mycocepurus smithii is an attine fungus-growing ant from latin america .

while a widely reported study from 2009 claimed that the species consists exclusively of females which reproduce asexually a latter work in 2011 found evidence of sexual reproduction in some populations .

in many colonies queen reproduces by parthenogenesis and all other ants in a colony are female clones of the queen .

the ants cultivate a garden of fungus inside their colony grown with pieces of dead vegetables and other insects .

it is this capacity for farming which initially prompted research into the species as a basal genus member would provide insight into the natural history of the fungal-cultivating ant tribe , attini .

mycocepurus smithii

senior synonym of mycocepurus attaxenus , mycocepurus bolivianus , mycocepurus borinquenensis , mycocepurus eucarnitae , mycocepurus manni , mycocepurus reconditus , mycocepurus tolteca , mycocepurus trinidadensis : kempf , 1963b pdf : 425 .
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the above specimen data are provided by antweb . please see mycocepurus smithii for further details
eucarnitae . mycocepurus smithii var . eucarnitae forel , 1913l : 235 ( w . ) cuba . junior synonym of smithii : kempf , 1963b : 425 .
comparative study of nest architecture and colony structure of the fungus - growing ants , mycocepurus goeldii and m . smithii
comparative study of nest architecture and colony structure of the fungus - growing ants , mycocepurus goeldii and m . smithii
thus mycocepurus smithii is only mostly asexual , rather than entirely so , and the transition among reproductive modes happens repeatedly .
comparative study of nest architecture and colony structure of the fungus - growing ants , mycocepurus goeldii and m . smithii .
comparative study of nest architecture and colony structure of the fungus - growing ants , mycocepurus goeldii and m . smithii
comparative study of nest architecture and colony structure of the fungus - growing ants , mycocepurus goeldii and m . smithii [ 2007 ]
distribution of sexual ( starred ) and asexual ( circled ) mycocepurus smithii . adapted from figure 1 of rabeling et al ( 2011 ) .
comparative study of nest architecture and colony structure of the fungus - growing ants , mycocepurus goeldii and m . smithii . - pubmed - ncbi
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fingerprinting
dna of the ant species - mycocepurus smithii - they found them all to be clones of the colony ' s queen .
dr himler ' s interest in mycocepurus smithii was originally sparked not by their unusually biased sex ratio , but by their ability to cultivate crops .
a brazilian population of the asexual fungus - growing ant mycocepurus smithii ( formicidae , myrmicinae , attini ) cultivates fungal symbionts with gongylidia - like structures .
bolivianus . mycocepurus bolivianus weber , 1938b : 155 , fig . 8 ( w . ) bolivia . junior synonym of smithii : kempf , 1963b : 425 .
borinquenensis . atta ( mycocepurus ) smithi var . borinquenensis wheeler , w . m . 1907c : 718 ( w . ) puerto rico . combination in mycocepurus : emery , 1924d : 335 . junior synonym of smithii : kempf , 1963b : 425 .
a brazilian population of the asexual fungus - growing ant mycocepurus smithii ( formicidae , myrmicinae , attini ) cultivates fungal symbionts with gong . . . - pubmed - ncbi
manni . mycocepurus manni weber , 1938b : 156 , figs . 1 , 2 ( q . ) bolivia . junior synonym of smithii : kempf , 1963b : 425 .
reconditus . mycocepurus reconditus borgmeier , 1937b : 246 , figs . 34 - 36 ( w . q . ) brazil . junior synonym of smithii : kempf , 1963b : 425 .
trinidadensis . mycocepurus smithi var . trinidadensis weber , 1937 : 378 , fig . 1 ( w . q . ) trinidad . junior synonym of smithii : kempf , 1963b : 425 .
cultivated fungi and gongylidia observed in gardens of mycocepurus smithii ( a , b , c , d ) and some species of higher attini ( e , f , g , h ) .
tolteca . atta ( mycocepurus ) smithi var . tolteca wheeler , w . m . 1907c : 718 ( w . ) mexico . junior synonym of smithii : kempf , 1963b : 425 .
to cite this page : schott , c . and c . strey 2011 .
mycocepurus smithii
( on - line ) , animal diversity web . accessed july 09 , 2018 at urltoken
to test the monophyly of m . smithii and reconstruct whether asexuality evolved once or multiple times from a sexually reproducing ancestor , we conducted a global phylogenetic analysis of the genus mycocepurus and a local analysis of only m . smithii taxa representing a sample from each of the genotyped populations ( table s5 ) .
comparison of gongylidia diameter from fungi cultivated by m . smithii and four species of higher attini .
fernandez - marin , h . , j . zimmerman , w . wcislo , s . rehner . 2005 . colony foundation , nest architecture and demography of a basal fungus - growing ant , mycocepurus smithii ( hymenoptera , formicidae ) .
recent work has shown that , in the amazonian fungus - growing ant mycocepurus smithii , queens use exclusively asexual reproduction and the male sex seems to have disappeared from the species . this finding illustrates the remarkable diversity of reproductive systems in ants .
kellner k , fernández - marín h , ishak hd , sen r , linksvayer ta , mueller ug ( 2013 ) data from : coevolutionary patterns and diversification of ant - fungus associations in the asexual fungus - farming ant mycocepurus smithii in panama .
rabeling , c . , verhaagh , m . & engels , w . 2007 . comparative study of nest architecture and colony structure of the fungus - growing ants , mycocepurus goeldii and m . smithii . journal of insect science 7 , article 40 .
( a ) nest chamber and pendant fungus garden of mycocepurus smithii ; ( b ) gongylidia organized in staphylae in the fungus garden of m . smithii ( 8x magnification ) ; ( c ) staphylae in a m . smithii cultivar . gongylidia in the fungus gardens of m . smithii ( d ) , t . fuscus ( e ) , ac . disciger ( f ) , a . laevigata ( g ) , and a . sexdens ( h ) . the scale bar in figure c represents 100 µm ; scale bars in figures d , e , f , g , and h represent 50 µm .
this week’s pnas contains detailed follow - up research on an intriguing story that emerged a couple years ago about a purportedly asexual , all - female ant species . a 2009 report looked for , and failed to find , any genetic signal for sexual reproduction in a population of the fungus - growing ant mycocepurus smithii in panama . this data , along with a lack of observed males over the extensive range of the species , prompted researchers to conclude mycocepurus smithii may be the first documented entirely non - sexual ant species .
rabeling , c . , j . lino - neto , s . cappellari , i . dos - santos , u . mueller , m . bacci jr . . 2009 . thelytokous parthenogenesis in the fungus - gardening ant mycocepurus smithii ( hymenoptera : formicidae ) .
at present , mycocepurus smithii seems to be the only representative of type a asexual ant species . natural history studies in puerto rico [ 25 ] and the brazilian amazon [ 26 ] , [ 27 ] suggested an unusual reproductive biology for m . smithii . fernández - marín et al . [ 25 ] first proposed the hypothesis that m . smithii might reproduce thelytokously . this study shows that m . smithii queens from the rio claro , brazil population are thelytokous . himler et al . [ 28 ] dissected six queens from a panamanian population , reaching the same conclusion . based on these three geographic data points , brazil , panama , both confirmed , and puerto rico , still unconfirmed , it seems that thelytoky is widespread across m . smithii ' s distribution range .
very interesting that asexuality would emerge many times from different populations along the amazon and rio negro , but only within a single species . what predisposes m . smithii to asexuality as opposed to other species of mycocepurus ? i can’t wait for this ant to be the next model organism…
citation : rabeling c , lino - neto j , cappellari sc , dos - santos ia , mueller ug , bacci m jr ( 2009 ) thelytokous parthenogenesis in the fungus - gardening ant mycocepurus smithii ( hymenoptera : formicidae ) . plos one 4 ( 8 ) : e6781 . urltoken
kellner k , fernández - marín h , ishak hd , sen r , linksvayer ta , mueller ug ( 2013 ) coevolutionary patterns and diversification of ant - fungus associations in the asexual fungus - farming ant mycocepurus smithii in panama . journal of evolutionary biology 26 ( 6 ) : 1353 - 1362 . urltoken
matching the ‘kerr males’ to m . obsoletus provides only circumstantial evidence for obligate thelytoky , but it does remove the strongest potential counter evidence , i . e . : that the ‘kerr males’ are actual males of m . smithii . while we cannot rule out the possibility that m . obsoletus and m . smithii males are morphologically identical ; this is most unlikely , however , given the observed morphological variation within and between mycocepurus species ( rabeling , pers . obs . ) .
attaxenus . trachymyrmex attaxenus menozzi , 1936b : 85 ( w . q . ) brazil . junior synonym of smithii : kempf , 1963b : 425 .
this is a word document which contains information about microsatellite genotypic data of the ant m . smithii in panama . see readme file for further explanation .
alternative to a nuptial flight , young queens could establish themselves in a new chamber that is part of the maternal colony . such colony expansion would be consistent with the observation that multiple reproductively active m . smithii queens inhabit the same colony [ 25 ] , [ 27 ] . however , the ability of independent nest foundation seems to persist in m . smithii , as infrequently alate m . smithii queens can be found in the leaf litter ( rabeling , pers . obs . ) and fernández - marín et al . [ 25 ] , [ 78 ] observed claustral nest founding of single dealate queens in puerto rican m . smithii .
mackay , w . , j . maes , p . fernandez , g . luna . 2004 . the ants of north and central america : the genus mycocepurus ( hymenoptera : formicidae ) . .
kellner et al . ( 2015 ) - mycocepurus smithii is an unusual attine ant since it is the only known asexually reproducing fungus - farming ant ( himler et al . 2009 ; rabeling et al . 2009 ) , such that each colony is comprised of a single ant clone tending a garden with a single fungus clone ( kellner et al . 2013 ) .
the existence of three unidentified mycocepurus males collected by w . e . kerr in rio claro , são paulo state in brazil [ 29 ] appear to challenge the hypothesis that m . smithii is obligately asexual . until recently , only two mycocepurus species were known from são paulo state : m . goeldii and m . smithii . the males of m . goeldii had been previously described [ 30 ] , and hence , by process of elimination , the males collected by kerr were tentatively assigned to m . smithii . interestingly , kempf [ 22 ] described the morphology of these males in his taxonomic revision of the genus , but had the foresight to not name them , because the specimens were unassociated with a maternal colony , making it impossible to identify them with confidence . the ‘kerr males’ therefore could belong to an undescribed species or to a described species not yet known from são paulo state .
in arthropods , the evolution of asexuality is often associated with hybridization ( 30 , 41 ) , a mechanism so far unknown in social hymenoptera ( 36 ) . given the monophyly of m . smithii and the phylogenetic congruence between nuclear and mitochondrial markers , hybridization is also unlikely to explain the origin of asexuality in m . smithii .
alternatively , microorganisms such as wolbachia , cardinium , and rickettsia have been shown to induce parthenogenesis in parasitoid wasps ( 42 – 44 ) . even though wolbachia infections have not been detected in social hymenoptera ( 45 ) , including m . smithii ( 16 ) , other parthenogenesis - inducing symbionts cannot be ruled out in m . smithii .
in the global analyses , the monophyly of the genus mycocepurus was unequivocally supported [ fig . s1 ; bayesian posterior probability ( bpp ) = 1 ; maximum likelihood bootstrap proportion ( mlbp ) = 100 ] , which is consistent with a previous analysis ( 35 ) . within the genus mycocepurus , nine reciprocally monophyletic , highly supported groups were recognized [ fig . s1 ; bpp = 1 , mlbp ≥ 92 ] , supporting the existence of five new species ( fig . s1 ) . the monophyly of m . smithii was well - supported [ fig . s1 ; bpp = 1 , mlbp = 92 ] , suggesting that extant m . smithii populations derive from a single most recent common ancestor ( mrca ) . an undescribed species from the colombian amazon was found to be the sister lineage of m . smithii , but with only weak statistical support ( fig . s1 ; bpp = 0 . 72 , mlbp = 56 ) .
to supplement the results of the m . smithii nest excavations and provide direct evidence for polydomy , different nest casting methods were tried , following methodologies described by williams and lofgren [ 31 ] and tschinkel [ 32 ] – [ 34 ] . however , the tunnel diameter of m . smithii nests was as narrow as 1 mm , which did not permit plaster of paris or molten zinc to trace the tunnels . other casting techniques will be required in the future to illustrate that nests of m . smithii are connected via tunnels .
behaviors specific to sexual reproduction might degenerate rapidly in parthenogenetic lineages [ 77 ] . in m . smithii , modifications of the nest related to nuptial flight activity were never observed during six field - seasons ( 2003–2008 ) , even though alate queens were present in m . smithii nests . at the same time sympatric sexual species showed modified nest structures and performed nuptial flights . the observation that m . smithii queens were still in the pupal stage whereas sexually reproducing species performed nuptial flights suggests that m . smithii queens may be produced over a wider time interval than in sympatric , sexual congeners . in the absence of males , it is unnecessary for m . smithii queens to depart in a synchronized population wide nuptial flight . consequently , it appears unnecessary to produce young queens in synchrony and to open the nest entrance for mating flights , which could possibly increase the risk of predation and parasitism .
wheeler , w . m . ; mann , w . m . 1914 . the ants of haiti . bull . am . mus . nat . hist . 33 : 1 - 61 ( page 42 , combination in mycocepurus )
several lines of direct evidence ( queen reproductive physiology ) and indirect evidence ( taxonomy & natural history ) implicate obligate thelytokous parthenogenesis for a brazilian population of the fungus - gardening ant species m . smithii . the presence of mature oocytes and yellow bodies indicate that dissected queens had oviposited and were reproductively active , whereas the empty spermathecae prove that queens were not inseminated . in combination , these characteristics indicate thelytokous parthenogenetic propagation and support fernández - marín’s [ 25 ] hypothesis of asexuality in m . smithii , formulated to explain the absence of males in a puerto rican m . smithii population .
for most mycocepurus species only the worker caste is described . to find males that match the ‘kerr males’ , special emphasis was given to collecting workers , queens , and males from any mycocepurus colonies . mycocepurus obsoletus is one of the species known only from workers and we collected entire colonies in a cerrado habitat at the margin of a gallery forest at the reserva ecológica do ibge , 35 km south of brasília , in the federal district of brazil ( 15 . 9436°s , 047 . 8767°w ; elevation 1087 m ) . to collect reproductives , the entrances of several nests were flagged during the dry season in july 2008 and on november 12 th 2008 , with the first rains of the season , alate mycocepurus males ( n = 19 ) and queens ( n = 13 ) were collected next to the previously marked nests . the external morphology of the males was compared to the ‘kerr males’ under a leica z6 apo dissecting scope .
in order to test if thelytoky is widespread in queens of m . smithii and to investigate the behavioral ecological consequences of thelytoky in field populations , we studied the population biology , reproductive physiology of queens and workers , and the mating behavior of a m . smithii population in brazil , and compared this population with two sexually reproducing , sympatric congeners . in addition , we provide taxonomic evidence for the identity of the ‘kerr males’ .
myrmecologist christian rabeling decided to probe the m . smithii problem in greater depth by sampling ants from a broader distribution . he found what researchers often discover when gathering additional data : the system is messier than thought .
this is an exciting find indeed . mycocepurus is one of those rare animals that apparently does both , rendering it a suitable model for studying why sex evolves . and being a fully social ant , it is primed to become a model species for studying sexual evolution in social contexts .
none of the 50 examined m . smithii workers was reproductively active . each worker had a pair of ovaries with a single ovariole per ovary . ovarioles did not contain mature oocytes and yellow bodies were absent . all workers were lacking a spermatheca .
fieldwork was conducted from september 29 th to october 26 th 2008 on the campus of são paulo state university ( unesp ) in rio claro , brazil ( 22 . 3955°s , 047 . 5424°w ; elevation 608 m ) . in the state of são paulo , the nuptial flights of most attine ant species , including those of mycocepurus [ 29 ] , occur during late september and october , which mark the beginning of the rainy season . in order to maximize the chance of finding males in the nests of m . smithii , we focused collecting on the probable mating season .
kempf , w . w . 1963b . a review of the ant genus mycocepurus forel , 1893 ( hymenoptera : formicidae ) . stud . entomol . 6 : 417 - 432 ( page 425 , queen described , senior synonym of attaxenus , bolivianus , borinquenensis , eucarnitae , manni , reconditus , tolteca , trinidadensis )
mackay , w . p . ; maes , j . - m . ; fernández , patricia rojas ; luna , g . 2004 . the ants of north and central america : the genus mycocepurus ( hymenoptera : formicidae ) . journal of insect science ( tucson ) 4 ( 27 ) : 1 - 7 pdf
the stem - group age ( i . e . , earliest possible origin ) of the fungus - gardening ants was estimated to be 52 million years ( ma ) [ confidence interval ( ci ) = 44 , 60 ] and the crown - group age was 50 ma ( ci = 43 , 58 ) , consistent with estimates in schultz and brady ( 35 ) . the estimated crown - group age of the genus mycocepurus is ∼10 ma ( ci = 6 , 14 ) , whereas the stem - group age is considerably older with 37 ma ( ci = 27 , 46 ) , which is also indicated by a long branch leading to the mrca shared with the sister lineage myrmicocrypta ( fig . s1 ) . the stem - group age of mycocepurus smithii is ∼1 . 65 ma ( ci = 0 . 57 , 2 . 84 ) , whereas the crown - group origin was estimated to be considerably more recent at 0 . 5 ma ago ( ci = 0 . 01 , 1 . 19 ) . this relatively recent estimate for the evolutionary origin of m . smithii is consistent with the almost complete absence of genetic variability observed in the nuclear dna sequences .
colonies of mycocepurus goeldii and an undescribed mycocepurus species served as a positive control for sexually reproducing species . nests of m . goeldii were shown to have a single nest entrance [ 27 ] . single , spatially separated colonies were chosen and a trench of 250 cm depth was dug in 50 cm distance from the first nest entrance , since pilot study showed that m . goeldii chambers could be encountered as deep as 190 cm below the surface ( rabeling , unpublished ) . vertical slivers of soil were sliced away with a spade and the position of all nest chambers were also recorded in a grid . the digging was continued 50 cm past the last excavated chamber .
this is one of the two species in which the workers lack the well - developed promesonotal spines in the center of the crown . generally there is a very low , sharp crest in this region . in most cases , it can be separated from the other species that lacks these spines , the brazilian mycocepurus obsoletus , on the basis of the distributions . otherwise , it can be separated by the lateral pronotal spines , which are long and sharp , not short and blunt as in m . obsoletus . specimens that have bumps in the center of the crown of spines , could be confused with mycocepurus tardus , but can be usually separated as m . tardus has definite promesonotal spines . the females have small lateral pronotal spines , and tiny inferior lateral pronotal spines . the females are slightly larger than those of mycocepurus curvispinosus ( ~ 3 mm total length ) . additionally the propodeal spines are bent slightly downward . these characters would easily separate it from females of m . curvispinosus . ( mackay et al . 2004 )
clade 1 of the leucocoprineae includes four primary clades of attine cultivars and closely related free - living fungi , here indicated as subclades a–d ( sensu mehdiabadi et al . ) . the clade within subclade b that contains the gongylidia - bearing fungal cultivar of m . smithii from rio claro is indicated by dashed lines and is enlarged in the inset . within the inset , the name of the gongylidia - bearing cultivar is indicated in red . the names of other cultivars of m . smithii in subclades a and b are indicated in red ; m . smithii additionally cultivates fungi in clade 2 ( not shown here ; see mehdiabadi et al . ) . the phylogram results from maximum - likelihood analyses of its sequence data of 305 fungal taxa . in the inset , numbers above branches are bayesian posterior probabilities ; numbers below branches are maximum - likelihood bootstrap proportions .
plot of 3d object coordinates resulting from an nmds analysis derived from individual genetic distances . colored circles represent genotypes of sexual m . smithii populations ( blue , belém ; orange , parintins ; green , caldeirão ; yellow , são gabriel da cachoeira ) and red triangles represent genotypes of asexual lineages .
under thelytoky and in the absence of males , the mating behavior is expected to degenerate , including the behavioral preparations of workers when modifying nest entrances for mating flights . preliminary behavioral observations in previous years showed that the sexually reproducing mycocepurus goeldii modifies its nest mounds in preparation of the nuptial flights of the alates ( see results for details ) . these modified nest entrances serve as a proxy if a particular colony is preparing for a nuptial flight . between september 29 th and october 15 th the nest entrances of ten m . goeldii colonies were monitored three times per day and compared to the ten entrances of m . smithii in the study plot .
rabeling confirmed with a combination of genetic and physiological data that indeed , most populations of m . smithii are asexual . but several sexual populations occur along the amazon river in northern brazil , and the genetic data suggest that each is more closely related to some of the asexual lineages than to each other .
the combined phylogenetic and population - genetic evidence is consistent with the hypothesis that sexual reproduction was lost in ancestors of parthenogenetic m . smithii populations . the spontaneous loss of sexual reproduction has been proposed for the little fire ant wasmannia auropunctata , in which sexual populations in the native range of this invasive species are likely the source of asexual invasive populations ( 36 ) . the proximate genetic mechanisms causing the loss of sexuality are not well - understood . however , studies of cape honey bees ( 37 ) and of parthenogenetic lineages of drosophila melanogaster ( 38 ) show that a single recessive allele can cause thelytoky . these examples suggest that the high propensity for switching from sexual to asexual reproduction in m . smithii may be controlled by a small number of genes . breeding experiments could test whether thelytoky is a qualitative or a quantitative trait in m . smithii by introgressing sexual genes into an asexual genetic background and observing the segregation pattern of the offspring .
eidmann [ 85 ] first hypothesized that m . smithii colonies were polygynous , because “in every of the numerous examined fungus chambers” he found “several old , dealate queens” ( p . 86 ) . unfortunately , eidmann never tested his hypothesis by dissecting the queens . fernández - marín et al . [ 25 ] further substantiated eidmann ' s hypothesis , suggesting that m . smithii colonies might be secondarily polygynous , because pleometrosis , the foundation of colonies by multiple queens , does occur in some colonies . dissections confirmed that multiple queens from the same nest were reproductively active , as indicated by the presence of developed ovaries and yellow bodies . however , it remains unclear if the dissected queens were found in the same fungus chamber or if they could have been reproductive females from different colonies . in addition , amazonian m . smithii colonies had several dealate queens in the same fungus chamber , but their reproductive status was not confirmed through dissections [ 27 ] .
our analysis of fragments of a single fungus garden shows that microbial communities within m . smithii gardens are not homogeneous . gardens of m . smithii have a hanging structure , with curtain - like strands hanging from the ceiling of the nest cave , which is unlike the relatively solid globule - shaped gardens typical of most leaf - cutting ants and other higher derived attines ( seal and tschinkel 2008 ) . within a single strand , the fungus grows from top to bottom ( the ants elongate the strand as it grows ) . because m . smithii in the field and in the lab start new gardens in the center of a chamber ceiling rather than near the ceiling periphery , it is likely that each garden has an age profile from the inside to the outside ( central older parts , peripheral younger parts ) . the nine garden fragments analyzed clustered into five likely groups , with the central ( presumably older ) parts forming one of these groups .
all mycocepurus specimens were stored in 95 % ethanol for further examination and censusing . voucher specimens of this study were deposited in the entomology collection of maurício bacci ' s molecular evolution laboratory at são paulo state university ( unesp ) , the museum of comparative zoology at harvard university ( mcz ) , the museu de zoologia da usp ( mzusp ) , and in c . rabeling ' s collection .
the wide geographic distribution could suggest an ancient origin of asexuality in m . smithii . low variability in dna sequence , however , does not support an ancient asexuality hypothesis , but rather argues for a recent origin of thelytoky followed by a rapid population expansion ( rabeling , unpublished ) . the widespread geographic distribution also suggests that geographic parthenogenesis , the occurrence of parthenogenetically reproducing lineages towards the margin of a distribution range [ 68 ] , [ 69 ] , unlikely applies to m . smithii , because both são paulo state and panama are located well within the continuous distribution range , which extends from northern mexico to argentina [ 22 , 23 ; rabeling , in prep ] .
geographic distribution of sexual ( stars ) and asexual ( circles ) m . smithii populations . localities refer to the sexual populations , distributed along the rio amazonas and the rio negro . asexual populations are widely distributed in latin america , ranging from northern mexico to northern argentina . lines of longitude and latitude are separated by units of 5° .
( a ) lateral view of ‘kerr male’ , ( b ) forewing and ( c ) hindwing in dorsal view . ( d ) male of mycocepurus obsoletus in lateral view , ( e ) forewing and ( f ) hindwing in dorsal view . morphological characters informative for species identification are the shape of the propodeal spine , of the petiole and the wing venation . scale bars represents 1 mm in all images .
wolbachia bacteria induce parthenogenetic reproduction in some parasitoid wasps [ 64 ] . several hymenopteran species have been tested for wolbachia infections , but wolbachia could never be demonstrated to induce parthenogenesis in any thelytokous eusocial hymenoptera species [ 28 ] , [ 41 ] , [ 52 ] , [ 65 ] . this supports the hypothesis that a sex determining system based on heterozygosity may form a limitation to wolbachia - induced parthenogenesis [ 65 ] . wolbachia endosymbionts induce the production of homozygous diploid offspring through gamete duplication [ 66 ] , [ 67 ] , which results in the exclusive production of diploid males . in ants and eusocial bees , females are required to be heterozygous at the sex - determining locus , a condition difficult to achieve when parasitized by a gamete duplicating endosymbiont [ 21 ] , [ 65 ] . because thelytoky seems to have evolved independently of wolbachia in ants and bees , it also seems unlikely that thelytoky of m . smithii queens was induced by wolbachia . the absence of wolbachia in m . smithii supports this expectation and additional antibiotic treatment could not ‘cure’ m . smithii from parthenogenesis [ 28 ] .
during this same time period , and until the end of the study three weeks later , the entrances of m . smithii nests remained unmodified , with some soil surrounding a single entrance hole . in addition , neither mating flights , nor modified nest mounds , nor males were ever observed throughout several years of field research in latin america ( rabeling , field observations 2003–2008 ) .
midpoint - rooted bayesian phylogram of m . smithii individuals representing each of the genotyped populations based on analyses of three mitochondrial gene fragments . bayesian posterior probabilities ( ×100 ) and ml bootstrap proportions are indicated as bpp / mlbp . red branches and bold font indicate taxa from sexually reproducing populations . all other taxa represent asexual populations . ( scale bar , number of substitutions per site . )
two nest architectures found in mycocepurus nests . ( a ) tree - like architecture of a m . goeldii nest ( cr 17 ) . eight of the eleven total fungus chambers are depicted in the cross section ; the fungus gardens are not shown . ( b ) necklace architecture of a m smithii nest ( cr 7 ) . the nest consisted of four chambers : the three lowermost ones were filled with a fungus garden , while the deepest chamber contained only a few mycelium strands , characteristic for recently built chambers . the uppermost chamber was abandoned , filled with organic matter from the decaying fungus garden and first rootlets started to grow into the cavity . because workers fill exiting tunnels of chambers with exhausted and / or parasitized fungus gardens with soil , the chamber was no longer connected to the main tunnel . both cross sections are drawn to the same scale . drawings by s . c . cappellari .
high relatedness is expected in colonies of m . smithii whether queens reproduce via apomictic or automictic thelytoky . kin conflict theory predicts conflict between individual kin or groups of kin ( i . e . , castes ) within a colony , because individuals or castes differ in relatedness , reproductive success or both [ 72 ] . in a society of highly related or even clonal organisms kin conflict over reproduction is expected to be reduced if not entirely absent . however , hamilton ' s rule [ 73 ] , [ 74 ] does not account for low frequency mutations with strong selective effects and therefore , relatedness alone is not always a good predictor of kin conflict over reproduction [ 75 ] , [ 76 ] . once the genetic underpinnings of reproduction in m . smithii societies are understood , this ant should be an interesting study organism for kin selection theory .
cyclical parthenogenesis , the alteration of asexual and sexual life stages ( 39 , 40 ) , is unlikely to occur in m . smithii . in each of the 218 parthenogenetic colonies collected in different seasons over an 8 - y period ( 2003–2010 ) , nestmates belonged only to one or very few clonal lineages . the nonrandom geographic distribution of sexual and asexual populations likewise suggests that the switch from sexuality to asexuality is unlikely triggered by season .
a - d : overview and details of a mycocepurus goeldii nest excavation . ( a ) mature fungus garden in a nest chamber . ( b ) adjacent chambers with fungus garden removed ; a connecting tunnel between them is marked with a stick . ( c ) excavation of two adjacent nests : on the far left side the excavation of a nest system is complete , on the right side it has just begun . ( d ) mycelial strands of an incipient garden hang from the ceiling of a recently constructed chamber . photos by c . rabeling .
firm evidence capable of identifying the cytological mechanism of thelytoky ( i . e . , apomixis versus automixis ) in m . smithii is currently lacking . a comprehensive genetic population study in combination with cytological evidence is necessary to distinguish between meiotic and mitotic parthenogenesis . himler et al . [ 28 ] tested queens and workers of 12 panamanian colonies for allelic variation at a single variable genetic locus and found queens and offspring to be genetically identical , concluding that m . smithii propagates clonally . however , during automixis with central fusion , two haploid nuclei are produced during the first meiotic division , which subsequently fuse to form the diploid zygote . gene recombination does not necessarily occur and thus , the genetic signature of apomicticly and automicticly produced offspring can be very similar if not identical [ 15 ] , [ 56 ] , [ 62 ] , [ 63 ] . a single genetic locus is hence insufficient to distinguish between mitotic and meiotic parthenogenesis .
additionally , these males were identified to species using molecular sequence information to ensure the males from brasília are in fact mycocepurus obsoletus males . the partial dna of three single - copy nuclear markers ( elongation factor 1 - alpha f1 copy , wingless and long wavelength rhodopsin ) and one mitochondrial gene ( cytochrome oxidase i ) were sequenced , resulting in 3512 base pairs , and compared to sequence information from m . obsoletus workers collected at the same site . the resulting molecular phylogeny will be presented in the context of an ongoing generic revision ( c . rabeling in prep . ) .
arrows drawn from the top left to the bottom right point to the spermatheca and arrows drawn from the top right to the bottom left point to yellow bodies . ( a , b ) the spermatheca of m . smithii is translucent ( empty ) , the ovaries are developed and yellow bodies are present . ( c , d ) a reproductively active m . goeldii queen has an opaque , sperm - filled spermatheca , developed ovaries and yellow bodies . the scale bar represents 2 mm in all images .
three males from rio claro in brazil [ 22 ] , [ 29 ] represented a challenge to the obligate asexuality hypothesis , because they were suspected to be males of m . smithii , indicating the potential for sex and recombination in this species . our morphological study , however , indicates that the ‘kerr males’ are the previously unknown males of m . obsoletus , not males of m . smithii . previously , m . obsoletus was not known to occur in rio claro , but we now know that m . obsoletus has a brazilian cerrado distribution ( fig . 4 ) , thus its occurrence in the rio claro area is certainly possible . in addition , m . obsoletus workers have been collected at two other relatively nearby localities in são paulo state ( fig . 4 ; see also [ 35 ] ) , further increasing the likelihood of its occurrence at or near rio claro . thus the ‘kerr males’ seem to represent only the southernmost presently known distribution record of m . obsoletus ( fig . 4 ) .
a total of 1 , 930 m . smithii individuals from 234 colonies collected at 39 different localities in latin america ( fig . 1 and table s1 ) was genotyped at 12 variable microsatellite loci yielding 106 alleles ( range : 2–15 alleles per locus ) . the number of alleles per locus per individual never exceeded two , indicating diploidy of females . of the genotyped populations , 89 . 7 % ( n = 35 ) showed population - genetic signatures of clonality , whereas 10 . 3 % ( n = 4 ) showed an increase of unique multilocus genotypes , indicative of genetic recombination caused by sexual reproduction .
in the rio claro population , only a single queen was found per fungus chamber ( n = 10 ) . the presence of several queenright chambers in one fungus chamber cluster below a single entrance suggests that one polydomous colony can contain multiple reproductively active queens , supporting functional polygyny . other clusters of fungus chamber , however , inhabited only a single reproductively active queen . in case these clusters are not part of the same polydomous nest , the rio claro m . smithii population would comprise a mosaic of monogynous and polygynous colonies , which would be consistent with fernández - marín et al . ' s [ 25 ] hypothesis of secondary polygyny . again , a genetic population study will have to test if queens and worker offspring share the same genetic signature or if multiple mothers produced the workers of a single colony .
among all m . smithii populations , 66 asexual genotypes were identified , 57 of them representing unique multilocus genotypes ( r = 0 . 86 ; tables s2 and s3 ) . five repeated multilocus genotypes were shared between 10 geographically proximate populations ( ∼10–40 km distance ) , and three unique genotypes were identified in seven geographically distant populations ( ∼700–2 , 600 km distance ; tables s2 and s3 ) . calculating the probability that repeated multilocus genotypes from different populations originated from distinct sexual events revealed that identical multilocus genotypes belong to the same clonal lineage ( p sex < 0 . 01 ) , indicating long - distance dispersal events of individuals from the same asexual lineage . no genetic variation was present within repeated multilocus genotypes ( f is = −1 ) , but significant genetic variance was structured among them [ analysis of molecular variance ( amova ) ; f st = 0 . 624 , p = 0 . 01 ] .
within eusocial hymenoptera , arrhenotokous parthenogenesis is a widely distributed phenomenon [ 15 ] , [ 36 ] and the reproductive modes in ants are particularly diverse [ 37 ] . in contrast , thelytokous parthenogenesis , the production of female offspring from unfertilized eggs , has been rarely confirmed in eusocial hymenoptera , although the number of recognized thelytokous species is increasing . so far , thelytoky has been demonstrated convincingly for the south african honey bee subspecies apis mellifera capensis [ 19 ] – [ 21 ] and seven ant species , which can be divided into three categories of asexual reproduction : queens produce queens and workers via thelytoky , but workers are sterile and males seem to be absent ( called here : type a thelytoky ) ; workers produce workers and potentially queens via thelytoky ( type b ) ; or queens produce queens thelytokously and workers sexually , and workers produce queens thelytokously ( type c ) . males are known for all type b and type c asexual ant lineages , suggesting the potential for sex and recombination , but males remain unknown for type a asexuals , of which m . smithii is the first known example in the formicidae .
a total of ten m . smithii nest entrances were identified in the 7 m 2 study plot ( 1 . 43 entrances / m 2 ) ( fig . 1 ) . the excavation of this population revealed a total of 59 fungus chambers ( see table s1 for exact chamber contents ) . the entire population consisted of 851 workers and 12 queens , averaging approximately 14 workers per chamber and one queen in every 5 th fungus chamber ( see table s1 ) . the maximum number of workers per chamber was 43 , the minimum one . all queenright chambers contained only a single reproductively active queen , however , the number of queens encountered was higher than the number of nest entrances in the study area . the brood was distributed in 21 ( 36 % ) chambers and consisted of 115 early instar larvae , 40 worker pupae , 51 queen larvae and 274 queen pupae . the sexual brood was restricted to ten chambers ( 17 % ) and in eight of them worker brood and sexual brood co - occurred . the maximum and minimum number of sexual brood per chamber was 80 and one , respectively . no male pupae and no alate queens and males were present in the nest chambers .
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an amazonian ant has dispensed with sex and developed into an all - female species , researchers have found .
the ants reproduce via cloning - the queen ants copy themselves to produce genetically identical daughters .
this species - the first ever to be shown to reproduce entirely without sex - cultivates a garden of fungus , which also reproduces asexually .
the finding of the ants '
world without sex
is published in the journal proceedings of the royal society b .
anna himler , the biologist from the university of arizona who led the research , told bbc news that the team used a battery of tests to verify their findings .
and when they dissected the female insects , they found them to be physically incapable of mating , as an essential part of their reproductive system known as the
mussel organ
had degenerated .
asexual reproduction of males from unfertilised eggs is a normal part of some insect reproduction , but asexual reproduction of females is
exceedingly rare in ants
, wrote the researchers .
in social insects , there are a number of different types of reproduction ,
explained dr himler .
but this species has evolved its own unusual mode .
she and her colleagues do not know exactly why this particular species has become fully asexual , and how long ago the phenomenon evolved .
they are carrying out further genetic experiments , which will enable them to estimate how long ago the evolutionary change occurred .
it avoids the energetic cost of producing males , and doubles the number of reproductive females produced each generation from 50 % to 100 % of the offspring .
if we ' re more diverse , we ' re more resistant to parasites and disease ,
explained laurent keller , an expert in social insects from the university of lausanne .
in a colony of clones , if one ant is susceptible to a parasite , they will all be susceptible . so if you ' re asexual , you normally don ' t last very long .
but in ants we ' re seeing more and more reports of unusual methods of reproduction ,
added professor keller , who was not involved in this study .
he also points out that social insects , like ants , may be particularly well suited to this type of reproduction because it enables the queen to control the caste and sex of all the offspring in her colony .
ants discovered farming long before we did - they have been cultivating fungus gardens for an estimated 80 million years .
they collect plant material , insect faeces and even dead insects from the forest floor and feed it to their crops ,
she said .
many different species of ant - including the famous leafcutter ants - cultivate fungi , relying on it for nutrition .
but this particular species is able to grow
a greater number of crops than other ant species
, she explained .
when we started to study this species more closely , we just weren ' t finding any males . that ' s when we started to look at them in a different way .
since the fungus crop reproduces asexually , dr himler thinks it might give the ants some kind of advantage
not to operate under the usual constraints of sexual reproduction
.
there is certainly more work to be done in this system ,
she added .
we ' re quite excited about the direction this research might take us , and its implications .
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a total of 1 , 647 individuals from 218 colonies in 35 populations exhibited genetic signatures of clonal reproduction . asexual reproduction was characterized by sharing of repeated multilocus genotypes among individuals ( table s1 ) , maximum deviation from random mating ( f is = −1 ; table s2 ) , and a low genotype - to - individual ratio ( i . e . , g : n approaching 0 , whereas a g : n of 1 indicates that each individual is genetically distinct from another ) ( table s1 ) . to determine the number of independently evolved asexual lineages that arose at different localities from the sexual population , we estimated the probability that slightly different multilocus genotypes originated from separate sexual events ( p sex > 0 . 01 ) instead of arising from accumulated mutations or scoring errors ( p sex < 0 . 01 ) . in addition , clonal diversity ( r ) was calculated .
a comparison of the 57 unique multilocus genotypes revealed high frequencies of low genetic distances between genotypes , resulting in a bimodal frequency distribution of genetic distances and indicating the potential existence of mutations or scoring errors in clones ( 34 ) . eleven multilocus genotype pairs differed from one other genotype only by a single allele , reducing the number of asexual lineages that potentially originated from distinct sexual events to 46 ( p sex < 0 . 01 , r = 0 . 69 ) . further lowering the threshold and allowing two to six alleles to be shared among multilocus genotypes within an independently evolved clonal lineage , we identified 43 ( r = 0 . 65 ) to minimally 38 ( r = 0 . 57 ) independently evolved clonal lineages .
in 20 clonal populations , only a single multilocus genotype was encountered across different colonies . in 15 populations , two to maximally six multilocus genotypes coexisted at a single site ( table s1 ) . in five populations , all or a subset of multilocus genotypes differed by one to six alleles , suggesting a single colonization event followed by diversification within clonal lineages due to the accumulation of mutations or scoring errors ( table s1 ) . in contrast , 12 populations harbored multilocus genotypes differing by 7–15 alleles , indicating independent colonization events of these sites by distantly related clonal foundress queens . the highest diversity of clonal lineages ( n = 5 ) was discovered at a peruvian lowland rainforest site ( los amigos ) .
four amazonian populations , distributed along the rio amazonas and the rio negro ( fig . 1 ) , exhibited population - genetic signatures of genetic recombination , indicative of sexual reproduction ( tables s1 and s2 ) . among 283 genotyped individuals , 210 multilocus genotypes were identified , resulting in high genotype - to - individual ( g : n ) ratios , ranging from 0 . 71 to 1 ( table s1 ) . recombinant populations were characterized by inbreeding indices diverging from genetic fixation ( f is = −1 ) , ranging from 0 . 03 to −0 . 77 , and observed and expected heterozygosities were similar , as expected for populations under hardy–weinberg conditions ( table s2 ) .
the genetically recombinant population from são gabriel da cachoeira showed that all nestmates ( n = 8 ) were genetically distinct ( g : n = 1 , h o = 0 . 365 , h e = 0 . 315 , f is = −0 . 172 ) , consistent with strict sexual reproduction ( tables s1 and s2 ) . however , in the belém colony ( g : n = 0 . 96 , h o = 0 . 451 , h e = 0 . 466 , f is = 0 . 034 ) and the parintins colony ( g : n = 0 . 71 , h o = 0 . 650 , h e = 0 . 398 , f is = −0 . 773 ) , few individuals shared a multilocus genotype , suggesting mixed sexual and parthenogenetic reproduction in these populations .
as test statistics for asexuality , we used the existence of repeated multilocus genotypes and maximum deviation from random mating ( f is ) ( 53 – 55 ) . the genotype - to - individual ratio ( g : n ratio ) was applied to identify multilocus genotypes ( 55 ) ( table s1 ) . independently evolved asexual lineages ( clones ) originating from separate sexual events were distinguished from slightly different multilocus genotypes that diversified through accumulation of mutations or scoring errors by calculating the probability , p sex , following the methodology outlined in ref . 34 and implemented in genclone 2 . 0 ( 56 ) . the observed and expected heterozygosity for each clonal lineage ( 57 ) , the proportion of clonal genotypes in a population , f statistics , and amova were calculated in genalex 6 ( 58 ) and genetic data analysis ( 59 ) . to reveal the underlying population - genetic structure of sexual and asexual populations , we used the multivariate statistical methods ( 60 – 62 ) nmds , principal component analysis , and dapc , as implemented in permap ( 63 ) , genalex ( 58 ) , and systat ( systat software ) .
we used a bayesian relaxed clock uncorrelated lognormal approach implemented in the program beast v1 . 4 . 8 with a yule process as the tree prior ( 73 – 75 ) . the root node was given a normal age prior distribution ( mean = 73 . 5 , sd = 4 . 5 ) , following methodology described in ref . 76 . based on fossil data , lognormal age prior distributions were assigned to three internal nodes , as outlined in ref . 35 . for more details on analyses and results , see si materials and methods and tables s1 – s8 .


the garden rose tortricid moth or fruit tortricid ( acleris variegana ) is a moth of the family tortricidae .

it has a palaearctic distribution .

the moth flies from july to september mainly at night and is attracted to bright lights .

the larvae feed on various trees and shrubs including rose and apple .

acleris variegana

acleris variegana may appear similar to other acleris species due to its highly variable forewing pattern .
agroatlas - pests - acleris variegana ( denis & shiffermuller ) - garden rose tortrix .
agroatlas - pests - area of distribution and damage of acleris variegana den . et schiff .
acleris variegana ( garden rose tortrix ) - norfolk micro moths - the micro moths of norfolk .
peronea variegana ( den . et schiff . ) , acalla variegana ( den . et schiff . ) .
kari pihlaviita added the finnish common name
pihatalvikääriäinen
to
acleris variegana denis & schiffermüller 1775
.
hans - martin braun added the german common name
rosenwickler
to
acleris variegana denis & schiffermüller 1775
.
hans - martin braun added the english common name
fruit tortricid moth
to
acleris variegana denis & schiffermüller 1775
.
hans - martin braun added the german common name
heidelbeer - wickler
to
acleris variegana denis & schiffermüller 1775
.
hans - martin braun added the english common name
garden rose tortrix
to
acleris variegana denis & schiffermüller 1775
.
acleris variegana may appear similar to certain forms of other acleris species , including a . keiferi , a . maculidorsana , and a . robinsoniana in north america and a . permutana in europe . a genitalic dissection can be used to confirm identity .
acleris variegana completes two generations per year in north america . adults fly from april to july and again from august to november . overwintering occurs as a second generation adult .
class insecta , order lepidoptera , family tortricidae , subfamily tortricinae , tribe tortricini , genus acleris .
acleris variegana is widely distributed throughout europe , northwestern africa , central asia , and china . it was first reported from north america in the early 1900 ' s where it is present on the east and west coasts .
obraztsov , n . s . 1963 . some north american moths of the genus acleris . proceedings of the united states national museum . 114 : 213 - 270 .
pustovarov v . v . 1975 . tortricidae ( lepidoptera ) of the genus acleris hbn . in forests of se armenia . biologicheskii zhurnal armenii ( yerevan ) , 28 ( 4 ) : 91 - 93 . ( in russian ) .
the forewing pattern of a . variegana is variable ; bradley et al . ( 1973 ) lists eight forms as occuring in the united kingdom . powell ( 1964 ) describes three distinct phenotypes found in western north america : 1 ) forewing white on inner half with a dark dorsal triangle , outer half dark - purplish ; 2 ) forewing divided into white basal and blue - black or purplish distal area ; 3 ) and forewing ground color tan with a basal dorsal triangle and purplish outer costal triangle . males lack a forewing costal fold .
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western europe , north america , northwest africa , asia minor , syria , iran , afghanistan and china . in the former ussr the species is distributed in the european part ( except in the far north ) , transcaucasia , volga river basin , ural , kazakhstan , central asia and southwest siberia .
the species is a significant pest of rosaceous fruit plants , especially of apple , pear , plum , sloe , apricot . caterpillars also cause damage to quince , cherry , hawthorn , almonds , cotoneaster , bird cherry . they have been documented on hazel , oak , elm , willow , and berry bushes of the genus vaccinium . mainly the first generation caterpillars cause harm .
agronomic measures include pruning old and diseased branches . biological measures include application of biological preparations . chemical measures include local insecticide treatments during development of first generation caterpillars before flowering of pome trees ( in red bud phase ) immediately after flowering of stone cultures in spring , and during development of 2nd generation caterpillars in mid - july . monitoring and forecast are possible with the use of pheromone traps .
galetenko s . m . 1964 . to morphology of fruit leafrollers . in : kochkin m . a . , rubtsov n . i . , ryndin n . v . , eds . 150 years to state nikitskii botanical garden . collection of scientific works . v . 37 . moscow : kolos . 531 - 395 p . ( in russian ) .
kostyuk yu . a . 1974 . family tortricidae . in : vasil . ev , v . p . , ed . pests of agricultural crops and forest plantations . v . 2 . arthropods . kiev : urozhai . 261 - 320 p . ( in russian ) .
kostyuk yu . a . 1980 . tortricidae . issue 10 . tortricinae . in : dolin v . g . , ed . fauna of ukraine . v . 15 . kiev : naukova dumka . 424 p . ( in ukrainian ) .
kuznetsov v . i . 1994 . family tortricidae . in : kuznetsov v . i . , ed . insects and mites . pests of agricultural plants . v . 3 ( 1 ) . lepidoptera . st . petersburg : nauka . 51 - 234 p . ( in russian ) .
vasil . ev v . p . , livshits i . z . 1984 . pests of fruit crops . moscow : kolos . 399 p . ( in russian ) .
© 2003 - 2009 project «interactive agricultural ecological atlas of russia and neighboring countries . economic plants and their diseases , pests and weeds»
) bushes , as the name suggests , but also on a range of other trees and shrubs .
flying from july to september , it is mainly nocturnal and is attracted by bright lights .
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powell , j . a . & p . a . opler , moths of western north america , pl . 17 . 23m , . 24m , . 25m ; p . 144 . book review and ordering
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the nominate form has a forewing demarcated centrally , with a white basal half showing a large dark patch on the dorsum and a ferruginous / black distal half .
recorded in 60 ( 87 % ) of 69 10k squares . first recorded in 1914 . last recorded in 2018 .
underlying maps using digital map data © norfolk online lepidoptera archive - nola™ 2018 . © james wheeler - n o r f o l k m o t h s 2007 - 2018 . data © nola™ 2018
synonyms : albana ( peronea ) , alpicolana ( form ) , argentana ( form ) , asperana ( pyralis ) , blandiana ( tortrix ) , brunneana ( form ) , caeruleoatrana ( ab . ) , cirrana ( peronea ) , costimaculana ( peronea ) , fuscana ( form ) , insignata ( tortrix ) , nyctemerana ( tortrix ) , osbeckiana ( tortrix ) , uniformis ( ab . )
larvae are yellowish or light green . the head is brown and the prothoracic shield is concolorous with the abdomen . legs are brown or pale . most individuals exhibit a black spot located laterally on the head and a dark posterolateral mark on each side of the prothoracic shield .
eggs are laid singly or in small batches on leaves of the host plant . early instar larvae tie together two leaves and skeletonize them from the inside ; later instars consume the entire leaf . pupation occurs in the larval shelter or among dead leaves on the ground .
larvae have been recorded feeding on a variety of host plants , primarily those in the family rosaceae .
dasiphora fruticosa ( l . ) rydb . ssp . floribunda ( pursh ) kartesz
bradley , j . d . , w . g . tremewan and a . smith . 1973 . british tortricoid moths - cochylidae and tortricidae : tortricinae . the ray society , london , england .
powell , j . a . 1964 . biological and taxonomic studies on tortricine moths , with reference to the species in california . university of california publications in entomology . vol . 32 . 317 pp .
powell , j . a . and p . a . opler . 2009 . moths of western north america . university of california press , berkeley . 369 pp .
razowski , j . 2002 . tortricidae of europe , vol . 1 , tortricinae and chlidanotinae . frantisek slamka , slovakia . 247 pp .
tortricids of agricultural importance by todd m . gilligan and marc e . epstein interactive keys developed in lucid 3 . 5 . last updated august 2014 .
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forewing pattern is variable . powell ( 1964 ) lists three distinct phenotypes found in california : 1 ) forewing white on inner half with a dark dorsal triangle , outer half dark purplish ; 2 ) forewing divided into white basal and blue - black or purplish distal area ; 3 ) forewing ground color tan with a basal dorsal triangle and purplish outer costal triangle . males lack a forewing costal fold .
this species completes two generations per year in california . adults are present from mid - june through late july and again from mid - september through november .
eggs are laid singly or in pairs . early instar larvae tie together two leaves and skeletonize them from the inside ; later instars consume the entire leaf . pupation occurs in the larval shelter . both adults and larvae overwinter .
larvae feed on members of the rosaceae in california , including : apple ( malus sp . ) , blackthorn ( prunus spinosa ) , cotoneaster ( cotoneaster sp . ) , firethorn ( pyracantha sp . ) , hawthorn ( crataegus sp . ) , pear ( pyrus communis ) ; rose ( rosa sp . ) , and sweet cherry ( prunus avium ) .
palearctic . england and northwest africa through europe , central asia , and china . first reported from north america in the early 1920 ' s .
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14 - 19 mm wingspan , forewings whitish ochreous to purplish , variable suffused with grey , frequently with much of the basal half white and with distinct , often black , scale tufts ; hindwings grey . a very variable species .
0 . 6 x 0 . 5 mm ; pale yellowish to reddish ; deposited singly or in small batches on the leaves of the food plant , usually along the mid - rib on either surface .
12 - 14 mm long ; head and prothoracic plate yellowish or greenish brown , head with blackish brown marking near stemmata , and postero - lateral markings ; abdomen yellowish green or light green , pinacula concolorous or paler ; anal plate green ; thoracic legs yellowish brown .
14 - 19 mm wingspan . head brownish grey to dark grey , labial palpus brownish grey , sometimes suffused with ochreous , thorax and abdomen brownish grey . forewing hardly expanding distally , costa bent outwards in basal third , apex short , pointed .
the variation in forewing colouration and markings is considerable . many ' forms ' are recognized . these forms are mostly distinct and do not intergrade .
tegumen with large terminal lobes ; socii broad , rounded laterally , almost straight at opposite side ; tuba analis broad . sacculus broad anteriorly , rounded basally , concave before middle , arced outwards in posterior part of ventral edge , delicately undulate or dentate before long spined termination . aedeagus rather broad with 8 or 9 fairly short cornuti and strong thorn .
sterigma with elongate , rounded apically , anterior projections ; antrum heavily sclerotized , large ; ductus bursae short , bulbous posteriorly ; ductus seminalis situated submedially ; signum somewhat elongate , dentate .
is a very variable species . the forewing can be sharply demarcated at the centre , the whole of the basal half being white except for a fuscous suffusion at the base of the costal margin and an indication of the sub - basal fascia on the dorsum , usually in the form of two short vertical black bars , the large scale - tuft below the submedian fold being white , while in contrast the distal half of the wing is blackish fuscous intermixed with ferruginous and shining plumbeous scaling . the basal half of the forewing can also be distinctly tinged with pale ochreous .
the forewing ground colour can be light ochreous , the sub - basal fascia developed on the dorsum , and the median fascia obsolescent dorsally but well developed on the costa and confluent with the pre - apical spot , forming a dark fuscous or blackish fuscous triangular costal blotch .
the markings can also be obsolescent except for the fuscous costal blotch , and the white ground colour sprinkled with blackish scales , especially in the distal area , or the forewing colour can be almost unicolorous white with only sparse fuscous scaling in the apical area , especially along the costa and termen . a melanistic form has the forewing blackish plumbeous with somewhat darker irregular transverse marbling ( after bradley et al . , 1973 ) .
adults appear from july to september . eggs are then laid in batches on either side of the leaves , usually along the mid - vein . they hatch in the spring . larvae feed from may to late june or early july , sheltered within loosely spun leaves or in folded leaf edges , eventually pupating in the larval habitation or amongst fallen leaves ( alford , 1995 ) .
europe to asia minor , central asia and china ; north - western africa ; north america .
( grav . ) ( ichneumonidae ) ( record from canada ; loan and doganlar , 1980 ) .
in general appearance but it may be distinguished by the erect scale - tuft in the submedian fold . also compare genitalia [
occurs from western , central and southern europe to asia minor and central asia . host plants are
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a scarce and local species , occurring in the southern parts of england and wales , and preferring coastal sandhills and limestone habitats .
the moths fly in august and september and are quite elusive by day , but are attracted to light after dark .
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the distribution in this summary table is based on all the information available . when several references are cited , they may give conflicting information on the status . further details may be available for individual references in the distribution table details section which can be selected by going to generate report .
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map was created based on information taken from open published literature and maps of scale 1 : 33 , 000 , 000 .
vector map . area of species distribution is shown by polygons . zones of recorded mass outbreak are shown by polygons .
babenko z . s . 1982 . phytophagous insects of fruit and berry plants in forest zone of ob river region . tomsk : tomsk university . 268 p . ( in russian ) .
barysheva yu . a . , bruk s . i . , dmitriev a . v . , stroev k . v . , sutormin a . i . , shashko d . i . , yagust s . i . 1992 . zones of soil - vegetation types . geographical atlas of the ussr . moscow : com . geodesy and cartography of the ussr . 48 p . ( in russian ) .
d . yakonov a . m . , kozhanchikov i . v . 1949 . lepidoptera . in : pavlovskii e . n . , shtakelberg a . a . , eds . pest animals of middle asia ( handbook ) . moscow - leningrad : an sssr . 179 - 198 p . ( in russian ) .
danilevskii a . s . , kuznetsov v . i . , fal . kovich m . i . 1962 . tortricidae ( lepidoptera ) in montane areas of kazakhstan . in : marikovskii p . i . , ed . materials on study of insects of kazakhstan . alma - ata : an kaz . ssr . 69 - 117 p . ( in russian ) .
dolzhenko v . i . , ed . 2004 . methodical instructions on registration tests of insecticides , acaricides , molluscicides , and rodenticides in agriculture . st . petersburg : vizr . 363 p . ( in russian ) .
dubatolov v . v . 2000 . tortricidae - collection of siberian zoological museum . siberian zoological museum of the institute of animal systematics and ecology , siberian branch of russian academy of sciences . novosibirsk :
fal . kovich m . i . 1962 . tortricidae ( lepidoptera ) of leningrad region . in : stackelberg a . a . , ed . fauna of leningrad region and karelia . materials on invertebrate fauna . moscow - leningrad : an sssr . 48 - 80 p . ( in russian ) .
galetenko s . m . 1960 . tortricid pests of orchards in crimea . in : livshits i . z . , ed . pests and diseases of fruit and decorative plants . proc . state bot . garden . v . 33 . yalta : nikitskii state bot . garden . 157 - 192 p . ( in russian ) .
koroljeva ie , vilchevskaya ev , ruhovich di . 2003 . digital arable land map . laboratory of soil information of the dokuchaev soil institute , moscow , russia [ based on : yanvareva lf . ( ed . ) , martynjuk kn . , kisileva nm . 1989 . map of land use , faculty of geography , moscow state university , moscow , russia . ] .
kostyuk yu . a . 1974 . family tortricidae . in : vasil . ev v . p . , ed . pests of agricultural crops and forest plantations . v . 2 . arthropods . kiev : urozhai . 261 - 320 p . ( in russian ) .
kostyuk yu . a . 1980 . tortricidae . issue 10 . tortricinae . in : dolin v . g . , ed . fauna of ukraine . v . 15 . kiev : naukova dumka . 424 p . ( in ukrainian )
krasil . nikova g . a . 1967 . review of lepidoptera damaging fruiters in sw turkmenistan . in : mizgireva o . f . , ed . proc . turkmen . res . station vir . issue 4 . ashkhabad : vir . 223 - 273 p . ( in russian ) .
kullberg j . , albrecht a . , kaila l . , varis v . 2001 . checklist of finnish lepidoptera - suomen perhosten luettelo . sahlbergia ( helsinki ) , 6 ( 2 ) : 1 - 190 .
romanova v . p . 1949 . leafrollers and their parasites on forest trees in environs of rostov - na - donu . proc . biol . inst . v . 15 ( 6 ) . rostov - na - donu : rostov university . 47 - 61 p . ( in russian ) .
sedykh k . f . 1974 . animal world of komi assr . invertebrates . syktyvkar : komi publishing house . 189 p . ( in russian ) .
sherniyazova r . m . 1974 . tortricidae , lepidoptera of trees and bushes in varzob gorge ( tajikistan ) . izvestiya an tadzhik ssr . biol . n . ( dushanbe ) , 54 ( 1 ) : 48 - 53 . ( in russian ) .
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) . lbam is currently found in 15 coastal and near coastal counties in california suggesting it has been present in the state for several years . to date the pest has not reached outbreak status at any location in california .
the united states department of agriculture ( usda ) posits that the potential distribution of lbam includes all areas having sufficient thermal units for the completion of three generations ( i . e . , california and the southern half of the united states ) with estimated losses of $ 118 million per annum from apple , grape , orange and pear production ( fowler et al .
) . in response to this perceived threat , the usda considers lbam to be an actionable quarantine pest , and has placed quarantine on california produce in affected counties as well as all counties in hawaii ( federal quarantine order of may 2 , 2007 ) . in late 2007 , the california department of agriculture ( cdfa ) began an eradication program using pheromones , sprays and other techniques . the pheromone spray program engendered considerable public protest concerning claimed public health effects , the lack of hard evidence that lbam is a serious pest and the proposed expenditure of nearly 100 million dollars for the program in california alone . because of public outcry , an eradication program based on the sterile insect technology ( sit ) has now been proposed .
the potential ecological and economic impact of invasive species can be influenced by many factors ( parker et al .
) , but for invasive arthropod pests , climate and temperature in particular have played a central role in the assessment of risk ( e . g . , fowler et al .
) . a variety of climate matching models have been used to predict the potential distribution of invaders ( coetzee et al .
) , but there have been few attempts to include biotic interactions in addition to abiotic drivers into such models ( e . g . , gutierrez and baumgärtner
) . here , we analyze the potential distribution of lbam in arizona and california using a temperature - driven demographic model to provide a more quantitative and realistic approach for assessment of the risk of this invasive pest .
) described the biology of lbam , and this body of work is the basis for our analysis . the developmental biology of
light brown apple moth does not have a diapause period and feeds on numerous plant species across family lines ; the lower threshold for development of all stages was estimated as 7 . 5°c ( danthanarayana
) , but reanalysis of the data for eggs and larvae yields a non linear relationship with a slightly lower threshold of 6 . 8°c ( fig .
) . the mean duration of the egg stage is 5 . 7 days at 28°c ( 120 . 8 degree days ;
) with no eggs hatching above 31 . 3°c . larval and pupal developmental times are 335 and 142
, respectively with an upper threshold for development for larvae and pupae of approximately 31 . 5°c . the generation time is 594
; mean fecundity is approximately 384 but varies considerably ( 0–1492 ) with female size , larval food and temperature ( danthanarayana et al .
) , but only temperature is used in the model . the optimum temperature for oviposition is near 20°c . at constant temperatures of 20 and 25°c , > 50 and 80 % of the eggs are laid by the fourth and the seventh day after emergence respectively with peak oviposition occurring when adult females are 2–3 days old . body weight of both male and female moths at emergence increases with increasing temperature up to 25°c . danthanarayana (
) provides information on the diel and lunar periodicities , seasonal variation in flight activity and the influence of food and temperature on flight performance in the field . flight duration increases with temperature in the range 15–28°c , with males being stronger fliers than females with the longest flights occurring on day four . mating in females begins within 24 h of emergence with nearly all being mated within 4 days . the average number of matings is 2 , although by day 11 some females may mate as many as five times ( gu and danthanarayana
) found that a small proportion of lbam females produce predominantly female progeny resulting in the proportion of males in field samples to range from 0 . 43 to 0 . 45 . based on lbam’s biology , theoretical considerations suggest that this value is near the theoretical optimum assuring that nearly all females are mated ( see geier et al .
) . the resulting sex ratio in our model is 1 . 1 ♀ : 0 . 9 ♂ .
) were used to estimate the effects of adult age and temperature on per capita fecundity ( fig .
a–c ) . the data are illustrated in days for ease of interpretation , but all calculations in the model are made in physiological time units . the effect of temperature (
three sources of lbam mortality are included in the model : ( 1 ) intrinsic mortality in the favorable range between 6 . 8 and 28°c is accounted for by the differing progression of the population members with temperature through the distributed maturation time model , ( 2 ) mortality due to extremes of temperatures , and ( 3 ) mortality due to the combined action of natural enemies and other factors . only mortality sources ( 2 ) and ( 3 ) require further explanation .
data on the lethal effects of low temperature on lbam are sparse , and hence we used estimates from an ongoing study by buergi and mills ( unpublished observations ) showing that 30 % of larvae die within 1 day at −10°c and all larvae died within 1 day at −16°c . we computed the daily mortality rate from low temperatures as the integral of time and temperature below the developmental threshold (
) , and used it to model the daily over - wintering mortality rate .
$ $ \ begin { gathered } { \ text { if } } \ , t { } _ { \ min } < 6 . 8 ^ \ circ c , \ ; 0 . 01 < \ mu _ { t } = 1 - 0 . 015 \ , \ exp ( 0 . 323dd _ { { < 6 . 8 ^ \ circ { \ text { c } } } } ) < 1 \ quad { \ text { i } } \ hfill \ \ { \ text { if } } \ , t { } _ { \ max } > 28 ^ \ circ c , \ ; 0 . 01 < \ mu _ { t } = 1 - 0 . 125 \ , \ exp ( 0 . 025dd _ { { > 28 ^ \ circ { \ text { c } } } } ) < 1 \ quad { \ text { ii } } \ hfill \ \ { \ text { else } } \ , \ mu _ { t } = 0 \ hfill \ \ \ end { gathered } $ $
) . the daily proportional mortality was computed from the percent mortality at the four highest temperatures shown in the figure by dividing by the egg to adult period ( 477 . 5
) , and dividing by 100 . the mortality rate increases with average daily temperature above 28–45°c ( eq .
d ) . mean daily temperatures above 45°c are unlikely to be encountered in the field .
) in victoria , australia estimate the action of various mortality factors affecting lbam life stages . total generation mortality was approximately 98–99 . 5 % with egg mortality being a key factor during spring and autumn , and egg mortality and losses of first instar during dispersal being key factors during summer . predation of eggs and first instar larvae by generalist predators acting in a density independent manner increased with hot dry weather and nearly eliminated summer generations . similar high rates of mortality were also observed by geier and briese (
despite having been present in california for several years , lbam has failed to reach outbreak levels , even though it has invaded a mediterranean climate similar to that of se australia . recent field observations in california have also shown that the egg and larval stages of lbam are attacked by a suite of generalist predators and parasitoids ( nj mills , unpublished observations ) . in the absence of field life tables for lbam in california , we take a composite approach to incorporating this mortality in a manner that reflects the population levels of lbam observed in the invaded areas . we develop a functional response model ( eq .
, and was chosen iteratively to produce the low population levels that are currently observed in the field .
$ $ 0 \ le \ mu _ { \ text { c } } = 1 - { \ text { e } } ^ { { - 0 . 0025 ( n _ { \ text { eggs } } + n _ { \ text { larvae } } ) \ updelta t } } < 1 $ $
rainfall can have an important indirect effect on lbam dynamics through host plant availability . outside of irrigated crops , herbaceous weedy plants are important hosts for lbam , and as it is an insect with no winter diapause these are often the only host plants available for the period from late summer through spring ( geier and briese
l . ; yst ) as a surrogate annual plant species ( see gutierrez et al .
$ $ { \ frac { { { \ text { d } } n _ { i } } } { { { \ text { d } } t } } } = { \ frac { k \ updelta x } { \ updelta } } [ n _ { i - 1 } ( t ) - n _ { i } ( t ) ] - \ mu _ { i } ( t ) n _ { i } ( t ) . $ $
) includes the proportional net loss rate of births as modified by temperature and age ( fig .
) ) between age classes depends on the numbers in the previous age class ( fig .
, the narrower is the erlang distribution of developmental times . the variance of developmental times reported by danthanarayana (
\ ( { { 335 . 3 \ times ( 0 . 0003 ( t - \ tau ) } \ mathord { \ left / { \ vphantom { { 335 . 3 \ times ( 0 . 0003 ( t - \ tau ) } { ( 1 + 1 . 9 } } } \ right . \ kern - \ nulldelimiterspace } { ( 1 + 1 . 9 } } ^ { ( t - 31 . 5 ) } ) ) \ )
\ ( f ( x , t ) = \ phi ( t ) \ cdot a ( x - 0 . 5 ) / b ^ { x - 0 . 5 } \ )
daily weather data ( maximum and minimum temperatures , solar radiation , precipitation , rh and runs of wind ) for the period 1 january 1995 to 15 november 2006 from 151 locations across california and arizona ( fig .
a ) were used in across year simulations . an initial population density of 50 first instar larvae m
was assumed at all locations in california and arizona and the simulations were run continuously over the 1995–2006 period . data from the first year of simulation were not included in the analysis to allow populations to adjust to the effects of site - specific weather .
weather stations used in the regional study ( a ) , and maximum and minimum temperatures and simulation dynamics of lbam egg and larval dynamics at salinas ( b , c ) and davis , california ( d , e ) during 1 / 1 / 2005 and 15 / 10 / 2006 . the lower ( 6 . 8°c ) and upper ( 28°c ) thresholds below and above which temperature mortality accrues are shown as dashed lines in sub figs . b and d . the location of salinas ( open circle ) and davis ( filled circle ) are illustrated in a
data from the simulation for all locations were geo - referenced and written to files for mapping at elevations below 1 , 000 m . the open source geographic information system (
) ) . raster - based triangulation kriging on a one km grid was used to interpolate the simulation data . the cumulative larval days per year from the simulation was used as an index of the favorability of the climate for the occurrence of lbam at a particular location . this index should not be considered a prediction of either absolute population densities of lbam or potential damage caused by lbam at a particular location . its purpose is merely to reflect the favorability of the climate for lbam population growth .
the relative abundance and phenology of lbam under mild conditions of salinas , ca and more extreme conditions common to davis , ca to show the effects of temperature assuming non limiting host plant availability .
a regional analysis of the favorability of temperature for lbam across all locations in california and arizona assuming non limiting host plant availability .
a regional analysis of climatic favorability for lbam using the limiting effect of rainfall on a surrogate annual plant to illustrate the potential effects of rainfall on the availability of weedy host plants for lbam in california and arizona .
simulations of egg and larval population dynamics using observed maximum and minimum temperatures at salinas , monterey county in coastal northern california and davis , yolo county in central california ( fig .
) were conducted for the period 1 january 2005 to 15 october 2006 to illustrate the effects of differing climatic temperatures on the relative abundance and phenology of lbam . temperatures in salinas were generally mild throughout the year ( fig .
c ) . in contrast , temperatures at davis were colder during winter though rarely freezing , while summers were often very hot ( fig .
e ) with extremes of temperature reducing fecundity and increasing mortality and restricting population growth to the milder times of the year . consequently the relative abundance of egg and larval populations at salinas were predicted to be six and two fold greater , respectively than those at davis .
a map of the locations where lbam adults have been caught in pheromone traps was reported in fowler et al . (
a ) . the moth catches are concentrated in the san francisco and monterey bay areas , but some catches have occurred in coastal regions of southern california .
locations where the moth has been recovered in north - central california ( from fowler et al .
for the period 1996–2006 is used as an index of lbam favorability with respect to temperature for regional mapping ( fig .
b ) . the simulation shows good correspondence between the predicted distribution of lbam and the recorded trap catches in california , especially in the san francisco and monterey bay areas . the model also suggests that the potential distribution of lbam could extend southward along the coastal into southern california ( fig .
b , areas of red–orange - yellow ) , and eastward from san francisco bay into parts of the central valley ( e . g . , davis ) where areas of intermediate favorability ( yellow ) are created by the moderating influence of easterly ocean breezes during summer . these areas of intermediate to high favorability include some important apple , pear and grape growing regions of california . in contrast , most of the central valley shows low to intermediate favorability ( green ) , while the desert valleys of southern california and the deserts of arizona are predicted largely unfavorable because of very high summer temperatures that reduce fecundity and increase mortality ( very light blue to absence of color ) .
) to assess the relative influence of high and low extremes of temperature on the index of favorability . the dependent variable was regressed on cumulative yearly
$ $ \ begin { gathered } { \ text { larval } } \ , { \ text { days } } = 251 . 09 - 0 . 223dd _ { { < 6 . 8 ^ \ circ { \ text { c } } } } - 0 . 297dd _ { { > 28 ^ \ circ { \ text { c } } } } - 0 . 0007dd _ { { < 6 . 8 ^ \ circ { \ text { c } } } } \ times dd _ { { > 28 ^ \ circ { \ text { c } } } } \ hfill \ \ n = 1490 , \ quad r ^ { 2 } = 0 . 80 , \ quad f = 1989 . 2 \ hfill \ \ \ end { gathered } $ $
\ ( \ partial y / \ partial dd _ { { > 28 ^ \ circ { \ text { c } } } } > \ partial y / \ partial dd _ { { < 6 . 8 ^ \ circ { \ text { c } } } } \ )
many mediterranean annual plants germinate during fall and early winter periods of rain and favorable temperatures . for these plants to persist without irrigation , sufficient rainfall for them to grow and produce seed must occur . the invasive yellow starthistle ( yst ) is used here to illustrate the potential effect of host plant availability on the distribution of lbam . yst occurs widely in california except in the southern half of the central valley and the desert regions of arizona and california ( fig .
a ) where it is limited by low variable rainfall that on average is below the 350 mm isohyets ( fig .
) . the combined effects of temperature and limiting rainfall for annual host plants on the potential distribution of lbam may be viewed by overlaying fig .
c . the combination of the two factors reinforces the conclusion that the potential distribution of lbam is largely near coastal with some extension into the sacramento valley . its distribution in southern california and the deserts of arizona is likely to be limited by both high summer temperatures and low levels of precipitation across much of the region , with temperature alone as the limiting factor in irrigated crops .
the simulated average distribution of yellow starthistle in california ( see gutierrez et al .
exotic pests are thought to cause losses in excess of $ 137 billion annually in the us ( pimentel et al .
) , and potential losses due to light brown apple moth are estimated to be $ 118 million per annum for apple , grape , orange and pear crops across the usa ( fowler et al .
, and personal communication ) . leaf rollers are generally not considered primary pests of agricultural crops , and while many , including lbam , have the potential to have direct impact by grazing on the surface of fruit or facilitating
infection within grape clusters , they remain classified as secondary pests due to their amenability to integrated pest management . the experience in new zealand where lbam is also an invasive pest and where climatic conditions of temperature and rainfall are conducive to lbam population development , serves as a valuable example of how its damage can be readily managed as part of an ongoing integrated fruit production program ( walker et al .
in contrast , the usda has classified lbam as an actionable quarantine pest in california based on estimates of its potential geographic distribution in the united states and projected economic losses to apple , grape , orange and pear crops within this predicted distribution ( fowler et al .
) was modeled using nappfast ( north carolina state university animal and plant health inspection service plant pest forecasting system ) based on two simple criteria estimated from a 10 years daily climatology database ; the frequency of degree days sufficient for the development of three or more lbam generations , and the frequency of limiting winter temperatures below −16°c for one full day that would preclude winter survival .
the proposed usda geographic distribution of favorableness for the establishment of lbam ( from fowler et al .
it is well accepted that climate plays a major role in limiting the distribution and abundance of species ( e . g . , andrewartha and birch
) . for poikilotherms , temperature has a strong influence on growth and reproduction , and consequently is often a major determinant of geographic distribution .
many approaches have been used to characterize the favorability of climatic zones for poikilothermic species : comparisons of time series of daily , weekly or monthly temperatures , rainfall , relative humidity , and other variables from the area of origin and the invaded area . a widely accepted approach is the use of physiological indices to estimate the tolerance limits of species to temperature , moisture and other factors ( fitzpatrick and nix
) called these and related methods the ‘climate envelope’ approach , and suggested that conclusions may be misleading because higher tropic levels and important temporal factors and events that affect the dynamics and distributional limits of a species are not included .
the weather - driven demographic model used in this study circumvents many of the criticisms raised by davis et al . (
) , and was used here to map the potential geographic distribution of lbam in arizona and california . this approach requires considerably more weather data to drive the model , which increases biological realism and captures important temporal factors that affect the distribution and abundance of a species ( e . g . , gutierrez et al .
) . lbam populations , although currently established in california , remain sparsely distributed and generally low . here the species is also affected by patterns of temperature and plant availability similar to the patterns in se australia , and lbam in california is also attacked by a suite of native generalist predators and parasitoids ( n . j . mills , unpublished observations ) .
using only temperature , our demographic model predicts the distribution of lbam in california to be restricted largely to near coastal areas with some extension eastward into the northern part of the central valley ( fig .
b ) . the majority of the central valley shows less than intermediate favourability due to summer temperatures that exceed the tolerance of lbam . this suggests that if populations could persist in these areas , the relative abundance of larvae would be lower than those at davis and approximately one - third the abundance predicted in more favorable coastal areas . the limiting effect of rainfall on the potential distribution of yst , a surrogate annual host plant , reinforces the greater favorability of the coastal regions and further highlights the lack of suitable environmental conditions for lbam in the southern half of the central valley ( fig .
) . these predictions of our demographic model differ markedly from those of the fowler et al . (
) . the differences stem primarily from the level of biological detail built into the two models , that of fowler et al . (
) being a simple degree - day accumulation model versus our mechanistic simulation model . our dynamic simulation model has the advantages of incorporating biotic as well as abiotic influences , and daily temperatures rather than weekly , monthly or yearly means , and can more readily track the influence of limiting climatic conditions on lbam populations . the disparity in predictions of the two models also provides support to sutherst and maywald (
) , who have highlighted both model structure , and format and choice of climate variables , in influencing predictions of species distributions .
critics may ask why we have not validated the model by predicting the distribution of lbam in australia , and may posit that lbam is present in regions of australia that are much hotter than the regions of california that our model predicts as unfavorable . in the absence of sufficient daily climate data to validate the model for the whole country , we examined daily climate data for the period 1 / 1 / 1998–1 / 1 / 2009 from five locations in se australia ( victoria and new south wales ) where lbam is known to occur . mean daily maximum summer temperatures at mildura ( vic ) and griffith ( nsw ) , the two hottest locations , prove similar to those at davis , ca , while winter temperatures are lower ( fig .
) . our model predicts moderate favourability for lbam at the four hotter inland locations in australia , as was the case for davis in california , and greater favourability for the coastal location of adelaide ( fig .
) found lbam infestations were most frequent ( and most damaging ) in the areas seemingly least favorable for its development , namely cooler southern victoria and tasmania . as for the comparison between davis and salinas in california ( fig .
) , the relative abundance of lbam is also predicted to be approximately twice as high in adelaide as at the inland locations in australia ( fig .
any assessment of the pest status of an invasive species must be based on a sound understanding of its likely range of distribution in the new region , in addition to an appraisal of the potential ecological and economic impact that it could have on production and natural ecosystems . the current designation of lbam as an actionable quarantine pest in the united states by usda is in part based upon the predicted distribution of fowler et al . (
) . the strong decline in favourability from coastal to interior regions of california and its likely absence from arizona , as predicted by our model , question the accuracy of the usda assessment of risk to production and natural ecosystems and suggest the need for re - evaluation of the current pest status of lbam in the united states .
this article is distributed under the terms of the creative commons attribution noncommercial license which permits any noncommercial use , distribution , and reproduction in any medium , provided the original author ( s ) and source are credited .
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