fujiwara , 2005 , trans . lepid . soc . japan 56 ( 1 ) : 82 ( note )
thiotricha chrysopa meyrick , 1904 ; proc . linn . soc . n . s . w . 29 ( 2 ) : 293 ; tl : brisbane , queensland
thiotricha clidias meyrick , 1908 ; j . bombay nat . hist . soc . 18 ( 2 ) : 439 ; tl : khasi hills ; maskeliya , celon
thiotricha cuneiformis ; [ nhm card ] ; ueda & fujiwara , 2005 , trans . lepid . soc . japan 56 ( 1 ) : 82 ( note )
thiotricha delacma meyrick , 1923 ; exot . microlep . 3 ( 1 - 2 ) : 24 ; tl : s . india , palnis , kodaikanal , 7000ft
thiotricha margarodes meyrick , 1904 ; proc . linn . soc . n . s . w . 29 ( 2 ) : 294 ; tl : rosewood , queensland
thiotricha pteropis meyrick , 1908 ; j . bombay nat . hist . soc . 18 ( 2 ) : 443 ; tl : khasi hills ; maskeliya , ceylon
thiotricha tethela ; [ nhm card ] ; ueda & fujiwara , 2005 , trans . lepid . soc . japan 56 ( 1 ) : 82 ( note )
thiotricha animosella ; meyrick , 1908 , j . bombay nat . hist . soc . 18 ( 2 ) : 439 ; [ nhm card ] ; [ aucl ]
thiotricha majorella ; [ nhm card ] ; huemer , 1993 , nota lepid . 16 : 47 ; [ me3 ] , 37 ( note ) ; [ fe ]
thiotricha niphastis meyrick , 1904 ; proc . linn . soc . n . s . w . 29 ( 2 ) : 296 ; tl : york , west australia
thiotricha arthrodes meyrick , 1904 ; proc . linn . soc . n . s . w . 29 ( 2 ) : 295 ; tl : sydney , new south wales
thiotricha oxyopis ; bradley , 1961 , bull . brit . mus . ( n . h . ) ent . 10 ( 4 ) : 132 ; [ nhm card ]
thiotricha paraconta meyrick , 1904 ; proc . linn . soc . n . s . w . 29 ( 2 ) : 295 ; tl : wollongong , new south wales
thiotricha bullata meyrick , 1904 ; proc . linn . soc . n . s . w . 29 ( 2 ) : 296 ; tl : broken hill , new south wales
thiotricha leucothona meyrick , 1904 ; proc . linn . soc . n . s . w . 29 ( 2 ) : 294 ; tl : murrurundi and sydney , new south wales
thiotricha oxytheces meyrick , 1904 ; proc . linn . soc . n . s . w . 29 ( 2 ) : 293 ; tl : brisbane , queensland ; sydney , new south wales
thiotricha prunifolivora ueda & fujiwara , 2005 ; trans . lepid . soc . japan 56 ( 1 ) : 76 ; tl : japan , honshu , osaka pref . , takatsuki city , bochikoen
thiotricha parthenica meyrick , 1904 ; proc . linn . soc . n . s . w . 29 ( 2 ) : 297 ; tl : sydney , new south wales ; george ' s bay , tasmania
thiotricha angelica bradley , 1961 ; bull . brit . mus . ( n . h . ) ent . 10 ( 4 ) : 131 , pl . 5 , f . 9 ; tl : guadalcanal , honiara
thiotricha eremita bradley , 1961 ; bull . brit . mus . ( n . h . ) ent . 10 ( 4 ) : 132 , pl . 5 , f . 111 ; tl : guadalcanal , honiara
thiotricha melanacma bradley , 1961 ; bull . brit . mus . ( n . h . ) ent . 10 ( 4 ) : 133 , pl . 5 , f . 12 ; tl : guadalcanal , honiara
thiotricha tethela bradley , 1961 ; bull . brit . mus . ( n . h . ) ent . 10 ( 4 ) : 131 , pl . 5 , f . 10 ; tl : guadalcanal , honiara
thiotricha subocellea ; [ nhm card ] ; huemer , 1993 , nota lepid . 16 : 51 ; ueda & fujiwara , 2005 , trans . lepid . soc . japan 56 ( 1 ) : 75 ; [ fe ]
thiotricha synodonta ; [ nhm card ] ; park , 1991 , ann . hist . - nat . mus . hung . 83 : 121 ; ueda & fujiwara , 2005 , trans . lepid . soc . japan 56 ( 1 ) : 75
korea , china ( jilin ) , japan , se . siberia . see [ maps ]
chinochrysa diakonoff , 1954 ; verh . k . akad . wet . amst . ( 2 ) 50 ( 1 ) : 10
gelechia ( ptocheuusa ) cleodorella zeller , 1877 ; horae soc . ent . ross . 13 : 355 , pl . 5 , f . 120 ; tl : bogota
ptocheuusa coleella constant , 1885 ; ann . soc . ent . fr . ( 6 ) 4 : 255 , pl . 10 , f . 16 ; tl : alpes - maritimes
polyhymno corylella ; ponomarenko , park & bae , 2006 , j . asia - pacif . ent . 9 ( 2 ) : 107
dissobola meyrick , 1935 ; exotic microlep . 4 ( 18 - 19 ) : 561
gemmulans meyrick , 1931 ; exotic microlep . 4 ( 2 - 4 ) : 63
polyhymno ? godmani walsingham , [ 1892 ] ; proc . zool . soc . lond . 1891 : 525 ; tl : west indies , st . vincent
polyhymno indistincta ; ponomarenko , park & bae , 2006 , j . asia - pacif . ent . 9 ( 2 ) : 108
polyhymno laterestriata walsingham , 1897 ; proc . zool . soc . lond . 1897 : 78
lindsayi philpott , 1927 ; trans . n . z . inst . 58 : 84
herzegovina , croatia , n . italy , s . france ( alpes - maritimes ) , greece . see [ maps ]
ptocheuusa majorella rebel , 1910 ; verh . zool . - bot . ges . wien 60 : 28 ; tl : herzegovina
microrrhoda meyrick , 1935 ; exotic microlep . 4 ( 18 - 19 ) : 586
oleariae hudson , 1928 ; butt . & moths n . z . : 254
polyhymno pontifera ; bae , lee & park , 2014 , ent . res . 44 : 19 ( list )
larva on symplocos prunifolia ueda & fujiwara , 2005 , trans . lepid . soc . japan 56 ( 1 ) : 81
eu , european russia , china ( gansu , shaanxi , jilin ) , japan . see [ maps ]
syncentritis meyrick , 1935 ; exotic microlep . 4 ( 18 - 19 ) : 586
mystax trapezoidella caradja , 1920 ; dt . ent . z . iris 34 ( 1 / 2 ) : 138 ; tl : kasakewitsch
mystax trichoma caradja , 1920 ; dt . ent . z . iris 34 ( 1 / 2 ) : 136 ; tl : kasakewitsch
polyhymno tylephora meyrick , 1935 ; mat . microlep . fauna chin . prov . : 68
anacampsis wollastoni walsingham , 1894 ; trans . ent . soc . lond . 1894 : 545 ; tl : madeira
[ afromoths ] de prins , j . & de prins , w . , 2013
sangmi lee , richard brown & sibyl bucheli . gelechioidea - a global framework ;
[ 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 .
microlepidoptera from the solomon islands . additional records and descriptions of microlepidoptera collected in the solomon islands by the rennell island expedition 1953 - 54
microlepidoptera of new guinea , results of the third archbold expedition ( american - netherlands indian expedition 1938 - 1939 ) . part iv
systematische bearbeitung der schmetterlinge von europa , zugleich als text , revision und supplement zu j . hübner ' s sammlung europäischer schmetterlinge , die schaben und federmotten , ( 1847 - ) 1853 - 1855 )
zeller , 1877 exotische microlepidopteren horae soc . ent . ross . 13 : 3 - 493 , pl . 1 - 6
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 .
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recorded in 22 ( 32 % ) of 69 10k squares . first recorded in 1884 . last recorded in 2017 .
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madecassophryne truebae is a species of frog in the microhylidae family .

it is in the monotypic genus madecassophryne .

it is endemic to madagascar .

its natural habitats are subtropical or tropical moist lowland forests , subtropical or tropical moist montane forests , and rocky areas .

it is threatened by habitat loss .

madecassophryne truebae

iucn ssc amphibian specialist group 2016 . madecassophryne truebae . the iucn red list of threatened species 2016 : e . t57867a84178804 . urltoken
- - . . . none , cophyla phyllodactyla , unknown . genus madecassophryne ( guibe , 1974 ) none ,
madecassophryne truebae guibé , 1974 , bull . mus . natl . hist . nat . paris , ser . 3 , zool . , 171 : 1192 . holotype : mnhnp 1973 . 1149 , by original designation . type locality :
chaînes anosyennes
, madagascar .
- - . . . [ 3 ] records returned . . . institution , genus , species , cnt . ummz , madecassophryne ,
see brief account by glaw and vences , 2007 , field guide amph . rept . madagascar , ed . 3 : 120 - 121 . see photograph , map , description of geographic range and habitat , and conservation status in stuart , hoffmann , chanson , cox , berridge , ramani , and young , 2008 , threatened amph . world : 452 . rakotoarison , scherz , glaw , and vences , 2017 , salamandra , 53 : 507–518 , reported on molecular phylogenetics , morphology , osteology , acoustics , and natural history of a population identified as madecassophryne cf . truebae .
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thank you for taking the time to provide feedback on the iucn red list of threatened species website , we are grateful for your input .
frost , d . r . 2015 . amphibian species of the world : an online reference . version 6 . 0 . new york , usa . available at : urltoken .
listed as endangered because its extent of occurrence is 2 , 317 km 2 , it occurs in fewer than five threat - defined locations , and there is continuing decline in the extent and quality of its habitat in southeastern madagascar .
this species is known only from extreme south - eastern madagascar in the anosyenne mountains , andohahela national park and tsitongambarika ( north of tolagnaro ) , between 700 - 1 , 900m asl . it is known from fewer than five threat - defined locations , and its extent of occurrence ( eoo ) is 2 , 317 km 2 .
it is apparently not common , but it is very hard to detect and probably has a patchy occurrence . due to ongoing declines in the extent and quality of the habitat , the population is suspected to be decreasing .
it occurs in forests in an area of extensive , continuous rainforest , and is thought to be associated with boulders and granitic outcrops . it is not known from disturbed habitats . the breeding biology of this species is completely unknown .
the major threat is habitat loss due to subsistence agriculture , timber extraction , charcoal manufacture , the spread of invasive eucalyptus , livestock grazing , and expanding human settlements .
conservation actions it occurs in andohahela national park and tolagnaro fivondronana classified forest . research needed research is needed to investigate the breeding biology of this species .
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a small - sized , ground dwelling microhylid with snout vent length 20 - 23 mm . colour in life is unknown . preserved specimens were dark in color . hindlimbs with bands . upper side of the limbs is yellowish while venter is whitish , with black spots on the throat . sometimes the venter is entirely dark . skin on the back is slightly granular , with large tubercles on the head . nostrils are equidistant between the eye and the tip of the snout . tympanum is rather indistinct , tympanum / eye ratio is 1 / 5 . tibiotarsal articulation reaches the eye . toe 3 is longer than toe 5 . a small inner metatarsal tubercle is present while outer metatarsal tubercle is absent . males are found with a distinct single subgular vocal sac . the call is unknown .
are normally smaller and smooth on the back . confusion is also possible with species of
an egg clutch was found in november near a male and female . it contained 18 eggs with a diameter of 4 mm . the gelatinous egg capsule measured 6 mm in diameter . tadpoles are unknown .
frank glaw and miguel vences ( m . vences at tu - bs . de ) , assistant professor and curator of vertebrates at the institute for biodiversity and ecosystem dynamics in the zoological museum at the university of amsterdam
> university of california , berkeley , ca , usa . accessed jul 9 , 2018 .
> university of california , berkeley , ca , usa . accessed 9 jul 2018 .
this species is known only from extreme south - eastern madagascar in the anosyenne mountains , andohahela national park and tsitongambarika ( north of tolagnaro ) , between 700 - 1 , 900m asl . it is known from fewer than five threat - defined locations , and its extent of occurrence ( eoo ) is 2 , 317 km
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trueb ' s madagascar treefrog ( frank and ramus , 1995 , compl . guide scient . common names amph . rept . world : 90 ) .
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strophedra weirana , the little beech piercer , is a moth of the tortricidae family .

it is found in most of europe , except the iberian peninsula , part of the balkan peninsula , ukraine , the baltic region , finland and ireland .

the wingspan is 10 – 12 mm .

adults are on wing in june .

the larvae feed on fagus species .

they attach two leaves of their host plant together with silk and feed from within .

strophedra weirana

strophedra weirana ( little beech piercer ) - norfolk micro moths - the micro moths of norfolk .
strophedra weirana §1 male ; st lawrence , isle of wight ; 30 / 05 / 2014 ; fw 5 . 1mm © chris lewis
this rather obscurely - marked species occurs sparsely in suitable habitat in the southern half of england , and parts of wales . it is similar to
occupying beech woods , the adults are on the wing during june , and fly in afternoon sunshine , as well as at sunrise .
) , attaching two leaves together with silk and feeding within , causing a noticeable blotch on the leaf surfaces .
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recorded in 18 ( 26 % ) of 69 10k squares . first recorded in 1874 . last recorded in 2018 .
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wingspan about 11 mm . this rather obscurely - marked species is similar to s . nitidana but is less distinctively marked than that species .
adults are on the wing during june , and fly in afternoon sunshine , as well as at sunrise .
the larva feeds on beech , attaching two leaves together with silk and feeding within , causing a noticeable blotch on the leaf surfaces .
this species occurs sparsely in suitable habitat in the southern half of england and parts of wales . in the butterfly conservation’s microlepidoptera report 2011 this species was classified as local .
rare in leicestershire and rutland . this is only the second record for vc55 .
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ws : 10 - 12mm ; jun ; beech ( fagus sylvatica ) ; local in beech wood in s . england synonym : s . flexana ( pierce & metcalfe )