leucinodes is a genus of moths of the crambidae family .

leucinodes species have been documented as eggplant fruit borer , posing medium threats to incoming crops from african nations .

leucinodes

chaetotaxy map of investigated leucinodes larvae ; blue elements illustrate variation found in leucinodes orbonalis and leucinodes africensis ; red elements illustrate the differences found in leucinodes laisalis compared to leucinodes orbonalis and leucinodes africensis .
a polytomous cluster comprising leucinodes rimavallis , leucinodes kenyensis and two undescribed ‘barcode species’ ( leucinodes spp . ) are sister to ( leucinodes africensis + leucinodes pseudorbonalis ) .
larvae of leucinodes . 36–37 leucinodes orbonalis 36 mid instar 37 late instar 38–39 leucinodes africensis 38 mid instar 39 late instar 40–41 leucinodes laisalis 40 early instar 41 late instar .
adult specimens of leucinodes . 1 leucinodes orbonalis , syntype ♂ ( bangladesh ) 2 leucinodes africensis ♀ ( angola ) 3 leucinodes rimavallis ♀ ( dr congo : kivu ) 4 leucinodes pseudorbonalis ♂ ( uganda ) 5 leucinodes kenyensis , holotype ♂ ( kenya ) 6 leucinodes malawiensis , holotype ♂ ( malawi ) 7 leucinodes laisalis ♀ , greyish form ( tanzania ) 8 leucinodes laisalis ♀ , brownish form ( tanzania ) 9 leucinodes ethiopica , holotype ♂ ( ethiopia ) 10 leucinodes ugandensis , holotype ♂ ( uganda ) . scale bar represents 5 mm .
for austral - asia , there remain twelve nominal leucinodes species ( nuss et al . 2003–2014 ) . other than leucinodes orbonalis , at least some of these species are certainly misplaced in leucinodes , e . g . leucinodes labefactalis swinhoe , 1904 from borneo and leucinodes perlucidalis caradja , 1933 from china . therefore , the asian leucinodes are in need of taxonomic revision . this also points to the question whether all leucinodes samples intercepted from asian exports refer to leucinodes orbonalis , or whether there are several leucinodes species of economic importance in asia as well ( hayden et al . 2013 , chang et al . 2014 , gilligan and passoa 2014 ) .
food plant records of african leucinodes species . cultivated plants are given in bold .
fact sheet leucinodes orbonalis guenée . lepintercept - an identification resource for intercepted lepidoptera larvae
male genitalia . 13 leucinodes orbonalis , vietnam ( prep . rm503 ) 14 leucinodes africensis , two - branched sacculus process , côte d’ivoire ( prep . rm330 , phallus omitted ) 15 leucinodes africensis , single - branched sacculus process , ghana ( import ) ( prep . rm501 ) 16 leucinodes rimavallis , kenya ( prep . rm667 ) 17 leucinodes pseudorbonalis , uganda ( prep . rm705 ) 18 leucinodes kenyensis , zimbabwe ( prep . rm694 ) 19 leucinodes malawiensis , malawi ( prep . rm683 ) 20 leucinodes laisalis , south africa ( prep . rm504 ) 21 leucinodes ethiopica , ethiopia ( bmnh pyralidae slide 23138 ) 22 leucinodes ugandensis , somalia ( bmnh pyralidae slide 23140 ) ; phallus mirrored . abbreviations : fi fibula , ga granulated area , sa sacculus , sb side branch of sacculus process , sp sacculus process . scale bar represents 500 µm .
analysis of the coi gene of the leucinodes species demonstrated that interspecific differences allow the use of the marker as a dna barcode for species identification . however , for leucinodes kenyensis and “ leucinodes spp . ” , we found little morphological differences but two distinct barcode species within leucinodes spp . moreover , we observed high intraspecific divergence in leucinodes laisalis with one specimen from south africa exhibiting a coi distance of 2 . 8 % .
female genitalia . 23 leucinodes orbonalis , thailand ( import ) ( prep . rm642 ) , ventral view 24 leucinodes africensis , ghana ( prep . rm640 ) , ventral view 25 leucinodes rimavallis , kenya ( prep . rm666 , smtd lep1592 ) , lateral view 26 leucinodes pseudorbonalis , uganda ( prep . rm706 ) , lateral view 27 leucinodes kenyensis , kenya ( prep . mn1134 ) , lateral view . scale bar represents 500 µm .
head profiles of adult leucinodes orbonalis . 11 male 12 female . figures at same scale .
leucinodes orbonalis is regularly intercepted in the eppo region : addition to the eppo alert list .
leucinodes , brinjal borer , fruit and shoot borer | zoology and entomology articles . ias zoology .
distinguished from the other leucinodes species by the dark , straight - framed forewing base and the absence of the subapical mark of the forewing termen . the male genitalia are similar to those of leucinodes ethiopica and leucinodes ugandensis , but are distinct in the long spinoid process of the posterior phallus apodeme .
this species’ forewing colour has more ochreous than the whitish species of leucinodes but less orange - brown to greyish than in leucinodes laisalis and leucinodes ugandensis . from leucinodes ugandensis and leucinodes laisalis it can be distinguished by the genitalia : in the male genitalia the transtilla arms each bear a dorsad spine ; in the female genitalia the ductus bursae lacks the fine granular sclerotization , the antrum is strongly sclerotized , tubular and widest at its anterior end , and the oval ostial sclerites in the lateral antrum pockets are larger .
female pheromone of brinjal fruit and shoot borer , leucinodes orbonalis : trap optimization and preliminary mass trapping trials .
the record of leucinodes laisalis from belgium by nyst ( 2004 ) is most probably a misidentification , since the illustrated imago resembles much more the whitish leucinodes species . apart from that , there is a european record of leucinodes laisalis from spain . additionally , it is frequently intercepted with the import of solanaceous fruits in great britain .
pupae of leucinodes . 42–43 leucinodes africensis 42 dorsal view 43 close - up of cremaster 44–46 leucinodes laisalis 44 dorsal view 45 lateral view 46 close - up of cremaster . abbreviations : hs hood - like structures dorsal to spiracles on abdominal segments 2 and 3 . scale bar refers to 42 , 44 and 45 and represents 5 mm .
leucinodes kenyensis is a species of moth in the crambidae family . it is found in kenya and possibly zimbabwe .
female genitalia . 28 leucinodes laisalis , kenya ( prep . rm308 ) , lateral view 29 leucinodes ethiopica , ethiopia ( bmnh pyralidae slide 23139 ) , ventral view 30 leucinodes ugandensis , somalia ( bmnh pyralidae slide no . 23137 ) , lateral view 31 leucinodes orbonalis , thailand ( import ) ( prep . rm642 ) , ventral close - up of antrum region 32 leucinodes africensis , côte d’ivoire ( prep . rm743 ) , dorsolateral close - up of antrum region ( phase contrast filter ) 33 leucinodes pseudorbonalis , uganda ( prep . rm706 ) , lateral close - up of antrum region 34 leucinodes kenyensis , kenya ( prep . mn1134 ) , lateral close - up of antrum region . abbreviations : as antrum sclerotizations ; scale bar in 28–30 represents 500 µm and in 31–34 represents 200 µm .
status of the pyraustid moths of the genus leucinodes in the new world , with descriptions of new genera and species .
leucinodes orbonalis guenée , 1854 ; hist . nat . ins . , spec . gén . lépid . 8 : 223
wings . forewing length ♂ 9 . 0 mm , ♀ 9 . 0 mm ; wing pattern as in leucinodes orbonalis .
leucinodes orbonalis guenée , 1854 ( lepidoptera : pyralidae ) , a species not previously recorded in the wild in great britain .
distinguished from most other members of leucinodes by the orange - brown to greyish forewing colour . distinguished from leucinodes ethiopica by the generally darker forewing colour with less amount of white . differs from both leucinodes ethiopica and leucinodes ugandensis in : male genitalia with large , oval sacculus ; broad , strongly sclerotized ventrad fibula ; saccus well elongated ; phallus coecum keeled , posteriodorsal apodeme with slim , fingerlike , well sclerotized process ; female genitalia with antrum broad , its anterior end coiled , with exocuticle diffusely sclerotized .
distinguished from the whitish species of leucinodes and leucinodes ethiopica by the predominantly brown forewing ground colour with minor white patches . distinguished from leucinodes laisalis in the male genitalia : less strongly sclerotized , valvae triangular , fibula small , tooth - like , a similarly shaped distal sacculus process present , saccus process shorter , phallus much shorter , dorsoposterior apodeme without slim , finger - like process .
similar revisionary work remains to be done for austral - asian leucinodes . a phylogenetic study including leucinodes and the new world euleucinodes , neoleucinodes and proleucinodes is needed in order to test the monophyly of these genera ( hayden and mally , in prep . ) .
field evaluation of eggplant cultivars to infestation by the shoot and fruit borer , leucinodes orbonalis ( lepidoptera , pyralidae ) in ghana .
leucinodes labefactalis swinhoe , 1904 ; trans . ent . soc . lond . 1904 ( 1 ) : 158 ; tl : kuching
female pheromone of brinjal fruit and shoot borer , leucinodes orbonalis : trap optimization and preliminary mass trapping trials . - gov . uk
this species is very similar to leucinodes rimavallis , but both coi barcoding data and constant morphological differences in genitalia separate the two species .
wings . forewing length ♂ 7 . 0–8 . 5 mm , ♀9 . 0–11 . 0 mm ; wing pattern as in leucinodes orbonalis .
analyta apicalis ( hampson , 1896 ) , comb . n . ( leucinodes ) . type locality : india , dharamsala . sri lanka .
uncorrected p - distances for the dna - barcoded species of leucinodes . values in bold denote intraspecific distances , plain values represent interspecific distances .
phylogeographical structure in mitochondrial dna of eggplant fruit and shoot borer , leucinodes orbonalis guenée ( lepidoptera : crambidae ) in south and southeast asia .
recently , several interceptions of larvae in solanaceous fruits imported from uganda have been recorded in england ( own observation ) and the netherlands ( marja van der straten , pers . comm . ) . leucinodes pseudorbonalis is one of the three african leucinodes species intercepted at european ports of entry .
wings . forewing length ♂ 7 . 5–10 . 5 m , ♀ 7 . 0–11 . 5 m ; wing pattern as in leucinodes orbonalis .
wings . forewing length ♂ 8 . 5–12 . 0 mm , ♀ 7 . 0–14 . 0 mm ; wing pattern as in leucinodes orbonalis .
leucinodes laisalis clusters in two barcode groups : one group containing all specimens imported with fruits from kenya , ghana and nigeria , and a second group comprising a single south african specimen . this single specimen shows high uncorr - p distances of 2 . 4–2 . 8 % to the other leucinodes laisalis specimens .
ty - jour ti - discovery of an unknown diversity of leucinodes species damaging solanaceae fruits in sub - saharan africa and moving in trade ( insecta , lepidoptera , pyraloidea ) t2 - zookeys ur - urltoken pb - pensoft publishers py - 2015 au - mally , richard au - korycinska , anastasia au - agassiz , david j . l . au - hall , jayne au - hodgetts , jennifer au - nuss , matthias kw - leucinodes leucinodes orb er -
here we taxonomically revise leucinodes and sceliodes and their species from continental sub - saharan africa , in order to delimit species and to allow their proper identification .
known from west africa ( côte d’ivoire , ghana , liberia , nigeria ) , angola , dr congo , gabon , and tanzania ; intercepted with plant imports from ghana and zimbabwe to great britain and the netherlands . at least in the southern dr congo ( lubumbashi ) leucinodes rimavallis occurs sympatrically with leucinodes africensis .
composition of greek pseud ( o ) ‘false’ and orbonalis , meaning ‘false orbonalis ’ , referring to the similarities in external and male genital characters with leucinodes orbonalis .
leucinodes cordalis is known to occur in new zealand , australia , and indonesia : sulawesi ( snellen 1880 , dugdale 1988 , shaffer et al . 1996 ) .
despite repeated search in the collection of the zmhb , original material of leucinodes translucidalis gaede , 1917 from tanzania , tendaguru , could not be traced . according to the original description , this taxon can be regarded as conspecific with leucinodes laisalis due to all details given in the original description . especially the white triangle at the anterior line , another white triangle , though often somewhat inconspicuous , at the middle of costa , and the dark brown area below apex support the conspecifity with leucinodes laisalis .
thea lautenschläger ( technische universität dresden , germany ) and gerard van der weerden ( radboud university nijmegen , the netherlands ) identified the angolan host plants of leucinodes africensis .
the african populations comprise at least three cryptic sibling species . their economical importance remains unclear at the moment . specimens originating from from africa should be determined as “leucinodes sp . ”
@ article { bhlpart216626 , title = { discovery of an unknown diversity of leucinodes species damaging solanaceae fruits in sub - saharan africa and moving in trade ( insecta , lepidoptera , pyraloidea ) } , journal = { zookeys } , url = urltoken publisher = { pensoft publishers 2015 } , author = { mally , richard and korycinska , anastasia and agassiz , david j . l . and hall , jayne and hodgetts , jennifer and nuss , matthias } , year = { 2015 } , keywords = { leucinodes leucinodes orb | } , }
the nature of damage to egg plant ( solanum melongena l . ) in ghana by two important pests , leucinodes orbonalis gn . and euzophera villora ( fldr . ) ( lepidoptera pyralidae ) .
there are further male specimens with indistinctive wing pattern and very similar genitalia , but dna barcode data suggest that among them are at least two further species . for more information , see under leucinodes spp .
male genitalia . as in leucinodes africensis , but with the fibula short , more triangular and robust , straight or slightly curved ; distal sacculus process short and always bent apically ; valva apex rounded or stout .
note : leucinodes orbonalis has now been added to the eppo a1 list . a full datasheet is being prepared , in the meantime you can view here the data which was previously available from the eppo alert list .
leucinodes currently includes twelve old world species distributed in tropical to mediterranean regions . the species share a white or light wing color and a calico pattern ranging from simple brown or dark fields to complex line patterns of light - to dark - brown color . a typical feature is the half moon - shaped spot on the forewing outer margin . several of these species are misplaced in leucinodes due to the superficial resemblance of wing color and pattern .
bishop s , matthews l , macleod a ( 2006 ) csl pest risk analysis . york , uk . urltoken cabi ( 2007 ) crop protection compendium . datasheet on leucinodes orbonalis . urltoken van der gaag dj , stigter h ( 2005 ) pest risk analysis leucinodes orbonalis ( guénée ) . plant protection service , the netherlands . urltoken zhang b - c ( 1994 ) index of the economically important lepidoptera . cabi wallingford , uk , 468 pp .
- most leucinodes specimens found in solanaceous fruits imported from africa into europe during the last 50 years belong to l . africensis , and to a lesser extent to l . laisalis , l . pseudorbonalis and l . rimavallis .
leucinodes orbonalis clusters in two groups , separated by 1 . 1–1 . 8 % uncorr - p distance . within - subcluster distances are 0–0 . 5 % for the smaller and 0–0 . 9 % for the larger of the two subclusters .
lygropia is a polyphyletic genus containing 62 species ( nuss et al . 2003–2014 ) . we provisionally transfer lygropia aureomarginalis ( gaede , 1916 ) , comb . n . ( leucinodes ) from cameroon to this genus , as this species , according to wing pattern elements , is congeneric , if not conspecific , to lygropia vinanyalis viette , 1958 from madagascar . lygropia aureomarginalis can be distinguished externally from species of leucinodes by the shiny golden wing maculation and the presence of two frenulum bristles in the female .
leucinodes malawiensis resembles species of the neotropical genus neoleucinodes capps , 1948 : it shares the prominent diagonal line in the forewing base with neoleucinodes dissolvens ( dyar , 1914 ) , but lacks the long , sabre - like cornutus in the phallus . the absence of the half moon - shaped pattern at the anterior half of the forewing’s outer margin is also found in proleucinodes capps , 1948 . in the coi barcode neighbor joining tree leucinodes malawiensis clusters with neoleucinodes , but is weakly supported with 50 % bootstrap support .
why : since 2004 , more than 120 interceptions of solanum fruits infested by leucinodes orbonalis and imported from asia and africa have been made by several eppo member countries . the panel on phytosanitary measures recommended that this pest should be included into the eppo alert list .
due to the synonymisation of sceliodes , this species is provisionally transferred to leucinodes , as no proper generic placement has been found . compared to leucinodes , several differences can be found in wing pattern of grisealis kenrick , 1912 : in the fore wing , the postmedian line is originating in the apex , and its median protrusion is closely approaching the termen ; the half moon - shaped patch below apex is protruding beyond m 3 ; in the hind wing , the postmedian line is originating closer to the apex and is running closer to the termen .
holotype ♂ [ small blue label ] “gr . kamerunberg | buea 1 . –10 . xi . 10 | 1000–1200 m | e . hintz s . g . ” , [ large blue handwritten label with black border ] “ leucinodes | aureomarginalis | 83 : 8a type gaede” ( zmhb ) .
mally r , korycinska a , agassiz djl , hall j , hodgetts j , nuss m ( 2015 ) discovery of an unknown diversity of leucinodes species damaging solanaceae fruits in sub - saharan africa and moving in trade ( insecta , lepidoptera , pyraloidea ) . zookeys 472 , 117 - 162 .
our work shows that typological concepts of taxonomy based on superficial similarity were still the state of the art in the genus leucinodes . the discovery of a complex of highly similar species demonstrates that traditional morphological methods and dna barcoding are helpful tools to detect species diversity and to improve their classification based on sound arguments .
- african leucinodes correspond to several species , most of which are new to science : l . africensis , l . ethiopica , l . kenyensis , l . laisalis ( = sceliodes laisalis , l . translucidalis ) , l . malawiensis , l . pseudorbonalis , l . rimavallis , and l . ugandensis .
mally r , korycinska a , agassiz djl , hall j , hodgetts j , nuss m ( 2015 ) discovery of an unknown diversity of leucinodes species damaging solanaceae fruits in sub - saharan africa and moving in trade ( insecta , lepidoptera , pyraloidea ) . zookeys 472 : 117–162 . doi : 10 . 3897 / zookeys . 472 . 8781
recorded on several 250 hosts worldwide . some of the major pests of the indian region parasitised by this species are chilo partellus , c . infuscatellus , c . sacchariphagus indicus , c . suppressalis , c . auricilius , cnaphalocrocis medinalis , scirpophaga incertulas , s . excerptalis , raphimetopus ablutellus , leucinodes orbonalis , helicoverpa armigera , and plutella xylostella .
we apply the morphospecies concept to our study . the dna barcode is used as additional source of information and as an identification tool for all developmental stages of african leucinodes species . the solanaceae species names mentioned in this study refer to their former context and do not necessarily correspond to the revised solanum taxonomy by knapp et al . ( 2013 ) .
wings . forewing white translucent , light brown or orange - to grey - brown , basal area light to dark brown , delimited by white and dark brown double line or in species with brown forewing ground colour by dark brown antemedian line ; median area with pale to dark brown , sometimes very faint proximal discoidal stigma ( absent in leucinodes malawiensis ) ; distal discoidal stigma pale to dark brown , reaching from costa to forewing centre ; central dorsum with prominent orange to dark brown , broadly l - shaped or triangular spot connected or disconnected with distal discoidal stigma ; postmedian line sinuate , faint and grey to grey - brown , white edged , with prominent subcostal bulge ; apex brown to grey - brown coloured ( absent in leucinodes malawiensis ) , with slim strip of white at outer margin ; margin dotted at veins , with large dots at apex and m 3 ; fringe white to pale brown with dark interruption at apex and at m 3 ( absent in leucinodes malawiensis ) . hindwing in both sexes with one frenular bristle , ground colour whitish , middle of wing with one or two spots , often faint ; postmedian line inconspicuous , bent towards spot at middle of wing ; area below apex suffused by pale brown to grey ; margin dotted at end of veins , with large dot at end of m 3 .
< mods xmlns : xlink =
urltoken
version =
3 . 0
xmlns : xsi =
urltoken
xmlns =
urltoken
xsi : schemalocation =
urltoken urltoken
> < titleinfo > < title > discovery of an unknown diversity of leucinodes species damaging solanaceae fruits in sub - saharan africa and moving in trade ( insecta , lepidoptera , pyraloidea ) < / title > < / titleinfo > < name > < namepart > mally , richard < / namepart > < / name > < name > < namepart > korycinska , anastasia < / namepart > < / name > < name > < namepart > agassiz , david j . l . < / namepart > < / name > < name > < namepart > hall , jayne < / namepart > < / name > < name > < namepart > hodgetts , jennifer < / namepart > < / name > < name > < namepart > nuss , matthias < / namepart > < / name > < typeofresource > text < / typeofresource > < genre authority =
marcgt
> < / genre > < subject > < topic > leucinodes leucinodes orb < / topic > < / subject > < relateditem type =
host
> < titleinfo > < title > zookeys < / title > < / titleinfo > < origininfo > < publisher > pensoft publishers < / publisher > < / origininfo > < part > < date > 2015 < / date > < / part > < / relateditem > < identifier type =
uri
> urltoken < / identifier > < / mods >
the wing pattern of iriocapna meyrick , 1938 exhibits none of the features found in leucinodes . instead , the fore wings are pale yellow , with a yellowish costa , a dark spot in both outer edges of the cell , and a reddish undulating margin along the termen . the hind wings are of the same pale yellow ground colour , and the anterior half of the termen exhibits a similar margin as found in the fore wings . this wing pattern is common to the genus deanolis snellen , 1899 , where this species is correctly placed ( pers . comm . james e . hayden ) .
types : holotype albicillalis [ circular white label with red border ] “type” , [ beige label with brown border and triangular edges ] “amboina | doll . ” , [ rectangular white label ] “felder | collection . ” , [ rectangular white label ] “rothschild | bequest | b . m . 1939 - 1 . ” , [ rectangular beige handwritten label ] “amboina | dol . ” , [ rectangular beige handwritten label ] “ analyta | albicillalis m” , [ rectangular brown label with central white area , roundly bordered by dark brown and yellow ] “ albicillalis led . ” ( bmnh ) ; holotype apicalis ♂ [ circular label with red border ] “type” , [ rectangular white label ] “4 - 94” , [ rectangular white label ] “ceylon | 95 - 119” , [ rectangular white handwritten label ] “ leucinodes | apicalis | type ♂ hmpsn . ” , transparent capsule with abdomen ( bmnh ) ; holotype pseudoapicalis ♂ [ red rectangular label ] “holotypus” , [ white rectangular label ] “anping | formosa | h . sauter vi . 1911 . ” , [ rectangular white label ] “ analyta | pseudoapi | calis m . | strand det . ♂” ( sdei ) .
html public
- / / w3c / / dtd xhtml 1 . 0 transitional / / en
urltoken
host : this is a pest of brinjal ( solanum melongena ) but also attacks other solanaceous plants .
damage : younger larvae bore into the midrib and petiole of the leaf , resulting in drooping of the leaves . larvae also feed on the tender leaves and flower buds causing dead heart . fruits are attacked by making a hole near the basal part and tunnelling inside . fruit pulp is eaten up . a single larva can destroy several fruits .
life cycle : adult moth is whitish in colour with irregular reddish - brown markings on both wings and having a wing span of 1 . 5 - 2 . 0 cm . longevity of the adults is about a week and fecundity varies between 150 to 250 eggs per female . eggs , which are whitish and flat , are laid singly on the tender shoots or on fruits and hatch in 3 - 5 days . young caterpillars are creamish in colour but full - grown larva is pinkish with brownish head and scattered hairs and warts on its 1 . 5 cm long body . there are 5 larval instars and the larval period varies between 15 and 25 days . pupation takes place in a tough grayish to dull brownish elongated cocoon , usually in hidden portions of the plant . pupal period is 6 - 8 days .
distribution : countrywide distribution in india , burma , sri lanka congo , malaysia , south africa .
control : damaged shoots and fruits should be removed and burnt . spray of 0 . 1 % of carbaryl , sevimol , endrin , diazinon , malathion or endosulfan or cypermethrin 0 . 025 % should be timed with egg laying and larval emergence .
conservation of the following larval parasitoids brings down population : cremastus , pristomerus , bracon , shirakia , iphiaulax sp .
warning : the ncbi web site requires javascript to function . more . . .
richard mally , 1 anastasia korycinska , 2 david j . l . agassiz , 3 jayne hall , 2 jennifer hodgetts , 2 and matthias nuss 4
4 senckenberg natural history collections dresden , königsbrücker landstr . 159 , 01109 dresden , germany
corresponding author : richard mally ( on . biu . mu @ yllam . drahcir )
copyright richard mally , anastasia korycinska , david j . l . agassiz , jayne hall , jennifer hodgetts , matthias nuss
this is an open access article distributed under the terms of the creative commons attribution license ( cc by 4 . 0 ) , which permits unrestricted use , distribution , and reproduction in any medium , provided the original author and source are credited .
pcr of the 5’ half of the cytochrome c oxidase subunit i ( coi ) gene , the so - called dna barcode ( for metazoa ) , was performed using primers hyblco ( folmer et al . 1994 , wahlberg and wheat 2008 ) and hybnancy ( wahlberg and wheat 2008 ) . for degraded material primer pairs hyblco / k699 and ron / hybnancy ( wahlberg and wheat 2008 ) were used to amplify the coi barcode in two fragments . pcr was performed in 25 µl reactions comprising 0 . 4u bio - x - act short dna polymerase ( bioline ) , 2 . 5 µl 10×optibuffer , 1 . 5 mm mgcl 2 , 200 nm each primer , 200 nm dntp mix and 1–2 µl dna ( concentration as extracted ) . cycling conditions were as follows : initial denaturation for 5min at 95 °c , 40 cycles with 1 ) 30 sec at 95 °c , 2 ) 30 sec at 48 °c , 3 ) 90 sec ( hyblco / hybnancy ) or 60 sec ( primers for degraded material ) at 70 °c , final extension of 10 min at 70 °c . alternatively primers lepf and lepr ( hajibabaei et al . 2006 ) were used in 25 µl pcr reactions using bio - x - act short 2× mix ( bioline ) , 400 nm each primer and 1–2 µl dna ( concentration as extracted ) . cycling conditions were as follows : initial denaturation for 5 min at 94 °c , 35 cycles with 1 ) 30 sec at 94 °c , 2 ) 45 sec at 50 °c , 3 ) 1min at 72 °c , final extension of 10 min at 72 °c .
pcr products were visualised by separation in 1–1 . 5 % agarose gels in 1 × tbe buffer ( 89 mm tris - borate 2 mm edta ) containing gelred or ethidium bromide and visualised under uv light . pcr amplicons were cleaned up using exosap - it ( affymetrix ) or qiaquick pcr purification kit ( qiagen ) . sequencing of both strands was performed by eurofins mwg operon ( germany ) or in - house as follows . sequencing pcrs were performed with bigdye terminator v3 . 1 cycle sequencing kit ( applied biosystems ) using 5pm of the sequencing primer tails t7 / t3 ( wahlberg and wheat 2008 ) and 0 . 5–4 µl pcr product . final clean - up was done via sodium acetate - ethanol precipitation . sequencing was performed on a 3130 genetic analyzer ( applied biosystems ) . pcrs , pcr clean - up and sequencing pcrs were performed on a mastercycler ep gradient s ( eppendorf ) or geneamp9700 ( applied biosystems ) .
obtained dna sequences were proofread by eye and aligned using phyde 0 . 9971 ( müller et al . 2008 ) or mega version 4 . 1 ( tamura et al . 2007 ) . ambiguous barcode nucleotides were coded according to the iupac ambiguity code ( cornish - bowden 1985 ) . sequences were then checked for plausibility using blast with the blastn algorithm ( altschul et al . 1990 ; url : urltoken ) as well as the bold identification system ( ids , url : urltoken ) . a 615 basepair fragment was analyzed with mega version 6 ( tamura et al . 2013 ) , using the distance criterion and the neighbor joining ( nj ) algorithm ( saitou and nei 1987 ) with uncorrected p - distances ( srivathsan and meier 2011 ) . statistical support was evaluated through 1 , 000 bootstrap replicates . udea ferrugalis ( hübner , 1796 ) was used to root the nj tree .
label data of studied specimens were compiled in order to generate a distribution map . geographical coordinates , if not given on the label , were obtained via google earth , version 5 . 2 . 1 . 1588 and subsequently plotted on a map using diva - gis , version 7 . 2 . 3 ( hijmans et al . 2004 ) .
the data underpinning the analyses reported in this paper are deposited in the dryad data repository at doi : 10 . 5061 / dryad . kk0n9 .
daraba walker , 1859 ( synonymised by hampson 1899 ) . type species : daraba idmonealis walker , 1859
eretria snellen , 1880 ( synonymised by hampson 1899 ; shaffer et al . 1996 : junior homonym of eretria robineau - desvoidy , 1863 ) . type species : eretria obsistalis snellen , 1880
) , in females shorter than antennal radius ; vertex with whitish to brown scales at the collar and brown scales directed forward ; chaetosemata absent .
thorax . dorsal side whitish to brown with whitish and dark brown scales mixed in ; ventral side whitish ; legs predominantly whitish , foreleg femur , tibia and epiphysis light to dark brown ; tibial spurs 0 , 2 , 4 ( fore - , mid - , hindleg ) with outer spur 1 / 2 to 2 / 3 the length of inner spur .
abdomen . first segment whitish , remainder light - , orange - or dark brown to grey .
) ; phallus simple , with variously shaped sclerites at posterior apodeme , vesica with or without cornuti .
female genitalia . corpus bursae ovoid , membranous , without signa ; ductus bursae membranous with delicate granulation , partly reaching into posterior corpus bursae ; antrum short to long , slim to broader than ductus bursae , anterior part sometimes coiled , mesocuticula thickened ( strongly stained with chlorazol black ) and exocuticula ( inner layer ) partly sclerotized ; ostium bursae with lateral membranous pockets , with or without oval sclerites ; both apophyses pairs simple , apophyses anteriores normally stronger developed than posterior apophyses , with or without broadened central portion .
last instar larvae with pink dorsal integument , intersegmental areas cream or light pink , the ventral integument cream ; strength of the colouration very variable , pink colour on majority of abdominal segments often interrupted laterally by a transverse cream line ; head , prothoracic and anal shields mid brown with variable black markings ; early instar larvae white or cream with brown pinacula and black head , prothoracic and anal shields . in older larvae the dorsal integument turns beige , then increasingly deeper pink as the moults progress , head and prothoracic shield brown ; pinacula pale brown and prominent against the integument in all instars . the chaetotaxy of the thorax and first nine abdominal segments of the last instar is illustrated in fig .
) ; four pairs of long hooked setae ventral to cremaster ; cocoon stout leathery , made of silk , firmly attached to the substrate .
wings . forewing length ♂ 8 . 5–10 . 5 m , ♀ 9 . 5–12 . 0 m ; forewing ground colour white , basal area light - to dark brown , delimited by dark brown to grey antemedial line ; median area with pale brown , faint proximal discoidal stigma ; distal discoidal stigma pale brown , reaching from costa to forewing centre ; central dorsum with prominent orange to dark brown l - shaped or triangular spot leading to forewing centre and often meeting with distal discoidal stigma ; antemedial line sinuate , more or less distinct , but with prominent subcostal bulge ; subapical half of termen with half moon - shaped brown to grey - grown spot ; marginal line dotted ; fringe and marginal line darkened at the tips of the half moon - shaped spot ; hindwing ground colour white , internal area white , with discoidal spot , basicostally often with auxiliary spot ; medial line sinuate , distal half approaching the discoidal spot , then turning towards dorsum ; external area pale brown to gray ; marginal line dotted .
) , valva apex rounded , strongly granulated ; distal 2 / 3 of ventral valva margin loosely covered with long thin setae ; phallus simple , tapering posteriad , posterior apodeme dorsally elongate , ventrally with subapical , weakly serrated sclerite ; ventral and dorsal portion of posterior apodeme separated by a slim , less strongly sclerotized region .
) , with a sclerite process leading in each of the lateral pockets ; both apophysis pairs simple , slightly curved .
msd1 and msd2 of meso - and metathorax usually on a shared pinaculum , earlier instars frequently have the msd setae on separate pinacula on one or both segments ; dorsal abdominal pinacula show apparent differentiation between west - and east - asian populations : in live western specimens ( e . g . , from pakistan ) , the abdominal d1 pinacula usually have an unpigmented cream coloured area near to their anteriomedian margin . this unpigmented area may be contiguous with the unmelanized cuticle surrounding the pinaculum or be surrounded by the melanized cuticle of the pinaculum ( illustrated on a3–6 in fig .
) ; this cream white , unpigmented area may darken in preserved specimens , and in pre - pupae may be ringed in black . eastern specimens ( e . g . , from thailand ) often have dark spots on at least some of the d1 pinacula ( illustrated on a2 in fig .
) ; geographically intermediate populations ( e . g . , sri lanka ) often show an intermediate form with black spots on occasional pinacula , and any unpigmented area is usually ringed with black pigmentation ( illustrated on a7 in fig .
) . east - asian populations usually have mesally triordinal crochets , whereas the crochets of west - asian populations are mesally biordinal .
cremaster forms a variable shelf - like , sub - rectangular structure , much wider than long , usually with distinct distal corners and median notch ; dorsal surface spinulose , with additional small but distinct spines which are variable in extent , form and number ; cocoon of dark brown silk , may be white or beige when newly spun .
india , indonesia : java ( guenée 1854 ) , sri lanka ( walker 1859 , moore 1885 ) , myanmar ( burma ) , andaman islands ( pagenstecher 1900 ) , bangladesh , brunei , cambodia , china , japan , laos , malaysia , nepal , pakistan , philippines , singapore , taiwan , thailand , vietnam ( cabi 2012a ) , australia ( shaffer et al . 1996 ) ; imported to great britain , the netherlands ( pers . comm . m . van der straten ) , denmark ( pers . comm . o . karsholt ) and the u . s . a . ( boateng et al . 2005 , solis 2006 ) .
solanaceae : solanum melongena l . , solanum aculeatissimum jacq . , solanum aethiopicum l . , solanum erianthum d . don . , solanum anguivi lam . ( as solanum indicum l . ) , solanum integrifolium poir . , solanum lycopersicum l . , solanum macrocarpon l . , solanum mammosum l . , solanum nigrum l . , solanum torvum sw . , solanum tuberosum l . , solanum viarum dunal , solanum xanthocarpum schrad . , physalis minima l . , physalis peruviana l . , capsicum annuum l . ( van der straten 2005 ; hayden et al . 2013 ) .
west africa , 11 june 1848 , h . s . le marquand leg .
head . as for the genus , with frons moderately bulged , base of each meron of labial palps with white scales .
) ; ventrad fibula thin , spine - like , curved , crossing distal sacculus anterior to the sacculus process ; valva apex pointed ; posteriodorsal phallus apodeme with prominent oval saw blade - like sclerite , posterioventral apodeme with posteriodorsad oriented tapering process .
as for the genus , apart from : colliculum - antrum complex in sagittal plane of sigmoid shape ; dorsal surface of antrum exocuticle with longitudinal sclerotized strip running from sternite 8 , bearing transverse ridges ( fig .
) ; sternite 8 with anteriomedian recess , anteriolateral edges slightly dentate ; apophyses anteriores with broadened central portion .
, the majority of the abdominal d1 pinacula have a dark pigmented spot on the anteriomedian margin ( illustrated on a2 in fig .
latinized africensis , derived from the continent of africa from where the type material originates and referring to the widespread distribution of this species on the african continent .
solanaceae : solanum aethiopicum l . ( angola , leg . nuss 2013 ) , solanum lycopersicon l . , solanum melongena l .
kenya , mt elgon , 01°07 ' 06
n , 34°41 ' 30
e , february 1952 , t . h . e . jackson leg .
thorax . as for the genus , with dorsal side brown , tegula scales whitish - brown .
female genitalia . as for the genus , apart from : anterior antrum with short sclerotized section , central posterior antrum with diffuse weak sclerotization ; sternite 8 on each side with anteriad triangular process extending into the lateral antrum pockets .
from latin rima for ‘rift’ and vallis for ‘valley’ , referring to the african rift valley , the main distributional area of this species ( as far as known ) .
burundi , eastern and southern democratic republic of the congo , kenya , rwanda , south africa , uganda ( import ) .
solanaceae : solanum melongena l . , withania somnifera ( l . ) dunal .
angola , huambo province , luimbale , mt moco , 1800 - 1900 m , 12°28 ' s , 15°10 ' s , 18 march 1934 , k . jordan leg .
type - specimen . holotype ♂ [ red - circled label ] “holo - | type” , “mt . moco , | luimbale , | 1800 - 1900m . , | 18 march 1934 . ” , “angola | ( dr k . jordan ) ” , “rothschild | bequest | 1939 - 1 . ” , bm pyralidae slide 23135 ( bmnh ) . – additional material . senegal . 1♂ dakar , 01 . viii . 1952 , leg . a . villiers ( bmnh ) ; uganda . 1♂ masindi , 29 . xii . 1897 , leg . ansorge , bmnh pyralidae slide no . 23125 ( bmnh ) ; 1♂ labonga , unyoro , 13 . xii . 1897 , leg . ansorge , bmbh pyralidae slide no . 23126 ( bmnh ) ; 1♂ nabagulo forest , 15 m from kampala , 25 . x . –06 . xi . 1921 , leg . w . feather , bmnh pyralidae slide no . 23145 ( bmnh ) ; 1♂ , ruwenzori range , ibanda , 4 , 700ft , 4 . –12 . ix . 1952 , leg . d . s . fletcher , bmnh pyralidae slide no . 23146 ( bmnh ) ; 1♂ masindi , 30 . x . 1897 , leg . ansorge , prep . rm707 ( bmnh ) ; 2♂ kampala , 1897 , leg . dr . ansorge , bmnh pyralidae slide no . 23149 , prep . rm705 ( bmnh ) ; 1♀ same data , prep . rm706 ( bmnh ) ; the netherlands ( import ) : 1♀ barendrecht , import uganda , 26 . ii . 2014 , leg . sluijs , on solanum melongena , prep . rm758 ( nppo ) ; great britain ( import ) . see suppl . material 2 .
male genitalia . as for the genus , apart from : juxta oval to rectangular ; valvae roughly rhombic ; sacculus process claw - shaped , extending dorsad , parallel to or crossing with fibula ; fibula slightly curved , spine - like , extending dorsad ; posteriodorsal phallus apodeme with a small oval or semicircular sclerite , posterioventral apodeme with simple rodlike process .
as for the genus , apart from : anterior antrum shortly coiled in coronal plane , with the exoculticle exhibiting a longitudinal sclerotized strip bearing transverse ridges ( fig .
) ; sternite 8 intruding into the posteriorly somewhat constricted antrum , giving it a globular appearence .
larva . only one specimen available , examined live . black spots present on dorsal pinacula ; in the final instar , msd1 and msd2 on a shared pinaculum on both meso - and metathorax ; crochets mesally triordinal .
we found this species among material from senegal , uganda and angola , leaving a considerable distribution gap in central africa .
kenya , eastern province , marsabit district , marsabit national park forest , 1158 m , 2°13 . 996 ' n , 37°55 . 676 ' e , 29 december 2003 , r . s . copeland leg .
type - specimens . holotype 1♂ “kenya : marsabit national | park forest . 1158 m . | 2°13 . 996 ' n 37°55 . 676 ' e . | 29 dec 2003 , a & m coll . # 2636 | r . s . copeland ; icipe / usda” , “reared from fruit : | withania somnifera” , dna barcode “usnm ent 007 / 19337” , “1133 | nuss prep . no . ” , coll . nmk . paratypes 1♂ , 1♀ , same data , dna barcodes usnm ent 719338 , 719339 , nuss prep . no . 1135 ( 1♀ nmk , 1♂ smtd ) ; 1♀ kenya , laikipia plateau , mpala research centre , 0 . 293°n 36 . 899°e , 1650 m , 21 . –24 . vi . 2005 , leg . s . e . miller , dna barcode usnm ent 719976 , nuss prep . no . 1134 ( usnm ) . – additional material . zimbabwe . 1♂ mashonaland , leg . h . b . dobbie , prep . rm694 ( bmnh ) .
female genitalia . as for the genus , apart from : anterior antrum with tubular sclerotized exocuticle ; sclerotized wall at antero - ventral edge of the ostium bursae which at rest closes the ostium bursae against abdominal segment 8 . this wall is not melanized and can be stained with chlorazol black . it is delimited dorso - laterally by sclerotized and melanized lobes arising from the anterior edge of segment 8 just ventral of the apophyses anteriores . anterior to the sclerotized wall there is a small , melanized colliculum .
the species is named after kenya , the only country from where it is confidently recorded so far .
so far only known from kenya . the record from zimbabwe needs confirmation by investigation of female specimens and molecular analysis .
malawi , central region , lilongwe district , ntchisi forest reserve , 1560 m , 13°18 . 99972 ' s , 34°02 . 99934 ' e , 18 february 2004 , l . aarvik leg .
type - specimen . holotype ♂ “malawi central | region , lilongwe district : | ntchisi forest reserve | 1560 m 18 . ii . 2004 | leg . l . aarvik” , dna voucher smtd lep1617 , prep . rm683 ( nhmo ) .
wings . forewing length ♂ 8 . 5 mm ; forewing base dark brown , its outer margin a straight diagonal line from the costa to the maculation of the central hind margin , a triangular patch leading lateroposteriad from the costa with the costal half reddish - brown and the central tip white ; outer median area with a faint brownish transverse streak ; subterminal line indistinct except subapical thickening ; apex white ; hindwing antemedial line indistinct ; dark discal spot ; postmedial line clear at costal margin , fading out posteriad ; anterior distal area with a faint brownish transverse streak ; terminal wing veins dark - spotted .
male genitalia . as for the genus , apart from : saccus elongated , u - shaped ; juxta short , oval , twice as broad as long ; valvae slender , tapering towards the dorsad bent apex ; fibula small , triangular , oriented ventrad ; sacculus process absent ; phallus with a spinoid posteriad sclerotization emerging from the ventroposterior apodeme .
megaphysa laisalis walker 1859 : 382–383 . ( hampson 1899 : 275 to sceliodes )
head . frons slightly bulged ; labial palps upturned , greyish to brown , first meron on ventral side with forward - directed tuft , third meron in males half as long as second meron , considerably longer in females ; maxillary palps minute ; haustellum well developed ; eyes large , hemispherical ; ocelli present ; antennae ciliate , cilia considerably longer in males ; vertex with creamy white scales ; chaetosemata absent .
thorax . dorsal side brown with greyish and dark brown scales mixed in ; ventral side grey to whitish ; legs predominantly whitish or grey , epiphysis present ; tibial spurs 0 , 2 , 4 with outer spur 2 / 3 the length of inner spur .
wings . forewing length 7 . 0–11 . 5 mm , the females being somewhat larger ; both sexes with one frenular bristle ; forewing ground colour orange - to grey - brown , with the general sceliodes wing pattern .
abdomen . first segment whitish , remainder light - brown ; older specimens often with darkened abdomen due to degeneration of abdominal fat body .
male genitalia . as for the genus , apart from : vinculum saccus conspicuously elongated anteriad ; juxta usually with small notch at median base ; valvae emerging in narrow angle from vinculum ; phallus with keeled coecum , posteriodorsal apodeme with slim , fingerlike , slightly curved and well sclerotized process , vesica with a short line of tiny cornuti .
female genitalia . as for the genus , apart from : antrum long , tubular , anterior end coiled ; apophysis pairs straight , apophyses anteriores with somewhat broader muscle attachment area at posterior quarter .
solanaceae : solanum incanum l . ( kenya , leg . muli & okuku 2005 ) , solanum anguivi lam . ( “ solanum sodomaeum l . , ” ) , solanum macrocarpon l . , solanum melongena l . , solanum linnaeanum hepper & p . - m . jaeger , solanum lycopersicon l . and capsicum annuum l . ( hayden et al . 2013 ) .
we found a significant dna barcode difference of 2 . 4–2 . 8 % uncorrected - p distance between the single south african specimen and the kenyan and ghanan / nigerian barcode clusters ( fig .
; see also section ‘dna barcoding’ below ) . these differences in the dna barcode are not reflected in a divergent morphology of the clusters .
neighbor joining tree of coi barcodes based on uncorr - p distances and rooted with udea ferrugalis . bootstrap support values derived from 1 , 000 bootstrap replicates ; scale bar represents 1 % uncorr - p barcode divergence .
ethiopia , dire dawa region , dire dawa district , dire dawa , december 1934 , h . ulenhuth leg .
type - specimens . holotype ♂ [ red - circled label ] “holo - | type” , “dire daoua , | abyssinia , | december 1934 . | ( h . uhlenhuth ) . ” ; 19 paratypes : 11♂ 8♀ same data as holotype , including one with bm pyralidae | slide 23138♂ ( bmnh ) . – additional material . eritrea . asmara , 20 . x . 1905 . leg . n . beccari ( without abdomen ) , 1♀ same data except 28 . i . 1905 ( bmnh ) ; ethiopia . 34 ex . same data as holotype except ii . 1935 , 4 ex . ditto except iv . 1935 , 7 ex . ditto except v . 1935 including bm pyralidae | slide 23139♀ , 1 ditto except ix . 1935 , 2 ex . labelled durleti [ = daleti ] ( bmnh ) ; saudi arabia . 2♀ taif ( bmnh ) .
wings . forewing length 6 . 0–8 . 0 mm . forewing mixed ochreous and white , an oblique dark ochreous fascia from above dorsum reaching halfway across wing , a blackish crescent before ochreous subterminal line , black dots along termen . hindwing white , a small black discal spot , a faint irregular dark subterminal line , ochreous suffusion in outer part of wing in middle and towards apex .
male genitalia . as for the genus , apart from : transtilla with short central notch , each transtilla arm with a dorsad spine ; vinculum saccus with rounded tip ; juxta oval to rectangular , apex with a short central notch ; apex of valvae dorsally curved ; small , spine - like dorsad fibula emerging from the central inner side of valva ; sacculus with a fibula - like spiny dorsad process emerging ventrodistally of the fibula ; posteriodorsal phallus apodeme with semicircular dentate sclerite .
female genitalia . as for the genus , apart from : ductus bursae with fine longitudinal ripples ; antrum tubular , with broader anterior end ; apophyses anteriores at posterior half laterally broadened for muscle attachment .
uganda , eastern uganda region , serere district , okulongo , 8 december 1958 , w . r . ingram leg .
type - specimens . holotype ♂ [ red - circled label ] “holo - | type” , “ [ transversally written ] 1608 | “serere | okulongo | 8 dec . 1958 | w . r . ingram | ex solanum sp . ” , “pres . by | com inst ent | bm1959 – 499” , “c . i . e . no . 16499” , with cocoon under specimen . 2 paratypes : 1♂ same data as holotype and “ pyralidae | brit mus | slide no . | 14696” . 1♀ same data as holotype except 9 dec . 1958 ( bmnh ) . – additional material . ethiopia . 2♂ diredaua , n . w . of harar , 1914 , leg . g . kristensen , 1♀ dire dawa , abyssinia , i . 1935 , leg . h . uhlenkuth ( bmnh ) ; south sudan . 1♀ tambura , southern bahr - al - ghasal ( bmnh ) ; somalia . 4♂ 1♀ mogadishu , 17 . –26 . xi . 1985 , 7 . vii . 1986 , 19–20 . viii . 1986 and 23 . vii . 1986 , leg . a . g . parker , bmnh pyralidae slides no . 23140 & 23137 ( bmnh ) ; 1♂ hargeison , 4300ft , v . 1939 , leg . m . portal hyatt ( bmnh ) ; kenya . 1♀ somaliland , mandera , 47km sw of hubera , 3000ft , 13 . xi . 1908 , leg . w . feather ( bmnh ) .
wings . forewing length 6 . 5–11 . 5 mm . forewing pale fuscous , an oblique brown partial fascia at halfway with white markings on either side , an orange triangle on dorsum beyond halfway , apex deep brown , separated by a whitish line . hindwing whitish , fuscous suffused near margin in middle and towards apex , a faint subterminal line in costal part of wing .
male genitalia . as for the genus , apart from : small , hooked dorsad fibula emerging from the ventrocentral inner side of valva ; distal sacculus with dorsad ridge forming a bulge , followed by a spiny , curved terminal process overlapping with the fibula ; phallus vesica with several small spiny cornuti .
female genitalia . as for the genus , apart from : diffusely sclerotized exocuticula reaching into posterior ductus bursae ; lateral antrum pockets rather small .
) . for the remaining specimens listed below , we did not obtain dna barcodes . in spite of the absence of further specimens for comparison , especially females , and the lack of convincing morphological differences , we are not going to describe these possibly distinct species here . this complex needs further study .
kenya . 1♂ rift valley , naivasha , 1900m , 0°46 ' 56
s 36°25 ' 23
e , 5 . xii . 2011 , leg . d . j . l . agassiz , prep . djla 1318 ( coll . djla ) ; zambia . 1♂ chiwefwe , ii . 1950 , leg . n . mitton , bmnh pyralidae slide no . 23133 ( bmnh ) ; namibia . 1♂ namibia , waterberg , 20°30 ' s 17°14 ' e , 13 . iii . 2010 , leg . f . koch , dna voucher mtd lep1872 , prep . rm708 ( zmhb ) ; south africa . 1♂ cape province , knysna , wilderness , iv . 1950 , leg . h . b . d . kettlewell , bm pyralidae slide 23128 ( bmnh ) ; 1♂ johannesburg , 21 . iv . 1906 , leg . a . t . cooke , bmnh pyralidae slide no . 23144 ( bmnh ) ; 1♂ e . cape prov . , katberg , 4 , 000ft , 1 . –15 . i . 1933 , bmnh pyralidae slide no . 23150 ( bmnh ) ; 1♂ kwazulu - natal , estcourt , leg . j . m . hutchinson , prep . rm779 ( bmnh ) ; 1♂ eastern cape [ pondoland ] , port st . john , ix . 1923 , leg . r . e . turner , prep . rm780 ( bmnh ) ; swaziland . 1♂ lebombo - mountains , ndzevane area near nsoko , acacia - rich bushland at the foot of the lebombo mountains , 23 . –24 . i . 2007 , leg . t . karisch , dna barcode bc mtd 01818 , prep . rm502 ( coll . mtd ) .
daraba extensalis walker , 1866 ( synonymised by hampson 1899 ) . type locality : new zealand , auckland .
eretria obsistalis snellen , 1880 ( synonymised by hampson 1899 ) . type locality : indonesia , sulawesi [ celebes ] , boelekomba ; bonthain .
sceliodes mucidalis guenée , 1854 ( synonymised by hampson 1899 ) . type locality : australia .


hippelates is a genus of flies in the family chloropidae and are often referred to as eye gnats or eye flies ( the name is also used for members of the old world genus siphunculina ) .

they are very small ( 1.5 – 2.5 millimetres or 0.06 – 0.10 inches long ) flies that frequently congregate around the eyes to lap at the fluids .

they are primarily a nuisance pest , and do not bite .

they have been linked with the spread of bovine mastitis in north america , and in certain tropical regions , they are capable of vectoring disease-causing bacteria ( e.g. , yaws ) .

hippelates pusio is considered to be the vector for anaplasmosis , bovine mastitis , and haemophilus spp. which cause bacterial conjunctivitis or ' pinkeye ' .

hippelates

lioppelates pusio was first named hippelates pusio by loew in 1872 , and liohippelates bishoppi was described by sabrosky as hippelates bishoppi in 1941 . in 1929 , duda described the genus liohippelates , which now contains the most important common species of eye gnats in the southeastern united states including what was hippelates pusio and hippelates bishoppi .
taplin d , zaias n , rebell g . 1967 . infection by hippelates flies . lancet 2 : 472 .
sabrosky cw . 1941 . the hippelates flies or eye gnats : preliminary notes . canadian entomologist 73 : 23 - 27 .
mulla ms . 1962 . the breeding niches of hippelates gnats . annals of the entomological society of america 55 : 389 - 393 .
bigham jt . 1941 . hippelates ( eye gnats ) investigations in the southeastern states . journal of economic entomology 34 : 439 - 444 .
what made you want to look up hippelates ? please tell us where you read or heard it ( including the quote , if possible ) .
liohippelates bishoppi is found from february through september and liohippelates pusio year round . liohippelates pusio can be found from washington to north dakota , south to pennsylvania , as well as california south to mexico and east to the southeastern united states and bermuda ( gni 2010 ) . liohippelates bishoppi is found from saskatchewan to quebec , south to colorado , texas and florida . other liohippelates species found in the southeastern and eastern u . s . are liohippelates bicolor ( coquillett ) and liohippelates pallipes ( loew ) , whereas hippelates species found in the same area include hippelates nobilis loew , hippelates plebejus loew , and hippelates dorsalis loew ( sabrosky 1941 ) .
sanders da . 1940 . hippelates flies as vectors of bovine mastitis ( preliminary report ) . journal of the american veterinary medical association 97 : 306 - 308 .
kumm hw . 1935 . the natural infection of hippelates pallipes loew with the spirochete of yaws . royal society of tropical medicine and hygiene 29 : 265 - 272 .
observations on predation of hippelates collusor and distribution in southern california of associated fauna < legner , e . f . , l . moore and r . a . medved
kumm hw , turner tb . 1936 . the transmission of yaws from man to rabbits by an insect vector , hippelates pallipes loew . american journal of tropical medicine 16 : 1 - 16
herms wb , burgess , rw . 1930 . a description of the immature stages of hippelates pusio loew and a brief account of its life history . journal of economic entomology 23 : 600 - 603 .
during 1961–62 four hymenopterous parasites of the eye gnat , hippelates collusor ( townsend ) , were discovered in the semiarid agricultural regions of southern california . these parasites were found by field - exposing host hippelates pupae for several days between discs of nylon bolting cloth for predator protection . three of the parasites , phaenopria occidentalis fouts , spalangia drosophilae ashmead , and eupteromalus nidulans thomson , were successfully cultured in the laboratory and also studied in field releases . the fourth , trichopria n . sp . , died before propagation could be attempted . in field releases of parasites s . drosophilae was the most efficient species in seeking hippelates pupae in the environments studied .
hall jr . dg . 1932 . some studies on the breeding media , development , and stages of the eye gnat hippelates pusio loew ( diptera : chloropidae ) . american journal of epidemiology 16 : 854 - 864 .
dow rp , bigham jt , sabrosky cw . 1951 . sequel to
hippelates ( eye gnat ) investigations in the southeastern states
by john t . bigham . proceedings of the entomological society of washington 53 : 263 - 271 .
hippelates probably contribute to the rapid spread of streptococci throughout the island , and to the frequency of infection with more than one serotype ; they may also contribute to the high incidence of streptococcal skin infection , but this requires further evidence .
will fly predators work for hippelates species , i . e . eye gnats ? these are flies , and they breed in rotting organic matter . but elsewhere in your q & r section it is said that fly predators do not work for any kind of gnat .
tondella mlc , paganelli ch , bortoloho im , tankano oa , trino k and brandileone mcc . 1994 . isolamento de harmophilus aegyptius associado a febre purpurica brasileira de cloropideos ( diptera ) dos generous hippelates e liohippelates . revista do instituto de medicina tropical de são paulo 36 : 105 - 109 .
an epidemic of acute glomerulonephritis in trinidad led to an investigation into streptococcal skin infection , as most nephritis cases were associated with this . hippelates in this region commonly feed on skin lesions , infected or otherwise . some of these flies captured in the vicinity of infected children were found to be contaminated with streptococcus pyogenes ; 3 species were involved , h . peruanus , h . flavipes and h . currani . the naturally - acquired infection lasted for 28 hours or more , and the flies would return to human bait while still infectious . the hippelates population and the streptococcal skin infection rate at one school showed similar changes .
the eye gnats , a genus of flies in the family chloropidae ( fruit flies ) that are attracted to the bodily secretions and fluids of animals and humans , particularly those in the eyes . hippelates is suspected of transmitting certain types of conjunctivitis ( such as pinkeye ) , bovine mastitis , and yaws ( frambesia tropica ) .
unfortunately the fly predators do not work on hippelates . primarily the fly predators are for flies in the muscidae family , though they will occasionally parasitize those in the calliphoridae and sarcophagidae families . their very limited host range makes them ideal for biological control as they do not pose a risk to non - pest species ; unfortunately , it also means they are unable to control every pest species of fly .
predatory and scavenger arthropods were surveyed in hippelates collusor ( townsend ) breeding habitats in southern california by using 3 collection methods . a total of 117 different species was collected . the importance of potential predatory species is discussed , and various intensities of natural predation are recorded . there was considerable heterogeneity in predatory activity as influenced by locality , habitat , and seasonal and hourly differences . eggs and 1st - stage larvae sustained the highest predation , with a substantial amount being associated with teneral adults after pupal eclosion .
both liohippelates spp . and closely related hippelates spp . occur throughout much of north america ( sabrosky 1941 ) and 270 species have been described ( sabrosky 1987 ) . bingham ( 1941 ) determined during his investigations of liohippelates in the southeastern u . s . that liohippelates pusio was the most common pest of this genus . later that year , sabrosky ( 1941 ) found and described a new species , liohippelates bishoppi , misidentified by bingham as liohippelates pusio , which made up a moderate portion of the southeastern liohippelates .
physical barriers , such as screened porches and windows on houses and fly masks or fly sheets ( which resemble a fine cloth mesh , largely impenetrable by even the smallest pests ) on livestock , can provide some relief from annoyance and protection from possible pathogen transmission . sanitation methods to eliminate potential breeding sites , such as reducing manure and organic matter in soil and reducing loose soil matter , may reduce breeding at the location . however , often reducing liohippelates and hippelates breeding sites is nearly impossible in livestock facilities as manure , moisture and soil disturbance are inseparable from animals .
members of the genus liohippelates , formerly hippelates , are very small true flies and a common occurrence in much of north and south america ( sabrosky 1980 , kumm 1935 ) . these non - biting pests are attracted to fluids secreted by the eyes , nose and ears on both humans and animals . some species are attracted to discharge from open wounds and excrement ( goddard 2007 ) . because of their propensity for hovering around the eyes , this genus has been referred to commonly as eye gnats , but are also known as grass flies , eye flies , and fruit flies .
in 2006 114 people were examined in santa catalina and san joaquin ; 21 patients were clinically diagnosed with trachoma ( 18 . 4 % ) , 15 ( 13 . 2 % ) of them children under 15 years old . all trachoma phases were observed . three women had corneal opacity with poor vision . in the remaining three communities , three women with advanced trachoma with corneal opacity and blindness were detected . the poor quality of living conditions without fresh water and adequate sanitary disposal systems , and the abundance of flies identified as hippelates sp . , are risk factors for the transmission of the disease .
advisories are not working correctly for plymouth . the next closest stations are rocky mount , williamston and buckland . begin sprays …
from cotton – july 2 , 2018 , 03 : 12 pm please consider attending a scouting school this summer . we will post as they become …
earworm populations ( known as bollworm in cotton ) were relatively low in our system from 2011 to 2016 . arguably , at …
this week i have noticed damage on many pin oaks ( quercus falcata ) by oak spider mites ( oligonychus bicolor ) . oak …
recent news stories about giant hogweed have raised awareness of this nasty invader . yes , we have giant hogweed in …
when cotton blooms , it’s time to switch sampling and thresholds for plant bugs . this previous article covered management of …
we received a confirmed report of basil downy mildew in north carolina . growers are advised to actively scout for …
corn is susceptible to damage at three stages ( roughly ) : v1 to v6 , v14 to vt and r1 to r4 . …
nc state extension is the largest outreach program at nc state university . based in the college of agriculture and life sciences , we reach millions of north carolina citizens each year through local centers in the state ' s 100 counties and with the eastern band of cherokee indians .
we have several topic based e - mail newsletters that are sent out periodically when we have new information to share . want to see which lists are available ?
integrated pest management ( ipm ) is a sustainable approach to managing pests that combines multiple approaches including prevention , avoidance , pest monitoring and suppression in a manner that minimizes public health , economic , and environmental risk . ipm serves as a framework to provide an effective , comprehensive , low - risk approach to protect people and resources from pests .
ultimately , the goal with an ipm program is to help stakeholders deal with pests—insects , plant diseases , weeds , and more—with methods that reduce risks to public health and the environmental while saving money .
the north carolina extension ipm program serves as a focal point for team building , communication and stakeholder participation in integrated pest management ( ipm ) within the state . program goals include promoting effective and economical management of pests , reducing risks to human health from pests and pest management practices , and minimizing environmental effects through the adoption of ipm on a variety of crops and settings in north carolina . these goals are achieved by the timely delivery of ipm technology and research information to stakeholders in all regions of the state .
leadership of the extension ipm program in north carolina is provided by an ipm coordinator ( designated by the director of the north carolina cooperative extension service ) , an established advisory board , and working groups . dr . danesha seth carley , the current ipm coordinator , accepted the nc ipm coordinator position in 2013 .
the extension ipm coordinator involves faculty and staff from north carolina state university and north carolina a & t state university in ipm activities across the state , communicates program successes , and maintains stakeholder input via the advisory board and the ipm portal , as well as multiple training sessions and meetings .
the advisory board provides advice to the extension ipm coordinator regarding the direction of the ipm program in the state . it is composed of a wide variety of ipm stakeholders , including north carolina state university and north carolina a & t state university faculty , non - governmental agencies , environmentalists , north carolina department of agriculture & consumer services personnel , farmers and agricultural consultants .
the north carolina extension ipm program focuses on delivering ipm content to our stakeholders and community members through a variety of activities . the primary activities include information delivery , pest monitoring and data managment , evaluation and needs assessment tools development , and programs to magnify statewide ipm impacts .
the extension ipm program is a cooperative effort of the usda nifa , north carolina cooperative extension service , north carolina state university , and north carolina a & t state university . it also works in partnership with the southern ipm center , located at north carolina state university in raleigh , and acts to promote ipm in north carolina and the southern united states .
funding for the north carolina extension ipm program is provided by competitive grants from the u . s . department of agriculture’s national institute of food and agriculture ( nifa ) .
nc state university and n . c . a & t state university work in tandem , along with federal , state and local governments , to form a strategic partnership called n . c . cooperative extension , which staffs local offices in all 100 counties and with the eastern band of cherokee indians .
nc state university and n . c . a & t state university are collectively committed to positive action to secure equal opportunity and prohibit discrimination and harassment regardless of race , color , national origin , religion , political beliefs , family and marital status , sex , age , veteran status , sexual identity , sexual orientation , genetic information , or disability .
html public
- / / w3c / / dtd xhtml 1 . 0 transitional / / en
urltoken
upcoming events 2018 bugguide gathering in virginia july 27 - 29 : registration and discussion photos of insects and people from the 2015 gathering in wisconsin , july 10 - 12 photos of insects and people from the 2014 gathering in virginia , june 4 - 7 . photos of insects and people from the 2013 gathering in arizona , july 25 - 28 photos of insects and people from the 2012 gathering in alabama photos of insects and people from the 2011 gathering in iowa photos from the 2010 workshop in grinnell , iowa photos from the 2009 gathering in washington
disclaimer : dedicated naturalists volunteer their time and resources here to provide this service . we strive to provide accurate information , but we are mostly just amateurs attempting to make sense of a diverse natural world . if you need expert professional advice , contact your local extension office .
contributors own the copyright to and are solely responsible for contributed content . click the contributor ' s name for licensing and usage information . everything else copyright © 2003 - 2018 iowa state university , unless otherwise noted .
moved from frit flies . id ' d from specimen , now a photo - voucher . thanks john .
select your preferred way to display the comments and click ' save settings ' to activate your changes .
pusio has been evaluated , showing that it is able to transmit the rickettsia up to three days after ingestion of infected erythrocytes [ 23 ] .
( diptera : chloropidae ) and tabanidae on cattle : its possible role in transmission of anaplasmosis .
en publicaciones anteriores ) ( mulla 1962 , legner y bay 1965 , legner et al .
all content on this website , including dictionary , thesaurus , literature , geography , and other reference data is for informational purposes only . this information should not be considered complete , up to date , and is not intended to be used in place of a visit , consultation , or advice of a legal , medical , or any other professional .
html public
- / / w3c / / dtd html 4 . 01 transitional / / en
urltoken
disclaimer : itis taxonomy is based on the latest scientific consensus available , and is provided as a general reference source for interested parties . however , it is not a legal authority for statutory or regulatory purposes . while every effort has been made to provide the most reliable and up - to - date information available , ultimate legal requirements with respect to species are contained in provisions of treaties to which the united states is a party , wildlife statutes , regulations , and any applicable notices that have been published in the federal register . for further information on u . s . legal requirements with respect to protected taxa , please contact the u . s . fish and wildlife service .
html public
- / / w3c / / dtd xhtml 1 . 0 strict / / en
urltoken
common name : eye gnats , grass flies , eye flies , fruit flies scientific name : liohippelates spp . ( insecta : diptera : chloropidae )
figure 1 . lateral view of an adult liohippelates sp . , from a horse farm in north central florida . photograph by lyle j . buss , university of florida .
extremely large aggregations of eye gnats are common in areas that have loose sandy soils , especially in the southern united states . these high concentrations of flies are a great nuisance to humans and animals in rural towns as well as agricultural , recreational and tourist areas . while they do not bite , liohippelates spp . have been implicated in the transmission of several diseases to humans and livestock including human acute conjunctivitis ( pink eye ) . due to the increase of transmission of pink eye and the reduction in labor efficiency caused by extreme numbers of eye gnats in the coachella valley , california , a control project was initiated by the bureau of entomology in the 1920s to focus on the life history and possible control measures for this pest . hall ( 1932 ) subsequently implicated liohippelates pusio ( loew ) as one of the limiting factors in the development of the coachella valley .
the majority of the flies encountered in the southeastern region of the united states are liohippelates pusio and liohippelates bishoppi ( sabrosky ) , and as a result the biological information below will focus broadly on these two species .
adults : adult liohippelates are true flies ( bearing only two wings ) and are very small , approximately 1 . 5 to 2 mm long , and shiny black or grayish in color with clear wings . some species have reddish or yellowish heads . legs tend to be reddish - yellow to brown with brown bands ( hall 1932 ) .
figure 2 . lateral view of an adult liohippelates , unknown species . photograph by erika t . machtinger , university of florida .
eggs : the eggs of liohippelates flies are a pearlescent white and approximately 0 . 5 mm long . they are half as wide ( 0 . 25 mm ) as long , resembling a banana with one side curved and one nearly straight ( hall 1932 ) . eggs are deposited below the surface of loose soils .
larvae : the larvae hatch in seven to 11 days and feed on organic matter ( kahn 2008 ) . the larvae average 3 mm in length and are a whitish color ( hermes and burgess 1930 ) . larvae are pointed towards the anterior end and rounded towards the posterior . the mouth hook is darker than the rest of the body and curved gently downward .
pupae : the pupae are approximately 2 . 25 mm in length , depending on the media in which larvae developed and grew ( hall 1932 ) , and a reddish - brown color , turning darker as they age . liohippelates pupate in or around sand mixed with organic matter , such as manure , hay and remaining harvested crop debris .
figure 3 . primary liohippelates breeding sites in florida include loose soils with organic matter and moisture , as is found on many livestock facilities throughout the state . photograph by erika t . machtinger , university of florida .
liohippelates spp . are holometabolous , meaning they have four distinct life stages ; egg , larvae , pupa and adult . the life cycle of liohippelates can range from 11 days to three months depending on development conditions such as temperature and moisture ( hall 1932 ) . development from egg emergence to the adult is completed in approximately three weeks during the summer in florida ( bigham 1941 ) and multiple generations can occur each year . breeding sites are primarily those with freshly disturbed soil mixed with organic matter , such as cut grass and hay , and moisture . disturbances can be caused by digging , plowing , harrowing , or even by livestock activities ( bigham 1941 , mulla 1962 ) .
liohippelates are not host specific and will feed on the fluid from wounds , eyes , nose , and other areas on humans as well as livestock and domestic pets in a variety of situations . while they do not bite , due to their persistent feeding behavior this group of insects is responsible for extreme annoyance to people and animals when they occur in large numbers . in livestock , it has been suggested that high numbers of liohippelates can cause losses in condition , such as weight , due to continuous and unrelieved attack ( hinkle et al . 2001 ) .
figure 5 . congregation of adult liohippelates spp . around the eye of a horse . photograph by erika t . machtinger , university of florida .
many species of liohippelates have been implicated in the mechanical transmission of several diseases and conditions in humans and livestock . mechanical transmission of pathogens causing disease from liohippelates to humans or livestock occurs when a fly comes in contact with a pathogen , such as a bacterium or virus , and subsequently feeds or rests on a human or animal , transmitting the pathogen from one location to another by contact . as feeding occurs on the blood , mucus , and other fluids around natural orifices of mammals , pathogens can be carried easily into the body .
human acute conjunctivitis , also known as
pink eye
or
sore eyes ,
is caused by one or more bacteria that can be mechanically transmitted by liohippelates . pink eye causes pain , swelling , changes in vision and extreme sensitivity to light ( mideha 2010 ) . incidents of conjunctivitis have been shown to increase during liohippelates outbreaks ( dow and hines 1957 , greenberg 1973 ) . in california , hall ( 1926 ) correlated pink eye to high populations of lioppelates flavipes ( loew ) .
liohippelates flavipes and liohippelates pallipes have been shown to mechanically transmit the spirochete treponema pertenue , the causative agent of yaws , in jamaica and parts of south america ( kumm 1935 , kumm and turner 1936 , saunders et al . 1936 ) . yaws is a skin infection that causes ulceration and , in severe cases , can affect bone and cartilage . yaws occurs mainly in warm and humid regions of the world , including much of south america , and primarily affects children under the age of 15 . yaws is especially common in poor communities and those in crowded conditions ( dnz 2010 ) .
figure 6 . this person has yaws with 8 month - old juxta - articular nodules on the left elbow . photograph by peter perine , cdc .
liohippelates has been implicated in mechanical transmission of pathogens between animals as well . in cattle , liohippelates can transmit the causative agents of acute bovine mastitis ( sanders 1940 ) and vesicular stomatitis ( taplin et al . 1967 ) . acute bovine mastitis is inflammation of the mammary glands , primarily caused by bacterial infection , which can be extremely painful and compromise milk quality . mastitis may also cause death in extreme cases ( bradley 2002 ) . treatment for mastitis includes the use of antibiotics , which has possible implications in public health as resistant bacterial strains may not respond to treatment ( bradley 2002 ) and milk from treated cattle cannot be sold for human consumption until the required withholding time has elapsed .
vesicular stomatitis is a reemerging viral disease that presents as lesions on the tongue , coronary bands and other body regions of cattle as well as horses , sheep , goats and pigs ( usda 2007 ) . this disease is remarkably similar in presentation to the deadly foot - and - mouth disease , which was eradicated from the united states in 1929 ( usda 2007 ) and infection can transfer to people who handle infected animals .
figure 7 . congregation of adult liohippelates around the puncture wound of a horse . photograph by erika t . machtinger , university of florida .
currently , there is no effective area - wide control . various insecticides , fogs , and soil treatments have been tested , but due to the incredible numbers of flies produced in the soil , area - wide treatment is nearly impossible . insecticides applied on a community level for mosquito control may reduce attack , but development sites quickly repopulate as the insecticide dissipates ( kahn 2008 ) .
repellents containing diethyl toluamide ( deet ) provide temporary protection from eye gnats ( hall and gerhardt 2009 ) , and many commercial livestock insecticides that contain pyrethrins , piperonyl butoxide or other common ingredients may provide some relief to animals .
( bpfsg ) the brazilian purpuric fever study group . 1992 . brazilian purpuric fever identified in a new region of brazil . journal of infectious disease 163 : 516 - 519 .
bradley a . 2002 . bovine mastitis : an evolving disease . the veterinary journal 164 : 116 - 128 .
( dnz ) dermnet nz . ( january 2010 ) . yaws . dermnet nz : the dermatology resource . urltoken ( 28 february 2011 ) .
dow rp , hines jd . 1957 . conjunctivitis in southwest georgia . public heath reports 72 : 441 - 448 .
francy db , moore lg , smith gc , jakob wl , taylor sa , calisher ch . 1988 . epizootic vesicular stomatitis in colorado , 1982 : isolation of virus from insects collected along the northern colorado rocky mountain front range . journal of medical entomology 25 : 343 - 347 .
( gni ) global names index . ( 2010 ) . global names index - index of scientific names . urltoken ( 28 february 2011 ) .
goddard j . 2007 . non - biting flies . pp . 191 - 200 . in physicians guide to arthropods of medical importance , 5th edition . crc press , boca raton , fl . 480 pp .
greenberg b . 1973 . flies and disease . vol . 2 . biology and disease transmission . princeton university press , princeton , nj . 856 pp .
hall rd , gerhardt rr . 2009 . flies ( diptera ) . in mullen gr , durden la ( editors ) , medical and veterinary entomology , second edition . elsevier , burlington , ma . 637 pp .
harrison ih , da silva ga , pitman m , fleming dw , vranjac a , broome cv . 1989 . epidemiology and clinical spectrum of brazilian purpuric fever . journal of clinical microbiology 27 : 599 - 604 .
hinkle nc , scholl pj , mock de and warner wb . 2001 . research and extension needs for integrated pest management for arthropods of veterinary importance . pp . 261 - 262 . in geden cj , hogsette ja ( editors ) , proceedings of a workshop in lincoln , nebraska . second edition . 328 pp .
( itis ) ( 2010 ) . liohippelates . integrated taxanomic information system . urltoken ( 28 february 2011 ) .
kahn c ( ed . ) . ( 2008 ) . eye gnats . the merck veterinary manual . urltoken ( 28 february 2011 ) .
looper m , stokes sr , waldner dn , and jordan er . ( 2001 ) . feeding waste milk to dairy calves . new mexico state university college of agricultural , consumer and environmental sciences . urltoken ( 28 february 2011 ) .
( mideha ) infectious disease and environmental health administration . ( 2010 ) . conjunctivitis (
pink eye
) . maryland department of health & mental hygiene . urltoken ( 05 may 2011 ) .
sabrosky cw . 1987 . chloropidae . pp . 1049 - 1067 . in mcalpine jf , et al . ( editors ) , manual of nearctic . diptera . volume 2 . research branch agriculture . canada monograph 28 : 675 - 1332 .
saunders gm , kumm hw , rerrie ji . 1936 . the relationship of certain environmental factors to the distribution of yaws in jamaica . american journal of hygiene 23 : 558 - 579 .
( usda ) united states department of agriculture . ( march 2007 ) . information about vesicular stomatitis for the beef producer . urltoken ( 06 may 2011 ) .
publication date : april 2011 . latest revision : may 2011 . reviewed : october 2015 .
confused by a class within a class or an order within an order ? please see our brief essay .
to cite this page : myers , p . , r . espinosa , c . s . parr , t . jones , g . s . hammond , and t . a . dewey . 2018 . the animal diversity web ( online ) . accessed at https : / / animaldiversity . org .
disclaimer : the animal diversity web is an educational resource written largely by and for college students . adw doesn ' t cover all species in the world , nor does it include all the latest scientific information about organisms we describe . though we edit our accounts for accuracy , we cannot guarantee all information in those accounts . while adw staff and contributors provide references to books and websites that we believe are reputable , we cannot necessarily endorse the contents of references beyond our control .
this material is based upon work supported by the national science foundation grants drl 0089283 , drl 0628151 , due 0633095 , drl 0918590 , and due 1122742 . additional support has come from the marisla foundation , um college of literature , science , and the arts , museum of zoology , and information and technology services .
the story of an imaginary word that managed to sneak past our editors and enter the dictionary .
take this quiz and discover 12 words for things you didn ' t know had words .
we use cookies to enhance your experience on our website . by continuing to use our website , you are agreeing to our use of cookies . you can change your cookie settings at any time .
i agree to the terms and conditions . you must accept the terms and conditions .
thank you for submitting a comment on this article . your comment will be reviewed and published at the journal ' s discretion . please check for further notifications by email .
you could not be signed in . please check your email address / username and password and try again .
most users should sign in with their email address . if you originally registered with a username please use that to sign in .
to purchase short term access , please sign in to your oxford academic account above .
natural infection of nyssomyia neivai ( pinto , 1926 ) ( diptera : psychodidae , phlebotominae ) by leishmania ( viannia ) spp . in brazil
analysis of covalent modifications of amyloidogenic proteins using two - dimensional electrophoresis : prion protein and its sialylation .
open repair of mycotic abdominal aortic aneurysms with biological grafts : an international multicenter study .
oxford university press is a department of the university of oxford . it furthers the university ' s objective of excellence in research , scholarship , and education by publishing worldwide
for full access to this pdf , sign in to an existing account , or purchase an annual subscription .
female eye gnats maintained on honey in the laboratory retained more than half their original radioactivity 5 days after being tagged with p 32 . when tagged gnats were released at the center of three concentric circles of traps on a windy day , some moved 25 feet directly upwind but an equally large number was caught 75 feet downwind in traps on the outermost circle , on both sides of the mean downwind direction . releases of tagged gnats ½ and 1 mile from a rural population center in southwestern georgia resulted in almost complete penetration of the small town on the day of release . in one test , traps more than a mile from the release box caught 15 gnats in less than 3½ hours after it was opened . chi - square analysis was used to test the hypothesis that tagged gnats are distributed like the wild ones . local departures from an overall equilibrium between tagged and untagged gnats could be recognized , and progress of the dispersal could be followed by comparing successive collections .
we use cookies to distinguish you from other users and to provide you with a better experience on our websites . close this message to accept cookies or find out how to manage your cookie settings .
this article has been cited by the following publications . this list is generated based on data provided by crossref .
vet , louise e . m . and lenteren , j . c . 2009 . the parasite - host relationship between encarsia formosa gah . ( hymenoptera : aphelinidae ) and trialeurodes vaporariorum ( westw . ) ( homoptera : aleyrodidae ) . zeitschrift für angewandte entomologie , vol . 91 , issue . 1 - 5 , p . 327 .
gordh , g . legner , e . f . and caltagirone , l . e . 1999 . handbook of biological control . p . 355 .
mann , j . a . stinner , r . e . and axtell , r . c . 1990 . parasitism of house fly ( musca domestica ) pupae by four species of pteromalidae ( hymenoptera ) : effects of host–parasitoid densities and host distribution . medical and veterinary entomology , vol . 4 , issue . 3 , p . 235 .
hammond , w . n . o . and neuenschwander , p . 1990 . sustained biological control of the cassava mealybugphenacoccus manihoti [ hom . : pseudococcidae ] byepidinocarsis lopezi [ hym . : encyrtidae ] in nigeria . entomophaga , vol . 35 , issue . 4 , p . 515 .
hammond , w . n . o . neuenschwander , p . and herren , h . r . 1987 . impact of the exotic parasitoid epidinocarsis lopezi on cassava mealybug ( phenacoccus manihoti ) populations . international journal of tropical insect science , vol . 8 , issue . 4 - 5 - 6 , p . 887 .
debach , paul huffaker , c . b . and macphee , a . w . 1976 . theory and practice of biological control . p . 255 .
legner , e . f . sjogren , r . d . hall , i . m . and washino , r . k . 1974 . the biological control of medically important arthropods . c r c critical reviews in environmental control , vol . 4 , issue . 1 - 4 , p . 85 .
horn , david j . 1971 . the relationship between a parasite , tetrastichus incertus ( hymenoptera : eulophidae ) , and its host , the alfalfa weevil , hypera postica ( coleoptera : curculionidae ) , in new york . the canadian entomologist , vol . 103 , issue . 01 , p . 83 .
( townsend ) , arranged in clumped or linear distributions , oviposited randomly producing equal numbers of progeny from cither host arrangement . mixed groups of linear and clumped puparia caused changes in behavioral patterns that resulted in reduced progeny production , thus showing parasitization to be nonrandom . host destruction was , nevertheless , greater in an all - clumped distribution , tile greater acceptance of clumped groups resulting in a greater number of progeny . mixed groups involving a choice between linear or clumped distributions demonstrated the greater acceptance of clumped groups . direct observation showed that the all - dumped distribution elicited the greatest overall initial attraction for hosts and subsequent movement to other areas . it was concluded that maximum host destruction resulted when completely random behavior was involved . a recognition of this , however , required a knowledge of behavior , host condition , and progeny production . the question of evaluating an exotic parasite’s effectiveness before introduction is discussed .
behavior changes the reproduction of spalangia cameroni , s . endius , muscidifurax raptor , and nasonia vitripennis ( hymenoptera : pteromalidae ) at increasing fly host densities
some mechanisms that affect the sex ratio of nasonia vitripennis ( walk . ) ( hymenoptera : pteromalidae ) reared from superparasitized house - fly pupae
email your librarian or administrator to recommend adding this journal to your organisation ' s collection .
[ clinical evidence of trachoma in colombian amerindians of the vaupés province ] . - pubmed - ncbi
warning : the ncbi web site requires javascript to function . more . . .
programa de enfermedades transmitidas por vectores , secretaría departamental de salud , mitú , vaupés , colombia .
trachoma is the leading cause of infectious blindness in the world . in 2008 there were 1 , 300 , 000 persons with blindness caused by trachoma and 8 million with trichiasis , which might eventually lead to blindness . in latin america it has been documented in brazil , guatemala and méxico .
to inform the presence of trachoma for the first time in colombia , amongst amerindians of the department of vaupés .
in 2003 and 2006 the amerindian makú communities of san joaquín and santa catalina , located 5 km from the border with brazil , were visited . from 2007 to 2009 , san gerardo , san gabriel and nuevo pueblo , at a 35 km distance from san joaquín were visited .
trachoma exists in colombia , and it is frequent among the studied communities . its focalized distribution makes it amenable to elimination . it is advisable to search for trachoma in other indigenous communities in vaupés with similar living conditions .
doctype html public
- / / w3c / / dtd html 4 . 0 transitional / / en
ask us a question and a member of our support staff will respond as soon as possible .
the checklists are descriptive documents of the species that are part of a concrete taxonomic group or a group of species that are part of an specific analysis .
these documents have a structure that includes a list of the authors , a description of the taxonomic group or subject , and a list of the species that are part of the list .
the checklists aren ' t updated regularly , since they require revisions from groups of authors before their publication .
a complete set of a checklist include a pdf document and a table in csv format .
gustavo jiménez - uzcátegui , david a . wiedenfeld , f . hernán vargas , howard l . snell .
nathalia tirado - sanchez , john mccosker , diego ruiz , angel chiriboga , stuart banks .
yves finet , nathalia tirado - sanchez , angel chiriboga , diego ruiz , stuart banks .
frank bungartz , frauke ziemmeck , alba yánez ayabaca , fredy nugra , andré aptroot .
anne guézou , susana chamorro , paola pozo , ana mireya guerrero , rachel atkinson , chris buddenhagen , patricia jaramillo díaz , mark gardener .
gustavo jiménez - uzcátegui , javier zabala , brian milstead , howard l . snell .
to get up - to - date information about our work , please subscribe to our e - newsletter or follow us on our social media platforms .
every single donation we receive , no matter how small , counts as we are completely dependent on the generosity of others to carry out our scientific projects . we need your passion , loyalty and continual support .
the “charles darwin foundation for the galapagos islands” , in french “fondation charles darwin pour les îles galapagos” , association international sans but lucratif ( “aisbl” ) , has its registered office located at drève du pieuré 19 , 1160 brussels , and is registered under the trade registry of brussels under the number 0409 . 359 . 103 .