schaefferia profundissima is a species of springtail ( arthropods ) endemic to the krubera-voronja cave system in georgia .

it is one of the deepest terrestrial animal ever found on earth , living at > 1,800 metres ( 5,900 ft ) below the cave entrance .

it was discovered in the cavex team expedition of 2010 .

schaefferia profundissima

.
a new species of schaefferia absolon , 1900 ( collembola : hypogastruridae )
.
key ; deepest cave of the world ; deuteraphorura kruberaensis n . sp . ; biospeleology ; cavernicolous fauna ; anurida stereoodorata n . sp . ; plutomurus ortobalaganensis n . sp . ; schaefferia profundissima n . sp .
in the black sea , researchers have found springtails in a deep cave – plutomurus ortobalaganensis at 1980 metres and schaefferia profundissima at 1600 meters . springtails are hexapods ( have six legs ) and these both like cheese !
.
trois espèces nouvelles du genre schaefferia absolon , 1900 ( insecte , collembole )
.
.
une nouvelle espèce de schaefferia d’une grotte des etats - unis d’amérique ( collembola , hypogastruridae )
.
amphipoda zenkevitchia sp . collembola schaefferia profundissima jordana & baquero , 2012 anurida stereoodorata jordana & baquero , 2012 deuteraphorura kruberaensis jordana & baquero , 2012 plutomurus ortobalaganensis jordana & baquero , 2012 coleoptera carabus sp . duvalius abyssimus reboleira & ortuño , 2014 catops cavicis giachino , 2011 diptera trichocera ( saltrichocera ) maculipennis ( meigen 1818 )
described by rafael jordana and enrique baquero from university of navarra ( spain ) , they are known for science as : anurida stereoodorata , deuteraphorura kruberaensis , schaefferia profundissima and plutomurus ortobalaganensis . the last one is the deepest arthropod ever found , at the remarkable depth of 1 . 980 meters ( 2 , 165 yards ) below ground surface .
the arthropod , known as plutomurus ortobalaganensis , was discovered 1980 metres below the surface , where it feeds off fungi and other decaying matter . three other new species were also found lurking in the cave : anurida stereoodorata , deuteraphorura kruberaensis and schaefferia profundissima . all four species have been classified as springtails , a type of small primitive wingless insect . living in total darkness , the species all lack eyes . however , a . stereoodorata compensates for this with a highly specialised form of chemoreceptor .
jordana , rafael , baquero , enrique , reboleira , ana sofia & sendra , alberto , 2012 , reviews of the genera schaefferia absolon , 1900 , deuteraphorura absolon , 1901 , plutomurus yosii , 1956 and the anurida laboulbène , 1865 species group without eyes , with the description of four , terrestrial arthropod reviews 5 , pp . 35 - 85 : 41
more information : reviews of the genera schaefferia absolon , 1900 , deuteraphorura absolon , 1901 , plutomurus yosii , 1956 and the anurida laboulbène , 1865 species group without eyes , with the description of four new species of cave springtails ( collembola ) from krubera - voronya cave , arabika massif , abkhazi , terrestrial arthropod reviews , volume 5 ( 2012 ) pp . 1 - 51
html public
- / / w3c / / dtd xhtml 1 . 0 strict / / en
urltoken
this site uses cookies . by continuing to browse the site you are agreeing to our use of cookies .
brill’s mybook program is exclusively available on brillonline books and journals . students and scholars affiliated with an institution that has purchased a brill e - book on the brillonline platform automatically have access to the mybook option for the title ( s ) acquired by the library . brill mybook is a print - on - demand paperback copy which is sold at a favorably uniform low price .
< ? xml version =
1 . 0
encoding =
utf - 8
? > < event > < eventlog > < portalid > brill < / portalid > < sessionid > vgcvjk2mti - gqyzfgjxurler . x - brill - live - 03 < / sessionid > < useragent > python / 3 . 5 aiohttp / 3 . 3 . 2 < / useragent > < identityid > guest < / identityid > < identity _ list > guest < / identity _ list > < ipaddress > 35 . 232 . 247 . 97 < / ipaddress > < eventtype > personalisation < / eventtype > < createdon > 1531172319074 < / createdon > < / eventlog > < eventlogproperty > < item _ id > urltoken < / item _ id > < type > favourite < / type > < / eventlogproperty > < / event >
1 : 1department of zoology and ecology , university of navarra , 31080 pamplona , spain e - mails : rjordana @ urltoken ; ebaquero @ urltoken ; 2 : 2department of biology , universidade de aveiro and cesam campus universitário de santiago , 3810 - 193 aveiro , portugal e - mail : sreboleira @ urltoken ; 3 : 3museu valencià d’història natural ( fundación entomológica torres sala ) paseo de la pechina 15 . 46008 valencia , spain e - mail : alberto . sendra @ urltoken
.
the taxonomy and distribution of the genus anurida ( collembola : neanuridae ) in the northern palaearctic
.
.
new palaearctic species of the genus anurida ( collembola , neanurida )
.
.
nuevas especies cavernícolas del género onychiurus del grupo de o . boneti gisin , 1953 ( collembola : onychiuridae ) del karst de navarra y gipuzkoa ( españa )
.
.
nuevos generos y especies de hipogastruridos de mexico ( collembola )
.
.
über einige theilweise neue collembolen aus den höhlen der gegen von letmathe in westfalen
.
.
sur un essai de classification des neanuridae holarctiques et sur quelques espèces de ce groupe ( collembola )
.
.
faune française des collemboles ( x ) espèces nouvelles ou peu connues des pyrénées et du sud - ouest
.
.
faune française des collemboles ix . les hypogastrura a . l . du massif du néouvielle ( hautes - pyrénées )
.
.
the collembola of north america , north of the rio grande
.
.
a new nearctic species of the genus tomocerus ( collembola : entomobryidae )
.
.
sur quelques collemboles de la région de banyuls ( pyrénées - orientales ) avec la description d’une espèce troglobie
.
.
a new species of the genus plutomurus yosii , 1956 ( collembola , tomoceridae ) from georgian caves
.
.
anurida frigida . a new species of collembola ( hypogastruridae ) from swedish lappland
.
.
arctic collembola . i – alaskan collembola of the families poduridae , hypogastruridae , odontellidae , brachystomellidae and neanuridae
.
north american springtails of the subfamily tomocerinae . proceedings of the united states national museum
.
notes sur les collemboles , avec description de cinq espèces nouvelles découvertes dans le canton de genève
.
.
a new species of the genus anurida laboulbene , 1985 ( collembola , neanuridae ) from romania
.
.
especies ibéricas de hypogastrura ( ceratophysella ) de seis ojos con descripción de tres nuevas especies ( collembola , hypogastruridae )
.
.
le genre plutomurus en russie et en géorgie ( collembola , tomoceridae )
.
.
etude de la faune coréenne des insectes collemboles vi . sur la famille des tomoceridae , édaphiques avec la description de quatre nouvelles espèces et d’une nouvelle sous - espèce
.
.
new species of springtails of the genus hypogastrura s . l . in the north - eastern asia fauna
.
.
onychiurus ( onychiurus ) diaelleni sp . n . , eine neue collembolenart aus der “raudner - höhle” ( steiermark )
.
.
nuevos hypogastruridae anoftalmos ( collembola ) de cuevas y suelos de méxico
.
.
plutomurus carpaticus sp . n . ( collembola : tomoceridae ) from the carpathian mountains
.
.
a new humicolous species of plutomurus ( collembola , tomoceridae ) from hokkaido , north japan
.
.
two new species of the family pseudachorutidae from mt . jizu , western yunnan , southwest china ( insecta : collembola )
.
.
un nouveau genre d’insectes collemboles onychiuridae cavernicoles des picos de europa ( espagne )
.
.
new species of the springtail genus anurida laboulb . ( collembola , neanuridae ) from the asiatic part of the ussr
.
.
generic revision of onychiurinae ( collembola : onychiuridae ) with a cladistic analysis
.
.
about the proserpinae group of hypogastrura ( collembola ) in the caves of pref
.
.
studies on the collembolan family tomoceridae , with special reference to japanese forms
.
.
two new species of collembola ( arthropleona : neanuridae , pseudachorutidae ) from shanghai , china
.
josé h . schoereder ; tathiana g . sobrinho ; marcelo s . madureira ; carla r . ribas and paulo s . oliveira
subterranean community of krubera - voronya cave
by alberto sendra and ana sofia p . s . reboleira
scholar commons > usf libraries > environmental sustainability > ijs > vol . 41 ( 2012 ) > iss . 2
alberto sendra , museu valencià d ' història natural ( fundación entomológica torres sala ) paseo de la pechina 15 . 46008 valencia , spain follow ana sofia p . s . reboleira , universidade de aveiro , portugal follow
subsurface biota extends over a wide variety of habitats that can be spatially interconnected . the largest communities of this subsurface biota inhabit cavities and are well known mainly in caves where biologists are able to have access . data about deep subterranean communities and arthropods living under one thousand meters was unknown .
the biocoenosis and the vertical distribution of invertebrate fauna of krubera - voronja are provided , from its entrance to the remarkable depth of 2140 meters , including the discovery of world’s deepest dwelling arthropod .
sendra , alberto and ana sofia p . s . reboleira . 2012 . the world’s deepest subterranean community - krubera - voronja cave ( western caucasus ) . international journal of speleology , 41 : 221 - 230 . available at : urltoken
© 2018 american association for the advancement of science . all rights reserved . aaas is a partner of hinari , agora , oare , chorus , clockss , crossref and counter .
this treatment is a stub . you can help plazi making the full treatment available by uploading the source publication .
no known copyright restrictions apply . see agosti , d . , egloff , w . , 2009 . taxonomic information exchange and copyright : the plazi approach . bmc research notes 2009 , 2 : 53 for further explanation .
the deepest - dwelling land animal in the world has been found almost 2 kilometres underground . fittingly , its home is krubera - voronja , the world ' s deepest cave , whose bottommost point is 2191 metres below its mouth . the cave is located near the black sea in abkhazia , a breakaway republic of georgia .
the animals were discovered by ana sofia reboleira from the university of aveiro , portugal , and alberto sendra of the valencian museum of natural history , spain . they were exploring the krubera - voronja cave as part of a 2010 expedition led by the ibero - russian cavex team .
before this discovery , springtails had been found just half a kilometre below ground . in 1986 , ongulonychiurus colpus was found living 550 metres down in spanish caves , and last year , tritomurus veles was found at 430 metres in caves in croatia . the discovery of species living deep underground in total darkness gives new insights into the extreme conditions in which animals can survive . [ new scientist ]
new scientist reports , explores and interprets the results of human endeavour set in the context of society and culture , providing comprehensive coverage of science and technology news .
kinja is in read - only mode . we are working to restore service .
html public
- / / w3c / / dtd xhtml 1 . 1 / / en
urltoken
enter your email address to follow this blog and receive notifications of new posts by email .
privacy & cookies : this site uses cookies . by continuing to use this website , you agree to their use . to find out more , including how to control cookies , see here : cookie policy
sendra , a . & reboleira , a . s . p . s . ( 2012 ) . the world’s deepest subterranean community - krubera - voronja cave ( western caucasus ) .
an expedition to world’s deepest cave , krubera - voronja in western caucasus , revealed an interesting subterranean community , living below 2000 meters and represented by more than 12 species of arthropods , including several new species for science . this deep cave biota is composed of troglobionts and also epigean species , that can penetrate until - 2140 m .
invertebrate fauna found in krubera - voronja cave : pseudoescorpiones neobisium ( blothrus ) birsteini lapschoff , 1940 opiliones nemaspela sp . araneae troglohyphantes sp . araneae gnaphosidae sp . indet acari sp . indet diplopoda chordeumatida sp . indet leucogeorgia sp . decapoda troglocaris sp .
scientists have recently described the deepest terrestrial animal ever found , together with 4 new species for science . these animals are springtails ( arthropoda , insecta , collembola ) , a minute primitive wingless insect with six - legs and without eyes that live in total darkness .
these animals were collected during the biospeleological works of sofia reboleira , from the university of aveiro ( portugal ) and alberto sendra , from the valencian museum of natural history ( spain ) , both who were members of the ibero - russian cavex team expedition to the world ' s deepest cave , during the summer of 2010 .
the world ' s deepest cave , krubera - voronja , reaching nowadays the remarkable depth of - 2 . 191 meters below ground level , is located in abkhazia , a remote area near the black sea in the mountains of western caucasus , being the only cave in the world with more than 2 kilometres of depth .
. in total absence of light and extreme low food resources , cave - dwelling animals have unique adaptations to subterranean life . they lack body pigmentation , they have no eyes and have been developing morpho - physological strategies for survival at such depth , during millions of years . one of the species has , for example , a spectacular chemoreceptor , a highly specialised type of the habitual post - antennal organ of the springtails .
last week the international journal of myriapodology published the first population genetic study of cave millipedes . this research highlights an important challenge in the conservation of cave biodiversity  that for . . .
( physorg . com ) - - aberdeen scientists have photographed for the first time fish and shrimps at europes deepest point - - 5111 meters or 3 . 2 miles deep below the surface of the mediterranean sea .
( physorg . com ) - - it waits blindly in the darkness of granite caves in yosemite national park , moving little to conserve energy .
a northern arizona university doctoral candidate and a national park service researcher have discovered a new genus of cave cricket .
a research team led by professor michael chazan , director of the university of toronto ' s archaeology centre , has discovered the earliest evidence of our cave - dwelling human ancestors at the wonderwerk cave in south africa .
they are dark , sometimes forbidding landscapes molded by volcanic eruptions or subterranean streams , but caves are also home to a host of creatures strangely adapted to the underworld .
the co - evolution of plant—pathogen interactions has been revealed in unprecedented detail in a study of one of the world ' s deadliest crop killers . this is the rice blast pathogen , which destroys enough food to feed more . . .
using genome editing to inactivate a protein called pcsk9 effectively reduces cholesterol levels in rhesus macaques , a species of monkey , according to researchers from the perelman school of medicine at the university of . . .
fish that ' farm ' their own patches of seaweed alter their ' cropping ' practices under high co2 conditions , researchers at the university of adelaide in australia have found .
mucus is able to protect us from infection thanks to ancient genes that have been conserved throughout 350 million years of evolution—dating back to our days as a jellyfish .
by proving a theory that was first proposed almost 40 years ago , researchers have confirmed a new way that cells conserve energy .
our bodies consist of many different kinds of cells , each with their own role . the japanese scientist shinya yamanaka had made earlier the discovery , earning the nobel prize in 2012 , that cells from adult skin can be converted . . .
please sign in to add a comment . registration is free , and takes less than a minute . read more
html public
- / / w3c / / dtd html 4 . 0 transitional / / en


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
html public
- / / w3c / / dtd xhtml 1 . 0 strict / / en
urltoken
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 .
by switching countries your current shopping cart will be cleared . are you sure ?
your account has been locked . enter your user name and click submit . an email will be sent to you with instructions .
don ' t see what you are looking for ? our technical support team may be able to help .
biosafety classification is based on u . s . public health service guidelines , it is the responsibility of the customer to ensure that their facilities comply with biosafety regulations for their own country .
this material is cited in a us or other patent and may not be used to infringe the claims . depending on the wishes of the depositor , atcc may be required to inform the patent depositor of the party to which the material was furnished .
customers located in the state of hawaii will need to contact the hawaii department of agriculture to determine if an import permit is required . a copy of the permit or documentation that a permit is not required must be sent to atcc in advance of shipment .
discover a faster , simpler path to publishing in a high - quality journal . plos one promises fair , rigorous peer review , broad scope , and wide readership – a perfect fit for your research every time .
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 .
kaya hk , gaugler r . entomopathogenic nematodes . annu rev entomol . 1993 ; 38 : 181–206 .
lewis ee , clarke dj . nematode parasites and entomopathogens . in : vega fe , kaya hk , editors . insect pathology . 2nd ed : elsevier ; 2012 . p . 395–424 .
dillman ar , guillermin ml , lee jh , kim b , sternberg pw , hallem ea . olfaction shapes host - parasite interactions in parasitic nematodes . proc natl acad sci usa . 2012 ; 109 ( 35 ) : e2324–33 . epub 2012 / 08 / 02 . pubmed central pmcid : pmc3435218 . pmid : 22851767
hallem ea , dillman ar , hong av , zhang y , yano jm , demarco sf , et al . a sensory code for host seeking in parasitic nematodes . current biology . 2011 ; 21 : 377–83 . pmid : 21353558
castelletto ml , gang ss , okubo rp , tselikova aa , nolan tj , platzer eg , et al . diverse host - seeking behaviors of skin - penetrating nematodes . plos pathogens . 2014 ; 10 ( 8 ) : e1004305 . pubmed central pmcid : pmc4133384 . pmid : 25121736
gang ss , hallem ea . mechanisms of host seeking by parasitic nematodes . molecular and biochemical parasitology . 2016 ; 208 ( 1 ) : 23–32 . pubmed central pmcid : pmcpmc4993646 . pmid : 27211240
morran lt , penley mj , byrd vs , meyer aj , o ' sullivan ts , bashey f , et al . nematode - bacteria mutualism : selection within the mutualism supersedes selection outside of the mutualism . evolution ; international journal of organic evolution . 2016 ; 70 ( 3 ) : 687–95 . pubmed central pmcid : pmcpmc4801668 . pmid : 26867502
murfin ke , dillman ar , foster jm , bulgheresi s , slatko be , sternberg pw , et al . nematode - bacterium symbioses - cooperation and conflict revealed in the
omics
age . biol bull - us . 2012 ; 223 ( 1 ) : 85–102 .
dillman ar , chaston jm , adams bj , ciche ta , goodrich - blair h , stock sp , et al . an entomopathogenic nematode by any other name . plos pathogens . 2012 ; 8 ( 3 ) : e1002527 . pmid : 22396642
conditions . j invertebr pathol . 2000 ; 75 : 55–8 . pmid : 10631058
sicard m , le brun n , pages s , godelle b , boemare n , moulia c . effect of native
hosts : contrasting types of interaction . parasitology research . 2003 ; 91 ( 6 ) : 520–4 . pmid : 14557877
sicard m , ramone h , le brun n , pages s , moulia c . specialization of the entomopathogenic nematode
bonifassi e , saux mfl , boemare n , lanois a , laumond c , smart g . gnotobiological study of infective juveniles and symbionts of
: a model to clarify the concept of the natural occurrence of monoxenic associations in entomopathogenic nematodes . j invertebr pathol . 1999 ; 74 ( 2 ) : 164–72 . pmid : 10486229
emelianoff v , sicard m , le brun n , moulia c , ferdy jb . effect of bacterial symbionts
nguyen kb . a new nematode parasite of mole crickets : its taxonomy , biology and potential for biological control . gainesville , florida : university of florida ; 1988 .
mole crickets . in : hajek ae , editor . use of microbes for control and eradication of invasive arthropods . progress in biological control . 6 . the netherlands : springer ; 2009 . p . 115–31 .
frank jh , walker tj . permanent control of pest mole crickets ( orthoptera : gryllotalpidae :
nguyen kb , hunt dj , mracek z . steinernematidae : species and descriptions . in : nguyen kb , hunt dj , editors . entomopathogenic nematodes : systematics , phylogeny and bacterial symbionts . boston : brill ; 2007 . p . 121–609 .
poinar go jr . nematodes for biological control of insects . boca raton : crc press ; 1979 .
adams bj , peat sm , dillman ar . phylogeny and evolution . in : nguyen kb , hunt dj , editors . entomopathogenic nematodes : systematics , phylogeny , and bacterial symbionts . nematology monographs and perspectives . 5 . leiden - boston : brill ; 2007 . p . 693–733 .
price pw . evolutionary biology of parasites . princeton , nj : princeton university press ; 1980 .
thompson jn . the coevolutionary process . chicago , il : university of chicago press ; 2009 .
kaya hk , stock sp . techniques in insect nematology . in : lacey l , editor . manual of techniques in insect pathology . san diego , ca : academic press limited ; 1997 .
mcmullen jg , stock sp . in vivo and in vitro rearing of entomopathogenic nematodes ( steinernematidae and heterorhabditidae ) . jove - j vis exp . 2014 ; ( 91 ) : e52096 .
white gf . a method for obtaining infective nematode larvae from cultures . science . 1927 ; 66 ( 1709 ) : 302–3 .
dillman ar , cronin cj , tang j , gray da , sternberg pw . a modified mole cricket lure and description of
( orthoptera : gryllotalpidae ) range expansion and calling song in california . environmental entomology . 2014 ; 43 ( 1 ) : 146–56 . pmid : 24472207
bedding ra . low cost in vitro mass production of neoaplectana and heterorhabditis species ( nematoda ) for field control of insect pests . nematologica . 1981 ; 27 : 109–14 .
neuhoff v , arold n , taube d , ehrhardt w . improved staining of proteins in polyacrylamide gels including isoelectric - focusing gels with clear background at nanogram sensitivity using coomassie brilliant blue g - 250 and r - 250 . electrophoresis . 1988 ; 9 ( 6 ) : 255–62 . pmid : 2466658
lee jh , dillman ar , hallem ea . temperature - dependent changes in the host - seeking behaviors of parasitic nematodes . bmc biology . 2016 ; 14 .
cassada rc , russell rl . the dauerlarva , a post - embryonic developmental variant of the nematode
balasubramanian n , hao yj , toubarro d , nascimento g , simoes n . purification , biochemical and molecular analysis of a chymotrypsin protease with prophenoloxidase suppression activity from the entomopathogenic nematode
toubarro d , avila mm , montiel r , simoes n . a pathogenic nematode targets recognition proteins to avoid insect defenses . plos one . 2013 ; 8 ( 9 ) : e75691 . pubmed central pmcid : pmc3787073 . pmid : 24098715
lewis ee , grewal ps , gauger r . hierarchical order of host cues in parasite foraging strategies . parasitology . 1995 ; 110 : 207–13 .
lewis ee , ricci m , gaugler r . host recognition behavior predicts host suitability in the entomopathogenic nematode
dillman ar , sternberg pw . entomopathogenic nematodes . current biology : cb . 2012 ; 22 ( 11 ) : r430–1 . epub 2012 / 06 / 09 . pmid : 22677279
do these subject areas make sense for this article ? click the target next to the incorrect subject area and let us know . thanks for your help !
html public
- / / w3c / / dtd xhtml 1 . 0 transitional / / en
urltoken
warning : the ncbi web site requires javascript to function . more . . .
the full text of this article is available as a pdf ( 1 . 7m ) .
the full text of this article is available as a pdf ( 381k ) .
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 .
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 (