steinernema scapterisci , the mole cricket nematode , is a species of nematode in the order rhabditida .

it is a parasite of insects in the order orthoptera , the grasshoppers , crickets and their allies .

it has been introduced into florida in the united states in an effort to provide a biological pest control of mole crickets .

steinernema scapterisci

steinernema scapterisci is a close relative of the insect - parasitic nematode steinernema carpocapsae .
11 . a formulation as claimed in claim 10 in which the two or more species of steinernema are selected from the group comprising : steinernema feltiae ; stinernema scapterisci ; steinernema riobravis ; steinernema carpocapsae ; and steinernema krussei .
16 . a formulation as claimed in claims 14 or 15 in which the at least one species of steinernema is selected from the group comprising : steinernema feltiae ; stinernema scapterisci ; steinernema riobravis ; steinernema carpocapsae ; and steinernema krussei .
infection of sound - trapped mole crickets , scapteriscus spp . , by steinernema scapterisci
12 . a formulation as claimed in claim 10 in which the two species of steinernema are selected from the group comprising steinernema feltiae , steinernema krussei , and steinernema carpocapsae .
. vertical dispersal of steinernema scapterisci . journal of nematology , 22 : 574 - 578 .
17 . a formulation as claimed in claim 16 in which the at least one species of steinernema is selected from the group comprising : steinernema feltiae ; steinernema krussei and steinernema carpocapsae .
. bacterial symbionts of steinernema scapterisci . journal of invertebrate pathology , 62 : 68 - 72 .
. addendum to the morphology of steinernema scapterisci . journal of nematology , 24 : 478 - 481 .
. pathogenicity of steinernema scapterisci to selected invertebrates . journal of nematology , 23 : 7 - 11 .
. life cycle of steinernema scapterisci nguyen & smart , 1990 . journal of nematology , 24 : 160 - 169 .
. mode of entry and sites of development of steinernema scapterisci . journal of nematology , 23 : 267 - 268 .
. steinernema scapterisci n . sp . ( steinernematidae : nematoda ) . journal of nematology , 22 : 187 - 199 .
nguyen kb , smart gc jr . 1990b . vertical dispersal of steinernema scapterisci . journal of nematology 22 : 574 - 578 .
anguillera mm , smart gc jr . 1992 . bacterial symbionts of steinernema scapterisci . journal of invertebrate pathology 62 : 68 - 72 .
nguyen kb , smart gc jr . 1992b . addendum to the morphology of steinernema scapterisci . journal of nematology 24 : 478 - 481 .
nguyen kb , smart gc jr . 1991a . pathogenicity of steinernema scapterisci to selected invertebrates . journal of nematology 23 : 7 - 11 .
. preliminary studies on survival of steinernema scapterisci in soil . soil and crop science society of florida , proceedings , 49 : 230 - 233 .
since 1985 steinernema scapterisci has been detected in mole crickets collected from several other sites in alachua county , undoubtedly spread from our original release sites by infected mole crickets . by november 1998 , about 10 to 30 percent of mole crickets collected in alachua county , florida were infected with steinernema scapterisci .
figure 1 . mole cricket nematodes , steinernema scapterisci nguyen & smart , emerging from a mole cricket . photograph by k . nguyen , university of florida .
figure 8 . mole cricket nematodes , steinernema scapterisci nguyen & smart , emerging from a cockroach nymph . photograph by k . nguyen , university of florida .
nguyen kb , smart gc jr . 1992a . life cycle of steinernema scapterisci nguyen and smart , 1990 . journal of nematology 24 : 160 - 169 .
nguyen kb , smart gc jr . 1991b . mode of entry and sites of development of steinernema scapterisci . journal of nematology 23 : 267 - 268 .
nguyen kb , smart gc . pathogenicity of steinernema scapterisci to selected invertebrates . j nematol . 1991 ; 23 ( 1 ) : 7–11 . pmid : 19283088
nguyen kb , smart gc jr . 1990 . steinernema scapterisci n . sp . ( steinernematidae : nematoda ) . journal of nematology 22 : 187 - 199 .
. development , reproduction , and pathogenicity of steinernema scapterisci in monoxenic culture with different species of bacteria . journal of invertebrate pathology , 62 : 289 - 294 .
aguillera mm , hodge nc , stall re , smart gc . bacterial symbionts of steinernema scapterisci . j invertebr pathol . 1993 ; 62 ( 1 ) : 68–72 .
this project was initiated in june 1999 to re - establish commercial production of steinernema scapterisci for mole cricket control in florida . in about one year , we have :
nematicides were used on golf courses against plant - parasitic nematodes and doubtless would kill steinernema scapterisci . however , such chemicals no longer are labeled for that purpose by u . s . environmental protection agency . studies in the early 2000s showed that steinernema scapterisci is remarkably resistant to chemical insecticides . the effect of chemical herbicides on this nematode is unknown .
nguyen kb , smart gc . steinernema scapterisci n . sp . ( rhabditida , steinernematidae ) . j nematol . 1990 ; 22 ( 2 ) : 187–99 . pmid : 19287709
figure 3 . spicules ( top ) and gubernaculum ( bottom ) of the mole cricket nematode , steinernema scapterisci nguyen & smart . photographs by k . nguyen , university of florida .
nguyen kb , smart gc jr . 1990a . preliminary studies on survival of steinernema scapterisci in soil . soil and crop science society of florida , proceedings 49 : 230 - 233 .
nguyen kb , smart gc . life - cycle of steinernema - scapterisci nguyen and smart , 1990 . j nematol . 1992 ; 24 ( 1 ) : 160–9 . pmid : 19283218
. inoculative release of steinernema scapterisci ( rhabditida : steinernematidae ) to suppress pest mole crickets ( orthoptera : gryllotalpidae ) of golf courses . environmental entomology , 23 : 1331 - 1337 .
figure 6 . infective juvenile of the mole cricket nematode , steinernema scapterisci nguyen & smart , with the head showing labial raising disc . photograph by k . nguyen , university of florida .
anguillera mm , smart gc jr . 1992 . development , reproduction , and pathogenicity of steinernema scapterisci in monoxenic culture with different species of bacteria . journal of invertebrate pathology 62 : 289 - 294 .
aguillera mm , smart gc . development , reproduction , and pathogenicity of steinernema scapterisci in monoxenic culture with different species of bacteria . j invertebr pathol . 1993 ; 62 ( 3 ) : 289–94 .
citation : lu d , sepulveda c , dillman ar ( 2017 ) infective juveniles of the entomopathogenic nematode steinernema scapterisci are preferentially activated by cricket tissue . plos one 12 ( 1 ) : e0169410 . urltoken
nguyen , k . b . , smart , g . c . , jr . 1990 . steinernema scapterisci n . sp . ( rhabditida : steinernematidae ) . journal of nematology 22 : 187 - 199 .
steinernema scapterisci was successfully introduced by inoculative applications ( parkman et al . 1993 , 1994 ) in florida . since 1993 , this nematode has been commercialized in florida to control mole crickets in golf courses and pastures .
biological control , entomopathogenic nematode , mole cricket parasite , morphology , new species , neoaplectana , scapteriscus , steinernema scapterisci n . sp . , taxonomy , light microscopy ( lm ) , scanning electron microscopy ( sem )
. biological control of mole crickets in the genus scapteriscus with the nematode steinernema scapterisci nguyen & smart , 1990 . rencontres caraibes en lutte biologique , guadeloupe , 5 - 7 november 1990 . ed . inra , paris 1991 .
30 . use of a formulation according to claim 1 to treat one or more of the tawny mole cricket and the southern mole cricket , and wherein the cohort of infective juvenile nematodes consists of steinernema scapterisci and two species of heterorhabditis .
parkman jp , frank jh , nguyen kb , smart gc jr . 1994 . inoculative release of steinernema scapterisci ( rhabditida : steinernematidae ) to suppress pest mole crickets ( orthoptera : gryllotalpidae ) of golf courses . environmental entomology 23 : 1331 - 1337 .
parkman jp , frank jh , nguyen kb , smart gc jr . 1993 . dispersal of steinernema scapterisci ( rhabditida : steinernematidae ) after inoculative applications for mole cricket ( orthoptera : gryllotalpidae ) control in pastures . biological control 3 : 226 - 232 .
24 . a method as claimed in claim 22 in which the target insect is one or more of the tawny mole cricket and the southern mole cricket , and wherein the cohort of infective juvenile nematodes consists of steinernema scapterisci and two species of heterorhabditis .
steinernema biddulphi n . sp . can be distinguished from other steinernema species by means of a combination of morphological and morphometric characteristics of males and ijs . based on these data , steinernema biddulphi n . sp . belongs to the “ bicornutum ” clade within the steinernematidae family . molecular data show that within this clade , steinernema biddulphi n . sp . is sister to the pair of s . pakistanense and s . bifurcatum and this group is related to the pair of s . yirgalemense and s . abbasi .
figure 5 . a , b - face views of first generation females ; c - female tail ; d , e , f - double flapped epiptygma ; of the mole cricket nematode , steinernema scapterisci nguyen & smart . photograph by k . nguyen , university of florida
parkman jp , frank jh , nguyen kb , smart gc . inoculative release of steinernema - scapterisci ( rhabditida , steinernematidae ) to suppress pest mole crickets ( orthoptera , gryllotalpidae ) on golf - courses . environmental entomology . 1994 ; 23 ( 5 ) : 1331–7 .
a multigene approach for assessing evolutionary relationships of xenorhabdus spp . ( gamma - proteobacteria ) , the bacterial symbionts of entomopathogenic steinernema nematodes
in 1985 , a nematode parasite of mole crickets was brought from uruguay to florida . that nematode , which was described as a new species , steinernema scapterisci ( nguyen and smart 1990 ) , appears to be a major factor in limiting populations of mole crickets in south america .
comparative morphometrics ( in μm ) of infective juveniles of steinernema biddulphi n . sp . ; mean ± sd with ranges given in brackets .
some commercial companies may sell steinernema nematodes other than s . scapterisci as biopesticides for use against mole crickets . if the label does not tell you the species of nematode , then ask the supplier to give the name in writing . it is true than other steinernema species will kill mole crickets , at least to some extent , but do not expect them to give any residual activity because they are not known to reproduce in mole crickets .
comparative morphometrics ( in μm ) of first generation males of steinernema biddulphi n . sp . ; mean ± sd with ranges given in brackets .
phylogenetic relationships of the species from “ bicornutum ” group and other related species of steinernema based on analysis of its rdna regions . steinernema nepalense and s . scapterisci was used as outgroup taxon . the percentage of replicate trees in which the associated taxa clustered together in the bootstrap test ( 10 , 000 replicates ) are shown next to the branches . branch lengths indicate evolutionary distances and are expressed in the units of number of base differences per site .
smart gc jr , nguyen kb , parkman jp , frank jh . 1990 . biological control of mole crickets in the genus scapteriscus with the nematode steinernema scapterisci nguyen and smart , , 1990 . rencontres caraibes en lutte biologique , guadeloupe , 5 - 7 november 1990 . ed . inra , paris 1991 .
scapterisci ( rhabditida : steinernematidae ) after inoculative applications for mole cricket ( orthoptera : gryllotalpidae ) control in pastures . biological control , 3 : 226 - 232 .
meeting held with a company in georgia to rear s . scapterisci in vivo ( 11 / 17 / 99 ) , confidentiality agreement completed and business plan submitted .
meeting held with uf , otl ( 1 / 20 / 00 ) , agreed to assist in locating a company to produce s . scapterisci using in vitro methods .
phylogenetic relationships of the species from “ bicornutum ” group and other related species of steinernema based on analysis of d2 - d3 expansion segments of the 28s rdna . steinernema nepalense and s . scapterisci was used as outgroup taxon . the percentage of replicate trees in which the associated taxa clustered together in the bootstrap test ( 10 , 000 replicates ) are shown next to the branches . branch lengths indicate evolutionary distances and are expressed in the units of number of base differences per site .
foreign company and uf , otl drafted an exclusive license agreement for production of s . scapterisci for sale in the u . s . ( 6 / 19 / 00 ) .
steinernema scapterisci was more pathogenic to insects tested in the order orthoptera than to those in the orders lepidoptera or hymenoptera ; it was not pathogenic to earthworms . the nematode also infected and killed the mole crickets scapteriscus acletus and s . vicinus when released four successive times at 10 - day intervals in containers of soil infested with the nematode .
9 . a formulation as claimed in any preceding claim in which the first and second genus are selected from the group consisting of : steinernema ; and heterorhapditis .
furthermore , the first generation females of steinernema biddulphi n . sp . differ from the females of s . pakistanense and s . bifurcatum by having a postanal swelling .
10 . a formulation as claimed in claim 9 in which the first genus ( majority species ) is steinernema and the second genus ( minority species ) is heterorhapditis .
14 . a formulation as claimed in claim 9 in which the first genus ( majority species ) is heterorhapditis and the second genus ( minority species ) is steinernema .
the whole genome shotgun project for s . scapterisci has been deposited at ddbj / embl / genbank under the accession [ azbw00000000 ] . the version described in this paper is version azbw01000000 .
nguyen kb , maruniak j , adams bj . diagnostic and phylogenetic utility of the rdna internal transcribed spacer sequences of steinernema . j nematol . 2001 ; 33 : 73–82 .
morphometric characters ( in μm ) of steinernema biddulphi n . sp . based on the holotype and 20 paratypes of each generations with mean ± sd and ranges given in brackets .
parasitism is a major ecological niche for a variety of nematodes . multiple nematode lineages have specialized as pathogens , including deadly parasites of insects that are used in biological control . we have sequenced and analyzed the draft genomes and transcriptomes of the entomopathogenic nematode steinernema carpocapsae and four congeners ( s . scapterisci , s . monticolum , s . feltiae , and s . glaseri ) .
meeting held with richard gaskalla , fdacs / dpi ( 1 / 13 / 00 ) , determined that they would be receptive to rearing s . scapterisci if provided with the necessary funding and direction .
report with background information on mole cricket infestations , damage and efficacy of control by s . scapterisci prepared for fdacs ( 2 / 25 / 00 ) , report sent with justification for a federal appropriation .
hox cluster architecture in steinernema . comparisons of the hox clusters of c . elegans , s . carpocapsae , and s . scapterisci . each cluster is mapped at the same scale , with the colored boxes representing different putative genes between the lin - 39 and ceh - 13 orthologs . genes marked in blue are specific to steinernema , not having orthologs in c . elegans . gray genes have a c . elegans ortholog , though they are not syntenic in the nematodes compared . genes marked in brown are unique , not having obvious orthologs in the other nematodes in this comparison
32 . use of a formulation according to claim 1 to treat a root weevil , and wherein the cohort of infective juvenile nematodes consists of heterorhabditis megidis , and two species of steinernema .
figure 7 . diagram of the life cycle of steinernema scapterisci . g1 = first - generation adults , g2 = second - generation adults , j1 = first - stage juvenile , j2 = second - stage juvenile that may be the preinfective or non - preinfective stage , j3 = third stage noninfective juvenile , pi = preinfective stage , ij = third - stage infective juvenile , j4 = fourth - stage juvenile .
26 . a method as claimed in claim 22 in which the target insect is a root weevil , and wherein the cohort of infective juvenile nematodes consists of heterorhabditis megidis , and two species of steinernema .
meeting held with uf , office of technology licensing ( 10 / 20 / 99 , 2 month delay ) , agreed that up front costs would be minimal for a license to sell s . scapterisci for mole cricket control .
29 . use of a formulation according to claim 1 to treat one or more of sciarid larvae and fungus gnats , wherein the cohort of infective juvenile nematodes consists of steinernema felitae and two species of heterorhabditis .
the phylogenetic trees of the its and 28s genes were obtained by the minimum evolution method ( rzhetsky and nei , 1992 ) in mega 6 . 0 ( tamura et al . , 2013 ) . steinernema nepalense and steinernema scapterisci were used as outgroup . the minimum evolution tree was searched using the close - neighbor - interchange ( cni ) algorithm ( nei and kumar , 2000 ) . the neighbor - joining algorithm ( saitou and nei , 1987 ) was used to generate the initial tree . the evolutionary distances were computed using the p - distance method ( nei and kumar , 2000 ) and are expressed as the number of base differences per site .
sleinemema scapterisci nguyen et smart , 1990 is recorded for the first time in argentina . this entomopathogenic nematode was found parasitizing the mole cricket scapteriscus borelli giglio - tos at the locality of colon ( province of buenos aires ) , argentina .
nadler sa , bolotin e , stock sp . phylogenetic relationships of steinernema travassos , ( nematoda : cephalobina : steinernematidae ) based on nuclear , mitochondrial and morphological data . syst parasitol . 2006 ; 63 : 161–81 .
23 . a method as claimed in claim 22 in which the target insect is one or more of sciarid larvae and fungus gnats , and wherein the cohort of infective juvenile nematodes consists of steinernema felitae and two species of heterorhabditis .
34 . use of a formulation according to claim 1 to treat one or more of root weevils , wood borers , and scarabs , and wherein the cohort of infective juvenile nematodes consists of heterorhabditis bacteriophora and two species of steinernema .
meeting held with uf , otl , and a foreign company to discuss an agreement to produce in vitro and market s . scapterisci in the u . s . ( 4 / 13 / 00 ) , the company requested a 60 - day non - compete agreement .
28 . a method as claimed in claim 22 in which the target insect is one or more of root weevils , wood borers , and scarabs , and wherein the cohort of infective juvenile nematodes consists of heterorhabditis bacteriophora and two species of steinernema .
lee mm , stock sp . a multigene approach for assessing evolutionary relationships of xenorhabdus spp . ( gamma - proteobacteria ) , the bacterial symbionts of entomopathogenic steinernema nematodes . j invertebr pathol . 2010 ; 104 ( 2 ) : 67–74 . pmid : 20102721
meeting held with fdacs , dpi and the mole cricket task force to determine the status of state funds requested to produce s . scapterisci and initiate a mole cricket control program ( 7 / 21 / 00 ) , appropriation is in a trust fund and project is unable to proceed .
figure 2 . males of the mole cricket nematode , steinernema scapterisci nguyen & smart n . sp . a ) spicules of the first - generation male showing angular head , ribs , and gubernaculum with anterior end bent upward . b ) variation in the tail shape of the first - generation males . c ) tail of the second - generation male showing elongate spicule head . d ) variation in tail shape of second - generation males . e ) entire body of the first - generation male . drawing by k . nguyen , university of florida .
31 . use of a formulation according to claim 1 to treat one or more of root weevils , the tawny mole cricket , and the southern mole cricket , and wherein the cohort of infective juvenile nematodes consists of steinernema riobravis , and two species of heterorhabditis .
25 . a method as claimed in claim 22 in which the target insect is one or more of root weevils , the tawny mole cricket , and the southern mole cricket , and wherein the cohort of infective juvenile nematodes consists of steinernema riobravis , and two species of heterorhabditis .
33 . use of a formulation according to claim 1 to treat one or more of armyworms , cutworms , webworms , root weevils , wood borers , artichokes , and plume moths , and wherein the cohort of infective juvenile nematodes consists of steinernema carpocapsae and two species of heterorhabditis .
nadler sa , bolotin e , stock sp . phylogenetic relationships of steinernema travassos , 1927 ( nematoda : cephalobina : steinernematidae ) based on nuclear , mitochondrial and morphological data . systematic parasitology . 2006 ; 63 ( 3 ) : 161–81 . epub 2006 / 03 / 17 . pmid : 16541298
meeting held with fdacs , dpi to plan production and prepare a budget , $ 281 , 000 year 1 ( 2 / 17 / 00 ) , planned a realistic structure and budget for solid culture of s . scapterisci , and delivered it to dr . martha roberts ( fdacs ) in the commissioner ' s office .
27 . a method as claimed in claim 22 in which the target insect is one or more of armyworms , cutworms , webworms , root weevils , wood borers , artichokes , and plume moths , and wherein the cohort of infective juvenile nematodes consists of steinernema carpocapsae and two species of heterorhabditis .
a . house crickets were anesthetized at 4°c and were then put individually into the wells of a cryo storage box . each well was lined with a small piece of paper towel loaded with approximately 100 ijs of s . scapterisci suspended in 100 μl of tap water . b . about 3–4 infected crickets were transferred to white traps [ 31 ] .
we have identified a set of expanded gene families that are likely to be involved in parasitism . we have also identified a set of non - coding motifs associated with groups of orthologous genes in steinernema and caenorhabditis involved in neurogenesis and embryonic development that are likely part of conserved protein–dna relationships shared between these two genera .
comparing activation rates of ijs exposed to different insect homogenates , mole cricket homogenate induced significantly higher activation than house cricket homogenate at 10 % ( p < 0 . 0001 ) , and 50 % ( p = 0 . 009 ) concentrations . both cricket homogenates induced significantly higher activation in s . scapterisci ijs than waxworm homogenate ( p < 0 . 0001 ) .
steinernema biddulphi sp . n . was reared on last instar of g . mellonella as described by kaya and stock ( 1997 ) . the first and second generation adults were obtained from the dissection of 3 - and 5 - d - old infected g . mellonella larvae , respectively . infective juveniles were collected approximately 1 wk after emergence from the cadavers .
dillman ar , macchietto m , porter cf , rogers a , williams b , antoshechkin i , lee mm , goodwin z , lu x , lewis ee , goodrich - blair h , stock sp , adams bj , sternberg pw , mortazavi a . comparative genomics of steinernema reveals deeply conserved gene regulatory networks . genome biol , 2015 ; 16 ( ) : 200
protein gels showing the quantity and diversity of proteins secreted by activated ( 12h ) and non - activated ( na ) ijs . equal percentage of total volumes of concentrated proteins from non - activated and activated s . scapterisci ijs was loaded onto each gel . ijs were activated with 50 % cricket homogenate . a . a protein gel stained with colloidal coomassie blue . b . a protein gel with silver staining .
the effects of environmental factors ( such as temperature , moisture , aeration , and soil type [ esp . texture and chemistry ] ) and biotic factors ( species of epn , targeted insect , age of insect , soil fauna ) have been documented by numerous researchers ( gaugler and kaya , 1990 ; kaya and gaugler , 1993 ; shapiro - ilan et al . , 2012 ; grewal et al . , 2005a ; georgis et al . , 2006 ) . temperature range for survival and infectivity will depend on the species of epn and its native habitat and center of origin ( kaya , 1990 ) . for example , steinernema feltiae can be infective from 2 - 30°c , whereas some heterorhabditids can infect host insects from 7 to 35°c and steinernema carpocapsae is nearly inactive at 10°c ( kaya , 1990 ; georgis et al . , 2006 ; lacey et al . , 2006a ) .
the life cycle of s . biddulphi n . sp . is similar to other steinernema species . in an experimental infection of g . mellonella with a dose of 50 ijs per insect at a temperature of 22°c , the majority of insects were dead after 48 h , and fourth stage juveniles were present in the hosts . two amphimictic generations occur inside the host , and the first and second generation stages could be observed after 2 and 8 d of initial infection , respectively . infective juveniles appeared 10 d postinoculation .
s . scapterisci ijs were exposed to three different host homogenates ; scapteriscus borellii , acheta domesticus , and galleria mellonella . the homogenates were made in different concentrations ( 10 % , 25 % , 50 % and crude ) . we were unable to test crude waxworm homogenate for technical reasons ; the crude homogenate was too thick to be separated from nematodes after exposure but we were able to test 10 % , 25 % , and 50 % waxworm homogenate . each of these activation experiments was collected and observed after 18 hours of exposure to the homogenate . ij activation was then quantified as described above .
steinernema biddulphi n . sp . lm of infective juvenile ( ij ) , male and female . a , c . first generation female . a . tail with postanal swelling . c . vulval region . b , d . second generation female . b . tail with postanal swelling . d . vulval region . e . first generation male , tail with spicules and gubernaculum . f . second generation male , tail with spicules and gubernaculum . g , h . ij . g . anterior portion showing rounded head and excretory pore ( arrow ) . h . tail with anus and hyaline region .
steinernema biddulphi n . sp . a . first generation male , tail with spicules and gubernaculum , lateral . b . first generation female , vulval region . c . first generation female , tail . d . second generation female , vulval region . e . second generation female , conical tail . f , g . infective juvenile . f . anterior region showing excretory pore and nerve ring . g . tail showing anus and hyaline region . ( scale bar in micrometers : a = 58 ; b = 143 ; c = 68 ; d = 71 , e = 61 ; f = 91 ; g = 75 ) .
in recent years , several surveys have been conducted in south africa resulting in the recovery of eight new steinernematid species , namely steinernema khoisanae ( nguyen et al . , 2006 ) , s . citrae ( stokwe et al . , 2011 ) , s . sacchari ( nthenga et al . , 2014 ) , s . tophus ( cimen et al . , 2014 ) , s . innovationi ( cimen et al . , 2015 ) , s . jeffreyense ( malan et al . , 2015 ) , s . beitlechemi ( cimen et al . , 2016 ) , and s . fabii ( abate et al . , 2016 ) .
based on the blast search and phylogenetic analysis of the concatenated sequences , sequences of the 16s rdna and reca and gyrb genes ( data not shown ) , the symbiotic bacterium of s . biddulphi n . sp . ( bacterial strain sgi - 246 ) seems to be most closely related to xenorhabdus indica ( blast similarities 98 % for 16s rdna , and 97 % for reca and gyrb genes ) . this bacterium was found in association with other nematodes from the “ bicornutum ” group , namely steinernema abbasi ( somvanshi et al . , 2006 ) , s . yirgalamense ( ferreira et al . , 2016 ) , and s . bifurcatum ( shahina et al . , 2014 ) .
steinernema biddulphi n . sp . scanning electron microscopy of infective juvenile ( ij ) , male and female . a–c . ij . a . head region with horn - like structures . b . lateral field in mid - body ( ridges numbered 1 to 6 ) . c . lateral field in tail region . d . first generation female , tail , and four projections on tip of the tail ( arrow ) . e , f . second generation female . e . tail with postanal swelling . f . vulva . g . first generation male , tail with paired genital papillae ( numbered ) and single papilla ( s ) , lateral . h , i . second generation male . h . tail with paired genital papillae , single papilla ( s ) and mucron ( arrow ) . i . tail with a part of paired genital papilae ( numbered ) , mucron ( arrow ) , and phasmid opening ( p ) , ventro - lateral .
s . scapterisci activation was categorized based on 3 morphological features ( fig 2 ) — ( 1 ) the mouth being open or closed , ( 2 ) the state of the anterior gut , or the opening of the gut immediately posterior to the pharyngeal bulb , and ( 3 ) expansion of pharyngeal bulb and how pronounced it appeared . if an individual nematode had all three characteristics ( an open mouth , an open and expanded anterior gut , and an expanded and visible pharyngeal bulb ) they were considered fully activated ( fig 2c ) . if the nematode had a visible pharyngeal bulb , though had not proceeded to a further stage of activation as indicated by the gut and mouth opening , it was considered partially activated ( fig 2d ) . if the nematode had no visible pharyngeal bulb , it was considered non - activated ( fig 2b ) . non - activated nematodes were generally still ensheathed , though presence of the l2 cuticle sheath was not used to determine nematode activation . dead nematodes were not counted . any nematodes that had developed beyond l3 into l4 or even adulthood were counted as fully activated .
proteins were denatured by sds sample buffer ( final 1x buffer : 50 mm trishcl ph6 . 8 , 2 % sds , 2 . 5 % ficoll , 0 . 01 % bromophenol blue , and 0 . 1 m dtt ) and heating for 10 min in boiling water before being loaded to the 4–15 % mini - protean® tgx™ precast gels ( biorad , 456–1084 ) . protein molecular weights were marked by the precision plus protein dual color standards ( biorad , # 1610374 ) . the electrophoresis was run at 100 volts for 60–90 min and the gels were stained by colloidal coomassie blue [ 35 ] or by the pierce silver staining kit ( pierce , # 24600 ) . equal percentages of the total volumes of collected proteins from non - activated and activated s . scapterisci ijs were loaded onto each gel . 10 % of protein volumes of activated ( total 210 μl x 1 . 08 μg / μl ) and non - activated ( total 200 μl x 0 . 008 μg / μl ) ijs were loaded onto the gel for colloidal coomassie blue staining ; for the silver staining , the volumes were 1 % of the total collected proteins .
conserved non - coding networks in steinernema and caenorhabditis . a a hierarchically clustered heat map of 30 derived regulatory motifs and the go terms that the target genes of these motifs are enriched in . only motif - go term associations that are shared between s . carpocapsae and c . elegans are shown . p - values depicted are from c . elegans associations . colored arrows point to single go term or groups of go terms that belong to the four developmental categories shown . b a network of conserved s . carpocapsae and c . elegans motif - target gene associations related to neurogenesis go terms . only nodes for motifs and downstream genes with degrees ≥5 are shown in the network . c a network of conserved s . carpocapsae and c . elegans motif - target gene associations related to muscle go terms . only nodes for motifs and downstream genes with degrees ≥5 are shown in the network . d zag - 1 gene model in s . carpocapsae and c . elegans showing conserved motifs , and well as conserved regulatory modules ( clusters of conserved motifs ) . sequence conservation tracks are displayed below each gene model . associations between zag - 1 and motifs are highlighted in red in the neurogenesis network . go gene ontology
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male , second generation : a second generation male is similar morphologically to that of the first generation except that it is about two - thirds as long and one - half as wide and the spicules have an elongate head .
female , first generation : anteriorly , the female is similar to the male but much larger . lateral fields and phasmids not observed . six labial papillae and four cephalic papillae present . stoma with prominent cheilorhabdions unusually thickened , appearing as a circular or hexagonal ring in face view . esophagus typical of family . esophago - intestinal valve large . excretory pore located anteriorly to mid - metacorpus .
excretory duct unusually prominent forming an elliptically - shaped structure seemingly with a hole at the center . gonads didelphic , amphidelphic , reflexed . vulva appears as a transverse slit with a prominent double - flapped epiptygma . tail length shorter than anal body width , with a mucron at end .
females , second generation : second generation female similar morphologically to that of the first generation with the following exceptions : about one - half as long and two - thirds as wide , valve in basal bulb of esophagus more prominent , elliptically - shaped structure less prominent , tail , which tapers to a point bearing a mucron , longer than body width at anus .
infective juveniles ( third stage ) : the infective stage , when newly formed , is always enclosed in the cuticle of the second - stage juvenile as a sheath . however , the sheath is lost rather easily , even in storage , and thus may not always be present . body thin , head with a labial raising disc ; lip region not offset , oral aperture not observed ; six labial , four cephalic papillae and an elevated oral disc prominent . esophagus degenerate and thus not seen clearly , but its basal bulb is elongate and has a valve . lateral field with six incisures . tail tapers gradually dorsally but abruptly ventrally .
three years after the nematode was released , a few mole crickets collected in a farm 10 miles from the nearest release site were found infected with the nematode . by the fifth year , 25 to 65 percent of the mole crickets collected at this farm were infected ( smart et al . 1990 ) .
in 1990 and 1991 , the nematode was released in pastures in six different counties and on nearly 30 golf courses in florida . it has become established and has reduced populations of mole crickets in all of the pastures .
mcsorley r . ( july 1997 ) . soil - inhabiting nematodes . uf / ifas featured creatures . eeny - 12 . ( august 1999 ) .
nguyen kb . ( may 1999 ) . mole cricket control by entomopathogenic nematodes . uf / ifas . ( no longer available online )
nguyen kb . ( february 1999 ) . taxonomy of entomopathogenic nematodes . uf / ifas . ( no longer available online )
nguyen kb . ( february 1998 ) . symbiotic bacteria of entomopathogenic nematodes . uf / ifas . ( no longer available online )
smart gc jr , nguyen kb . 1995 . biological control of orthoptera pest insects . united state patent , patent number 5 , 466 , 448 , date of patent : november , 14 1995 .
publication date : june 1999 . latest revision : june 2014 . reviewed : april 2017 .
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copyright : © 2017 lu et al . this is an open access article distributed under the terms of the creative commons attribution license , which permits unrestricted use , distribution , and reproduction in any medium , provided the original author and source are credited .
data availability : all relevant data are within the paper and its supporting information files .
funding : this work was supported by the national institutes of health , national institute of allergy and infectious diseases . k22 career transition award ai119155 to ard . the funders had no role in study design , data collection and analysis , decision to publish , or preparation of the manuscript .
galleria mellonella waxworms used for activation experiments were purchased from crittergrub ( urltoken ) . once received , waxworms were immediately sorted to remove dead ones , and the healthy waxworms were frozen at - 80°c until used . acheta domesticus house crickets ( american cricket ranch , lakeside , ca ) were immediately sorted to remove dead ones , and the healthy crickets were frozen at - 80°c until used . scapteriscus borellii mole crickets used for activation experiments were collected from the rio hondo golf course ( downey , ca ) as previously described [ 33 ] . adult mole crickets were fed a diet of wheat germ ( 15 g agar , 166 g wheat germ , 900–1000 ml h 2 o ) and water ad libitum until used . healthy adult mole crickets were frozen at - 80°c prior to use in activation experiments .
to recover nematodes from the small - scale activation assays , tap water was added to soak the sponge pieces in the petri dish for 5 min and the nematode suspension was transferred to a 15 ml tube . the process was repeated two more times and all nematodes were combined in one 15 ml tube . the nematodes were pelleted by centrifugation at 700 rcf for 1 min and washed 3 times with tap water . the supernatant was removed leaving ~ 1 . 5 ml of nematode suspension . aliquots of the nematode suspension were visualized with a compound microscope at the 10 x 40 magnification and scored for activation . higher magnification was occasionally used when the morphology was unclear . 3–4 aliquots were quantified per treatment meaning that 150–600 nematodes were used for each replicate .
the mouth , pharynx , and anterior gut are marked in these images by red arrows and labeled m , p , and g respectively . a . schematic drawing illustrating the morphological features of an activated ij on the left and a non - activated ij on the right . in the non - activated ij the pharynx and anterior gut can be difficult to visualize . b . two pictures showing the opaque morphology of non - activated ijs . the mouth is closed , the anterior gut is closed , and the pharyngeal bulb is not visible . c . two pictures showing the visible morphology of activated ijs . these fully activated ijs have visibly open mouths and stoma . they have expanded pharyngeal bulbs and the anterior gut is open . d . this picture shows a nematode scored as partially active because the pharyngeal bulb is only partially expanded .
ijs were exposed to insect homogenate for different amounts of time : 6 , 12 , 18 , 24 , and 30 hours . though a 3 - hour exposure was initially tested , it was too difficult to reliably distinguish morphological characteristics . for the time course we used 50 % acheta domesticus homogenate , made as described above .
statistical analysis was performed using graphpad prism 6 . 04 . standard statistical tests were used for all experiments , as described in the figure legends . one - way anovas were conducted to compare experimental effects among different exposure times to 50 % house cricket homogenate and the effects of the percentage of host homogenate on activation and partial activation . unpaired t tests were used to compare the experimental effects between two treatments within an experiment ( e . g . differences between two time points of the time course activation or differences between two homogenate concentrations within an experiment ) .
comparing activation rates for each type of host homogenate we found that for house cricket homogenate , 25 % , 50 % , and crude homogenate all produced significantly higher activation than 10 % homogenate ( p < 0 . 0001 ) ( fig 5a and s2 file ) . 50 % and crude house cricket homogenate both induced significantly higher activation than 25 % homogenate ( p < 0 . 0001 ) , and crude homogenate induced significantly higher activation than 50 % homogenate ( p = 0 . 04 ) . using mole cricket homogenate , there was no significant difference in activation between 10 % and 25 % homogenate , but 50 % and crude homogenate both induced significantly higher activation than 10 % or 25 % homogenate ( p≤0 . 03 ) ( fig 5b and s2 file ) . there was no significant difference between 50 % and crude homogenate . using waxworm homogenate , 25 % and 50 % homogenate both induced significantly more activation than 10 % homogenate ( p≤0 . 002 ) ( fig 5c ) , but there was no significant difference between 25 % and 50 % waxworm homogenate .
to study ij activation we first identified quantifiable morphologic changes that could be used to differentiate between activated and non - activated ijs ( fig 2 ) . we found that by examining the state of the mouth , pharyngeal bulb , and anterior gut , we could reliably distinguish between non - activated , partially activated , and fully activated ijs ( fig 2 ) . some early - activated ijs developed to l4 and young adult stages when incubated for more than 18 hours in cricket homogenate and in this study we categorized those as fully activated ijs . in future studies it may be informative to distinguish between activated ijs and nematodes that have developed beyond the activated l3 stage .
s2 file . activation data using different insect homogenates and different concentrations of homogenate .
we thank john rodriguez and the rio hondo golf course grounds crew for access to the grounds for collecting mole crickets ; sudarshan aryal and dennis chang for collecting and maintaining mole crickets and for providing feedback on the manuscript ; tiffany baiocchi , edwin lewis , and byron adams for insightful comments on the manuscript .
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ijs in the wells of a plastic cryo storage box with 100 slots designed for 1 . 5 ml microtubes ( e . g . , vwr cat # 40000–322 ) , with the lid removed . a sheet of transparent plastic was cut to fit the top of the box and cover all the wells to prevent crickets from escaping ; small holes were punched in the plastic for aeration . single layers of paper towels ( ~ 4 cm x 4 cm ) were pushed into the wells of the microtube storage box using a pen to make infection chambers . about 100 ijs of
) . the box was covered by the plastic sheet described above , which was pushed against the wells using a slightly smaller freezer box ( e . g . , vwr cat # 82007–162 ) . the boxes were securely bound together by rubber bands or tape . the infection boxes were incubated for 2 days in the dark at 25°c . dead crickets , presumably infected , were transferred to white traps [
] in which a piece of 5 . 5 cm filter paper was raised by the lid of a 3 . 5 cm petri dish in a 10 cm petri dish with a thin layer of tap water ( about 2 mm deep ) (
) — ( 1 ) the mouth being open or closed , ( 2 ) the state of the anterior gut , or the opening of the gut immediately posterior to the pharyngeal bulb , and ( 3 ) expansion of pharyngeal bulb and how pronounced it appeared . if an individual nematode had all three characteristics ( an open mouth , an open and expanded anterior gut , and an expanded and visible pharyngeal bulb ) they were considered fully activated (


hyles hippophaes ( seathorn hawk-moth ) is a species of moth in the family sphingidae .

the wingspan is 65 – 80 mm .

hyles hippophaes

hyles hippophaes hippophaes , male , upperside . france , basses alpes , st . andrer
hyles hippophaes hippophaes , male , underside . france , basses alpes , st . andrer
ovum : as subsp . hyles hippophaes hippophaes , with up to 500 being laid by each female .
synonym . hyles hippophaes miatleuskii eitschberger & saldaitis , 2000 , atalanta 31 : 213 .
habitat : in europe , hyles hippophaes settles hot , dry , stony habitats up to about 1300m above sea level , and preferably at the edge of wild floodplains close to or in the alps , where hippophaes grows in an early succession stage . hyles hippophaes inhabits also rocky slopes in these regions .
hyles hippophaes ( esper , 1789 ) = sphinx hippophaes esper , 1789 = crocea ( rebel , 1910 ) = flava ( denso , 1913 ) = obscurata ( dannehl , 1929 ) = teriolensis ( dannehl , 1929 ) = expallidata ( dannehl , 1933 ) = celerio hippophaes kiortsii koutsaftikis , 1974 .
as subsp . hyles hippophaes hippophaes ( esper , 1789 ) , northern spain , southern france , southern switzerland and northern italy to slovenia . then , as a separate population , romania , bulgaria , moldova , northern greece , the aegean islands and western turkey .
the early stages are very similar to subsp . hyles hippophaes hippophaes . fully grown , usually also very similar to those of subsp . hippophaes ; however , some are dark green with a dorsal lilac tint on the anterior segments and a broken , white ventro - lateral streak . in others , the basic body colour may even be dark grey / black .
pupa : similar to subsp . hyles hippophaes hippophaes ; during the summer months it remains in this stage for no more than 20 days . formed in a chamber in the soil , often up to 10cm deep ( chu & wang , 1980b ) . overwinters as a pupa .
celerio hippophaes kiortsii koutsaftikis , 1974 , annls mus . goulandris 2 : 99 - - 103 .
remarks : hyles hippophaes does not occur north of the alps . in europe , the moth is very rare in the alpine region , more frequently , for example , in the southwestern french alps , particularly in the valley of the durance . there i found , for example , caterpillars in mid - july 2005 and 2012 at xerothermic places . hyles hippophaes is distributed from the pyrenees locally across southern france , the southern alps , parts of italy ( apennines ) and obviously also of the balkans to the black sea . additionally , hyles hippophaes occurs from asia minor to central asia .
( ii ) there is some contact between subsp . hyles hippophaes hippophaes ( esper , 1789 ) and subsp . bienerti in the crimea and in western and southern turkey . this produces intermediate hybrids , such as f . malatiatus gehlen , 1934a , and f . kiortsii koutsaftikis , 1974 . the aegean population is most certainly subsp . hippophaes , but with a trace of subsp . bienerti visible in many examples . some adults of the crimean population are intermediate in coloration and pattern between subsp . hippophaes and subsp . bienerti . )
endangerment factors : hyles hippophaes is strongly threatened by river regulation , gravel and electricity industry , construction of roads , housing developments , agricultural intensivication ( creation of vineyarrds on rocky slopes in floodplains ) and by succession . most once mile - wide floodplains have been destroyed , the river forced into a straitjacket . this proved fatal for many specialist insects ( and other living beings ) such as many grasshoppers like bryodemella tuberculata . hyles hippophaes can be preserved only if the habitats are protected extensively . in southern central europe , hyles hippophaes is largely eradicated . last outpost is a residual presence in the rhone valley , valais , where it could also possibly be gone already .
from central turkey and the southern ukraine eastwards to liaoning ( china ) and mongolia , extending south to kashmir and north - west india and north to lake baikal and the tuva a . s . s . r . in russia as subsp . hyles hippophaes biernerti .
( taxonomic note . there is some contact between subsp . hippophaes and subsp . bienerti ( staudinger , 1874 ) in the crimea and in western and southern turkey . this produces intermediate hybrids , such as f . malatiatus gehlen , 1934a , and f . kiortsii koutsaftikis , 1974 . the aegean population is most certainly subsp . hippophaes , but with a trace of subsp . bienerti visible in many examples . some adults of the crimean population are intermediate in coloration and pattern between subsp . hippophaes and subsp . bienerti , but can be assigned to the latter . )
hyles hippophaes ( seathorn hawk - moth ) is a species of moth in the family sphingidae . it is found in afghanistan , armenia , azerbaijan , china , france , georgia , germany , greece , iran , iraq , kazakhstan , kyrgyzstan , mongolia , pakistan , romania , serbia and montenegro , spain , switzerland , syria , tajikistan , turkey , turkmenistan , and uzbekistan .
life cycle : hyles hippophaes forms one or two generations , but the second one is only partial . the moths fly from may to july and separated again in august / september . the larval time is from june to october , but by far the most caterpillars are observed in late june and july . the pupa hibernates as with all european sphingids ( except the migrant butterfly taubenschwänzchen ) .
transferred to hyles by hübner , [ 1819 ] , verz . bekannter schmett . : 137 . transferred to celerio by rothschild & jordan , 1903 , novit . zool . 9 ( suppl . ) : 714 ( key ) , 729 ; then back to hyles by pittaway , 1983 , entomologist ' s gaz . 34 : 80 . implicitly transferred to celerio by zhu & wang , 1997 , fauna sinica insecta 11 : 342 ( key ) , 344 . implicitly transferred back to hyles by danner , eitschberger & surholt , 1998 , herbipoliana 4 ( 1 ) : 202 .
it should be noted that in the french alps larvae of natural hybrids between this species and hyles vespertilio ( esper , 1780 ) are regularly found on epilobium dodonaei , with many being of the pink form ( j . - m . bompar , pers . comm . ) .
within its range , populations can be somewhat isolated . however , as this species is prone to wander , individuals may turn up at great distances from known breeding grounds , leading to confusing records . frequents river valleys in mountainous areas ( up to 500m in spain and switzerland ) , mountainous steppe and sand - dunes . river islands overgrown with hippophae rhamnoides are a favoured haunt in central europe . in some western european localities , as a result of river flood - control measures , hyles hippophaes is becoming increasingly rare as its hostplant cannot compete with riverine shrubs and trees which take over stabilized riverbanks .
after a very long period of inactivity on this blog , i’m finally posting some new shots . not “astro” ones , but nature - related ones . the following shots are of a hawk moth ( fam . sphingidae ) , hyles hippophaes ( esper , 1789 ) . this species is threatened by human activity , as many other insect species . in romania it occurs mainly in dobrogea , but can occasionally be found further to the west . the larva was found in august 2016 , and the adult emerged in may 2017 . the moth is active at night , and is attracted to artificial light .
synonymized with celerio euphorbiae as a subspecies by rothschild & jordan , 1903 , novit . zool . 9 ( suppl . ) : 720 ; then with hyles centralasiae as a subspecies by pittaway , 1983 , entomologist ' s gaz . 34 : 78 . reinstated as a species by danner , eitschberger & surholt , 1998 , herbipoliana 4 ( 1 ) : 205 , 280 .
wingspan : 60 - - 80mm . very unlikely to be confused with other hyles species . most individuals from the aegean population have more extensive black markings than those from central europe ; however , there is sufficient overlap in colour variation not to split the two groups into separate taxa . the original description and illustrations by esper ( [ 1789 ] ) clearly indicate that the romanian population is indistinguishable from that in the aegean . where multiple climatic conditions are present , such as along mountain chains , adults of this species are very variable in wingspan , markings and colour intensity . in fact , adult coloration is very much dependent on the temperature developing pupae are exposed to . heat produces more reddish and paler individuals , while low temperatures give rise to darker and greyer adults .
deilephila bienerti staudinger , 1874 , stettin . ent . ztg 35 : 91 . type locality : northeast persia [ iran ] , schahrud [ emamrud ] .
note . this subspecies can be very variable in both coloration and size where numerous climatic conditions occur in close proximity to each other , such as in mountainous areas . many of these forms were described as distinct subspecies but this is not warranted . subspecies ornatus , transcaucasica , anatolica , bucharana , shugnana , malatiatus , caucasica and baltistana were therefore synonymized with subsp . bienerti by pittaway ( 1993 ) and should be regarded as forms .
wingspan : 65 - - 80mm . considerably paler and browner than related subspecies . a pale , oblique median line is noticeable on the underside of the forewing ; hindwing patches more orange than red . some large specimens found above 2000m in north - west iran and kashmir tend to f . caucasica in coloration .
often common in mountainous , arid steppe , especially along rivers overgrown with hippophae or elaeagnus . although found at any altitude from 400 - - 3000m , most populations occur from 1000 - - 2000m where hippophae rhamnoides often forms discrete thickets away from rivers .
china : 29 . iv - 5 . v ( shihezi ) ; urltoken ( shihezi ) ; urltoken ( shaanxi ) ; 1 . vii ( zada ) ; vii ( yining / gulja , 1000m ) ; 5 - 31 . vii ( shihezi ) ; 28 . ix ( dingbian ) . mongolia : urltoken ( hovd prov . ) ; 19 . vii ( hovd prov . ) . russia : 19 - urltoken ( karasuk , novosibirsk area ) ; 14 . vii ( tuva ) ; 18 - 21 . vii ( altai ) ; 30 . viii ( karasuk , novosibirsk area ) .
there are two generations a year in northeastern and central china , with adults flying in may and july / august ( yang , 1978 ; chu & wang , 1982 ) . in xinjiang , the first brood emerges between late april and mid june , depending upon the weather .
ovum : pale greenish - grey , almost spherical ( 1 . 0 x 1 . 1mm ) . deposited on both the upper and under surface of leaves , usually near the edge , on the lower branches of the hostplant . thicket - edge or isolated shrubs are preferred , with up to 500 being laid by each female .
larva : full - fed 75 - - 80mm . dichromatic : unstriped or striped .
on hatching , the eggshell is ignored , the 3 - - 4mm - long larva proceeding to find a resting place below a leaf , a site to which it returns after each spell of feeding . at this stage it is dark green , thickly speckled with white and dark grey . the final instar has two colour forms . one is dark green ( in some cases suffused with pink ) , thickly speckled with white and grey ; superimposed on this are an off - white dorso - lateral line , often with orange eye - spots , and a broader , white , ventro - lateral stripe running just above the legs . horn long , thin , orange below , black above , with two elongated orange spots at its base ; head green , with two brown lines . the other colour form is silvery grey , with a black , broken dorso - lateral line from which emanate black , equally broken oblique lateral stripes with white , red , or yellow patches often present in between . head brown and grey ; horn as above . in a very rare colour form , all green coloration is replaced by pinkish brown .
larvae frequently sun themselves openly on the upper branches , amongst those they have already stripped of leaves . there is a very heavy mortality due to parasitoids . those that survive eventually become light brown before descending to find a pupation site , often after hours of perambulation on the ground .
this stage lasts as little as 28 days , during which the larva basks quite openly on the topmost branches of its hostplant .
pupa : 40 - - 50mm . pale yellowish brown , or light grey - brown , with dark brown striations . more elongated than others of the genus . enclosed in a flimsy cocoon amongst roots or under stones . the overwintering stage .
in the summer months it remains in this stage for no more than 20 days . formed in a chamber in the soil , often up to 10cm deep ( chu & wang , 1980b ) .
larval hostplants . recorded in china on elaeagnus angustifolia ( sensu lato ) and hippophae rhamnoides ( wang , 1978 ; pittaway & kitching , 2000 ) , which are also the hostplants in tajikistan ( shchetkin , 1956 ) .
china : xinjiang ( shihezi ; ürümqi ; yining / gulja ) ; nei mongol ( alxa zuoqi ; alxa youqi ) ; liaoning ; shaanxi ( dingbian ) ; ningxia ( yinchuan ; lingwu ; pingluo ) ; gansu ; xizang / tibet ( zada , 2950m ) .
mongolia hovd prov . ( bodonchijn river valley , elkhony - ekhen - tal , 30km s altai village , 1200m ( 45°43 ' n 92°05 ' e ) ; dzun - dzhargalant - khairkhan , ar - shatyn - gol ( river ) valley , 2100m ( 47°44 ' n 92°27 ' e ) ; bulgan river basin , bayan river valley , ulyastayn - sala river , 1900m ( 46°21 ' n 91°08 ' e ) ; bulgan river valley , 45km n bulgan , 1500m ; dzhungarian gobi , 45km sw bulgan , uvhod - ula mts . , 1350m ) .
russia : western siberia ( karasuk , novosibirsk area ) ; altai ( cherga ; 5km sww mikhailovskoe ) ; tuva ( khovu - aksy ) ; transbaikalia .
central ( rebel , 1933 ) , south - eastern and eastern turkey ( daniel , 1932 ; daniel , 1939 ; de freina , 2012 ) , southern ukraine ( efetov & budashkin , 1990 ; zolotuhin , pers . comm . ; vasilyuk & inozemtseva , 2003 ) , the caucasus and southern russia ( zolotuhin , pers . comm ; poltavsky , pers . comm 2003 ; anikin , 2004 ) , northern and central iran ( bienert , 1870 ; barou , 1967 ; kalali , 1976 ; ghassemi , alemansoor & alehossein , 2009 ) , turkmenistan ( danov & pereladov , 1985 ) and uzbekistan ( derzhavets , 1984 ) , the southern urals ( nupponen & fibiger , 2002 ; dubatolov , 2012 ) , eastern kazakhstan ( kondratiev coll . , nhmuk ) , western xinjiang province , china ( pittaway & kitching , 2000 ) , tajikistan ( shchetkin , 1956 ) , afghanistan ( ebert , 1969 ; daniel , 1971 ) , northern pakistan / azad kashmir ( karakoram mountains , juglot valley , 2550m , 26 . vii . 2011 ( leg . balázs benedek ) ) ( rafi et al . , 2014 ) , and north - west india / kashmir ( o . bang - haas , 1939 ) to the western tian shan .
also , china , from xinjiang province ( china ) north to the altai mountains ( izerskiy , 1999 ) and eastwards across the provinces of ningxia , gansu , shaanxi and nei mongol ( inner mongolia ) to liaoning ( chu & wang , 1980b ; pittaway & kitching , 2000 ) , and also mongolia ( derzhavets , 1977 ; yakovlev , gus ' kova , doroshkin & titov , 2015 ; yakovlev & doroshkin , 2017 ) . from these areas north to karasuk ( dubatolov , 2012 ) , the tuva a . s . s . r . ( viidalepp , 1979 ; izerskiy , 1999 ) , the altai kray ( yakovlev , dubatolov & titov , 2013 ) and lake baikal ( kondratiev coll . , nhmuk ) in russia .
holarctic ; palaearctic ( both eastern and western subregions ) . pleistocene refuge : polycentric - - caspian , iranian , turanoeremic , turkestan and mongoloeremic refugia .
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host plants : the caterpillar lives on sea buckthorn ( hippophae rhamnoides ) , in the southeast ( turkey ) also on other eleagnaceae ( eleagnus ) .
gb : seabuckthorn hawkmoth ; seathorn hawkmoth , f : sphinx de l ' argousier , d : sanddornschwärmer , ru : oblepikhovyi brazhnik ; yuzhnyi brazhnik , h : déli szender , e : esfinge del espino amarillo , fi : tyrnikiitäjä , it : sfinge dell ' olivella spinosa .
type locality : wallachei , milkowfluss bei foran [ wallachia region , southern romania ] .
holarctic ; western palaearctic region . pleistocene refuge : poly / monocentric - - adriatomediterranean and pontomediterranean subsections of the mediterranean refuge .
as with nearly all members of this genus , pairing is a short affair lasting not more than three hours , generally before midnight . afterwards , most females spend a few hours every night feeding , mainly before 23 . 00 hours and before dawn . oddly , this species does not fly very much and spends much of the night resting .
late april to early july , with a peak in mid - june . a partial second brood in august often occurs . it is not unusual for only three weeks to elapse between the two broods .
ovum : almost spherical ( 1 . 1 x 1 . 0mm ) , pale greenish grey . deposited on both the upper and under surface of leaves , usually near the edge , on the lower branches of the hostplant . thicket - edge or isolated shrubs are preferred , most eggs being laid in late june .
larva : full - fed , 75 - - 80mm . polymorphic : unstriped or striped .
on hatching , the eggshell is ignored , the 3 - - 4mm - long larva proceeding to find a resting place below a leaf , a site to which it returns after each spell of feeding . at first only the cells on the leaf upperside are eaten , leaving clear ' windows ' in the leaf ; in the second instar leaves are consumed in the normal fashion . initially pale grey with a white dorso - lateral line and grey horn , it gradually becomes dark green , thickly speckled with white and dark grey .
the final instar has several colour forms . the main one is dark green ( in some cases suffused with pink ) , thickly speckled with white and grey ; superimposed on this are an off - white dorso - lateral line , often with orange eye - spots , and a broader , white , ventro - lateral stripe running just above the legs . horn long , thin , orange below , black above , with two elongated orange spots at its base ; head green , with two brown lines .
a less common form is silvery grey , with a black , broken dorso - lateral line from which emanate black , equally broken oblique lateral stripes with white , red , or yellow patches often present in between . head brown and grey ; horn as above .
there are also two very rare colour forms in which all green coloration is replaced by either pinkish brown or dark grey / black . the latter appears more readily under cold conditions .
larvae frequently sun themselves openly on the upper branches , amongst those they have already stripped of leaves . there is a very heavy mortality due to parasitoids . those that survive eventually become light purple - brown before descending to find a pupation site , often after hours of perambulation on the ground .
most common during late june and july ; in some areas also during early september .
minor hostplants . elaeagnus angustifolia , an introduced oleaster from central and eastern turkey now established over much of southern europe . this is the main hostplant of the aegean population ( pittaway , 1982a ) . [ in captivity , the larvae will thrive on many species of ornamental elaeagnus , and will also accept epilobium angustifolium when larger . ]
pupa : 40 - - 50mm . pale yellowish brown , or light grey - brown , with dark brown striations . more elongated than others of the genus . enclosed in a flimsy yellowish cocoon amongst roots or under stones . the overwintering stage .
tachinidae : exorista fasciata ( fallén ) , exorista larvarum ( linnaeus ) , exorista grandis ( zetterstedt ) , masicera sphingivora ( robineau - desvoidy ) .
separated into two main areas which seem to be the remnants of a much larger , post - glacial range . from northeastern spain ( portbou , catalonia ( pittaway , 1983b ) ) across southern france ( frionnet , 1910 ; chanselme , 1997 ) , southern switzerland ( schweizerischer bund für naturschutz , 1997 ) and northern italy to slovenia . then , as a separate population , romania ( esper , [ 1793 ] ; székely & szabó , 1995 ; vlad dinca , pers . comm . 2007 ) , bulgaria ( beshkov , 1998 ; danner , eitschberger & surholt , 1998 ) , moldova ( derzhavets , 1984 ) , northern greece ( koutsaftikis , 1970 ; 1973 ; 1974 ) , the aegean islands ( de freina & piatkowski , 1999 ) and western turkey ( pittaway , 1982a ) .
this species will very probably also be found in more areas of northern spain , hungary and romania , countries with large areas of hostplant . it has been recorded as a vagrant in england ( gilchrist , 1979 ) , northwestern spain ( basque country ( gómez bustillo & fernández - rubio , 1976 ) ) , southern spain ( malaga ( ribbe , 1909 - 1912 ) ) , southern portugal ( near faro ) and slovakia ( danner , eitschberger & surholt , 1998 ) . recently recorded from the apennine mountains east of florenz , italy ( dapporto , fiorini , fiumi & flamigni , 2005 ) .
recorded in the past from germany ( bavaria ) ( heinemann , 1859 ; forster & wohlfahrt , 1960 ) , but no longer resident in that country ( danner , eitschberger & surholt , 1998 ) .
this species appears on the british list as a result of a supposed vagrant record from devon in approximately 1857 .
, and fly between april and june , sometimes with a second generation in august .
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die schmetterlinge in abbildungen nach der natur mit beschreibungen . fortsetzung der europäischer schmetterlinge 6
lectotype ♀ [ romania : ] wallechei [ wallachia region ] , near foxan , milkowfluss [ milkov river ] [ wmhg ] ; designated by hacker , 1999 , esperiana 7 : 453 . the lectotype is the largest female (
das größere weibchen
) in
kasten 20
.
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( taxonomic notes . ( i ) this subspecies can be very variable in both coloration and size where numerous climatic conditions occur in close proximity to each other , such as in mountainous areas . many of these forms were described as distinct subspecies but this is not warranted . subspecies ornatus , transcaucasica , anatolica , bucharana , shugnana , malatiatus , caucasica and baltistana have been synonymized with subsp . bienerti and should be regarded as forms .
wingspan : 65 - - 80mm . can be considerably paler and browner than related subspecies . a pale , oblique median line is noticeable on the underside of the forewing ; hindwing patches more orange than red . some large specimens found above 2000m in north - west iran and kashmir tend to f . caucasica in coloration .
often common in mountainous , arid steppe , especially along rivers overgrown with hippophae or elaeagnus . although found at any altitude from 400 - - 3000m , most populations occur from 1000 - - 2000m where hippophae rhamnoides often forms discrete thickets away from rivers . attracted to the flowers of cistanche at dusk ( shchetkin , 1956 ) .
larva : full - fed , 75 - - 85mm . dimorphic : unstriped or striped .
major hostplants . hippophae rhamnoides and elaeagnus spp . , especially elaeagnus angustifolia and elaeagnus hortensis in tajikistan ( shchetkin , 1956 ) . in the crimea it is found on elaeagnus argentea ( efetov & budashkin , 1990 ) .
central ( rebel , 1933 ) , south - eastern and eastern turkey ( daniel , 1932 ; daniel , 1939 ; de freina , 2012 ) , southern ukraine ( efetov & budashkin , 1990 ; zolotuhin , pers . comm . ; vasilyuk & inozemtseva , 2003 ) , the caucasus and southern european russia ( zolotuhin , pers . comm ; poltavsky , pers . comm . 2003 ; anikin , 2004 ) , the republic of georgia ( didmanidze , petrov & zolotuhin , 2013 ) , armenia ( wąsala & zamorski , 2015 ) and azerbaijan ( didmanidze , petrov & zolotuhin , 2013 ) , northern and central iran ( bienert , 1870 ; barou , 1967 ; kalali , 1976 ; ghassemi , alemansoor & alehossein , 2009 ; lehmann & zahiri , 2011 ) , turkmenistan ( danov & pereladov , 1985 ) and uzbekistan ( derzhavets , 1984 ) , the southern urals ( nupponen & fibiger , 2002 ; dubatolov , 2012 ) , eastern kazakhstan ( kondratiev coll . , nhmuk ; shovkoon , 2015 ) , western xinjiang province , china ( pittaway & kitching , 2000 ) , tajikistan ( shchetkin , 1956 ) , afghanistan ( ebert , 1969 ; daniel , 1971 ) , northern pakistan / azad kashmir ( karakoram mountains , juglot valley , 2550m , 26 . vii . 2011 ( leg . balázs benedek ) ( rafi et al . , 2014 ) ) , and north - west india / kashmir ( o . bang - haas , 1939 ) to the western tian shan .
as elaeagnus angustifolia is widely planted as a hedge and windbreak throughout eastern europe and central asia , this moth has not only expanded its range northwards ( dubatolov , 2012 ) but may turn up as a vagrant far from its resident range , e . g . the northern ukraine ( plyushch & sheshurak , 1997 ) .
the anatolian plateau forms the western resident limit of this subspecies , although individuals can penetrate as far west as istanbul .
extra - limital range . in china , from xinjiang province ( china ) north to the altai mountains ( izerskiy , 1999 ) and eastwards across the provinces of ningxia , gansu , shaanxi and nei mongol ( inner mongolia ) to liaoning ( chu & wang , 1980b ; pittaway & kitching , 2000 ) , and also mongolia ( derzhavets , 1977 ; yakovlev & doroshkin , 2017 ) . from these areas north into russia to karasuk ( dubatolov , 2012 ) , the tuva a . s . s . r . ( viidalepp , 1979 ; izerskiy , 1999 ) , the altai kray ( yakovlev , dubatolov & titov , 2013 ) and lake baikal ( kondratiev coll . , nhmuk ) .
lectotype ♂ [ turkey : toros mountains , ] bulghar dagh [ bolkar dağlari ] , lydia , [ bred ex larvae ( w . siehe ) ] [ mnhu ] ; implicitly designated by danner , eitschberger & surholt , 1998 , herbipoliana 4 ( 2 ) : 74 .
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2011 - 01 - 26 by & van nieukerken , dr erik j . karsholt , dr ole & by dr ian kitching
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austria , bulgaria , germany , greece , spain , italy , corsica , romania , the soviet union - the european part of turkey - european part of france , switzerland , yugoslavia .
regions of the russian federation : the volga - don , east caucasus , western caucasus , lower volga , tuva , southern urals .
bulgaria , the british isles , hungary ? , germany , greece ( mainland ) , spain ( mainland ) ? , italy ( mainland ) , corsica , moldova , russia , romania , north aegean , slovenia , ukraine , france ( mainland ) croatia , switzerland .
[ 10 ] de jong , y . s . d . m . ( ed . ) ( 2011 ) fauna europaea version 2 . 4 ( faunaeur . org )
[ 28 ] moths and butterflies of europe and north africa ( leps . it ) , 2012
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curators : konstantin efetov , vasiliy feoktistov , svyatoslav knyazev , evgeny komarov , stan korb , alexander zhakov .
species catalog enables to sort by characteristics such as expansion , flight time , etc . .
the wingspan is 65–80 mm . subspecies bienerti is paler and browner than related subspecies . a pale , oblique median line is noticeable on the underside of the forewing . the hindwing patches are more orange than red .
larvae of subspecies bienerti have been recorded on elaeagnus angustifolia and hippophae rhamnoides in china and tajikistan .