stalk-eyed flies are insects of the fly family diopsidae .

the family is distinguished from most other flies by the possession of " eyestalks " : projections from the sides of the head with the eyes at the end .

some fly species from other families such as drosophilidae , platystomatidae , richardiidae , and tephritidae have similar heads , but the unique character of the diopsidae is that their antennae are located on the stalk , rather than in the middle of the head as in all other flies .

the stalk-eyed flies are up to a centimeter long , and they feed on both decaying plants and animals .

their unique morphology has inspired research into how the attribute may have arisen through forces of sexual selection and natural selection .

studies of the behavior of the diopsidae have yielded important insights into the development of sexual ornamentation , the genetic factors that maintain such a morphological feature , sexual selection , and the handicap principle .

stalk - eyed fly

stalk eyed fly pc : rob knell ( cc by sa 2 . 5 )
signalling fitness : larger males sire more offspring . studies of the stalk - eyed fly
. signalling fitness : larger males sire more offspring . studies of the stalk - eyed fly
female choice response to artificial selection on an exaggerated male trait in a stalk - eyed fly .
female preference response to artificial selection on an exaggerated male trait in a stalk - eyed fly .
mating - induced reduction in accessory reproductive organ size in the stalk - eyed fly cyrtodiopsis dalmanni .
. female preference response to artificial selection on an exaggerated male trait in a stalk - eyed fly .
hingle a , fowler k , pomiankowski a . size - dependent mate preference in the stalk - eyed fly
lorch pd , wilkinson gs , reillo pr . copulation duration and sperm precedence in the stalk - eyed fly
burkhardt d , de la motte i . how stalk - eyed flies eye stalk - eyed flies : observations and measurements of the eyes of
burkhardt d . and motte i , 1983 . how stalk - eyed flies eye stalk - eyed flies : observations and measurements of the eyes of
university college london , department of biology ; 2003 . the evolution of multiple mating in the stalk - eyed fly ,
mating - induced reduction in accessory reproductive organ size in the stalk - eyed fly cyrtodiopsis dalmanni . - pubmed - ncbi
condition dependence of sexual ornament size and variation in the stalk - eyed fly cyrtodiopsis dalmanni ( diptera : diopsidae ) .
whether sexually selected traits are sex linked can have profound effects on their evolution . in the diopsid stalk - eyed fly ,
cotton s , fowler k , pomiankowski a . condition dependence of sexual ornament size and variation in the stalk - eyed fly
male sexual ornament size but not asymmetry reflects condition in stalk - eyed flies .
evolution of genetic variation for condition - dependent traits in stalk - eyed flies .
. coevolution of sperm and female reproductive tract morphology in stalk - eyed flies .
. evolution of genetic variation for condition dependent traits in stalk - eyed flies .
visual system of the stalk - eyed fly , cyrtodiopsis quinqueguttata ( diopsidae , diptera ) : an anatomical investigation of unusual eyes .
wilkinson gs , reillo pr . female preference response to artificial selection on an exaggerated male trait in a stalk - eyed fly .
hingle a , fowler k , pomiankowski a . the effect of transient food stress on female mate preference in the stalk - eyed fly
. male sexual ornament size but not asymmetry reflects condition in stalk - eyed flies .
wilkinson g . s . , 1993 . artificial selection alters allometry in the stalk - eyed fly cyrtodiopsis dalmanni ( diptera : diopsidae ) .
condition dependence of sexual ornament size and variation in the stalk - eyed fly cyrtodiopsis dalmanni ( diptera : diopsidae ) . - pubmed - ncbi
selective pressures , variability , and sexual dimorphism in stalk - eyed flies ( diopsidae ) .
. male eye span in stalk - eyed flies indicates genetic quality by meiotic drive suppression .
rogers dw , chapman t , fowler k , pomiankowski a . mating - induced reduction in accessory reproductive organ size in the stalk - eyed fly
. hunting for ayv 28—right and wrong approaches in an attempt to increase our knowledge about the stalk - eyed fly ( diopsidae , diptera ) .
visual system of the stalk - eyed fly , cyrtodiopsis quinqueguttata ( diopsidae , diptera ) : an anatomical investigation of unusual eyes . - pubmed - ncbi
panhuis tm , wilkinson gs . exaggerated eyespan influences male contest outcome in stalk - eyed flies .
wilkinson gs , fry cl . meiotic drive alters sperm competitive ability in stalk - eyed flies .
hurley i , pomiankowski a , fowler k , smith h . , 2002 . fate map of the eye antennal imaginal disc in the stalk - eyed fly
so , the benefit that the compound eye gives the fly is that the fly can see . it lets the fly know if something is coming towards it , where the fly is positioned in its environment , what’s there , and tells the fly that it’s moving in relation to other things .
rogers dw , grant ca , chapman t , pomiankowski a , fowler k . the influence of male and female eyespan on fertility in the stalk - eyed fly
stalk - eyed flies ( diopsidae ) : modelling the evolution and development of an exaggerated sexual trait .
presgraves dc , severance e , wilkinson gs . sex chromosome meiotic drive in stalk - eyed flies .
figure 16 : a species of stalk - eyed flies photographed in dairy farm . courtesy of nicky bay .
wilkinson gs , kahler h , baker rh . evolution of female mating preferences in stalk - eyed flies .
is the only stalk - eyed fly in the micropezidae family – and we only have one , slightly damaged specimen in the collection . on the right : the essay unwrapped .
wilkinson gs . genetic consequences of sexual selection in stalk - eyed flies . in : dugatkin la , editor .
cotton s , rogers dw , small j , pomiankowski a , fowler k . variation in preference for a male ornament is positively associated with female eyespan in the stalk - eyed fly
the full importance of the accessory glands in stalk - eyed fly reproduction is poorly understood . accessory gland products form the casing of the spermatophore and consequently are necessary for sperm transfer [
fry cl . juvenile hormone mediates a trade - off between primary and secondary sexual traits in stalk - eyed flies .
baker rh , denniff m , futerman p , fowler k , pomiankowski a , chapman t . accessory glands influence time to sexual maturity and mating frequency in the stalk - eyed fly ,
rogers dw , baker rh , chapman t , denniff m , pomiankowski a , fowler k . direct and correlated responses to artificial selection on male mating frequency in the stalk - eyed fly
lorch p . , wilkinson g . s . , reilo pr . , 1993 . copulation duration and sperm precedence in malaysian stalk - eyed fly , cyrtodiopsis whitei ( diptera : diopsidae ) .
baker rh , ashwell ris , richards ta , fowler k , chapman t , pomiankowski a . effects of multiple mating and male eye span on female reproductive output in the stalk - eyed fly
stalk - eyed flies ( diopsidae ) : modelling the evolution and development of an exaggerated sexual trait . - pubmed - ncbi
reguera p . , and pomiankowski , a . , 2004 . low cost of reproduction in female stalk - eyed flies ,
reguera p , pomiankowski a , fowler k , chapman t . low cost of reproduction in female stalk - eyed flies ,
wilkinson gs , reillo pr : female choice response to artificial selection on an exaggerated male trait in a stalk - eyed fly . proc r soc lond b . 1994 , 255 : 1 - 6 .
burkhardt d , de la motte i . selective pressures , variability , and sexual dimorphism in stalk - eyed flies ( diopsidae )
wilkinson gs et al . ,
sex - biased gene expression during head development in a sexually dimorphic stalk - eyed fly .
, plos one , 2013 mar 19 ; 8 ( 3 ) : e59826
big ‘antlers’ are favoured : female choice in stalk - eyed flies ( diptera , insecta ) , field collected harems and laboratory experiments .
david p , bjorksten t , fowler k , pomiankowski a . condition - dependent signalling of genetic variation in stalk - eyed flies .
sukontason kl , chaiwong t , piangjai s , upakut s , moophayak k , et al . ( 2008 ) ommatidia of blow fly , house fly , and flesh fly : implication of their vision efficiency . parasitol res 103 : 123–131 .
kotrba m . , 2004 . baltic amber fossils reveal early evolution of sexual dimorphism in stalk - eyed flies ( diptera : diopsidae ) .
the australian museum has the world ' s largest and most comprehensive collection of signal flies , including examples from 64 different species of stalk - eyed signal fly and a number of unique specimens found in no other museum collection .
sukontason kl , chaiwong t , piangjai s , upakut s , moophayak k , and sukontason k . 2008 . ommatidia of blow fly , house fly , and flesh fly : implication of their vision efficiency . parasitology research 103 : 123 - 131 .
burkhardt d . , and motte i . , 1985 . selective pressure , variability and sexual dimorphism in stalk - eyed flies ( diopsidae ) .
stalk - eyed signal flies occur mainly in new guinea ( with more than 90 recorded species ) and in queensland ( with five recorded species ) .
. big ` antlers ' are favoured : female choice in stalk - eyed flies ( diptera , insecta ) , field collected harems and laboratory experiments .
male stalk - eyed signal flies spend much of their time in rainforests on shaded tree trunks which they use as courtship territory and avidly defend against rivals .
the heritability of sexually dimorphic traits in the yellow dung fly scatophaga stercoraria ( l . )
a male big eyed fly . its entire head is a pair of eyes . ( diptera : pipunculidae ) pc : marcello consolo ( cc by sa 2 . 0 )
panhuis t . m . , wilkinson g . s . , 1999 . exaggerated male eye span influences contest outcome in stalk - eyed flies ( diopsidae ) .
wright tf , johns pm , walters jr , lerner ap , swallow jg , wilkinson gs . microsatellite variation among divergent populations of stalk - eyed flies , genus
^ wilkinson g . s . , and reillo p . r . , 1994 . female choice response to artificial selection on an exaggerated male trait in a stalk - eyed fly . proceedings of biological sciences . 255 ( 342 ) : 1 - 6 .
sometimes evolution just doesn’t care if you can see and it’s only important how sexy you are . that’s what happened to the stalk eyed fly . the outrageous stalks that the males carry around actual hinder their flying capabilities but they can still see relatively well .
a soldier fly . the red color acts as a pair of sunglasses . pc : eddie smith
studies on the egg morphology of stalk - eyed flies using scanning electron microscope ( sem ) demonstrated that egg morphology is useful for identification purposes and for cladistic analysis .
baker rh , wilkinson gs , desalle r . the phylogenetic utility of different types of molecular data used to infer evolutionary relationships among stalk - eyed flies ( diopsidae )
^ rihak g . , egge a . r . , swallow g . s . , 2009 . saccadic head rotations during walking in the stalk - eyed fly ( cyrtodiopsis dalmanni ) . proceedings of the royal society of biological sciences . 276 : 1643 - 1649 .
this video shows three stalk - eyed fly species from borneo ( teleopsis pallifacies , eurydopsis sarawakensis , cyrtodiopsis sp . ) and features information on their mating biology . it also shows a surprise attack ! ! follow me on twitter @ magsorger # crazybeautifulnature visit my website urltoken
figure 3 : eyes of a stalk - eyed flies , showing the position of antenna beside the compund eye . [ image courtesy of tronghieusg , used with permission . ]
. this suggest that the behavior of aggregating and intrasexual competition between males , besides sexual selection , is a selection force for sexual dimorphsim in the stalk - eyed flies .
fry cl , wilkinson gs : sperm survival in female stalk - eyed flies depends on seminal fluid and meiotic drive . evolution . 2004 , 58 : 1622 - 1626 .
baker rh et al . ,
genomic analysis of a sexually - selected character : est sequencing and microarray analysis of eye - antennal imaginal discs in the stalk - eyed fly teleopsis dalmanni ( diopsidae ) .
, bmc genomics , 2009 aug 5 ; 10 : 361
male and female of a blow fly . ( chrysomya rufifacies ) pc : sukontason et al . 2008
life returns to discovery channel in october - sundays starting 10 / 3 @ 8 and 9p ! urltoken stalk - eyed flies inflate their long , tube - like eye stalks .
the most notable example where this occurs is the family of flies called diopsidae . every species has eye stalks and so they are the ones commonly called the stalk - eyed flies .
it should be noted that stalk - eyed flies are not the only group of dipteran flies that posses eye stalks ( figure 2 ; click here for examples of dipteran with eye stalk ) . however , they are unique by having the antenna near the end of the eyes stalk , next to the eye instead of in the facial region ( figure 3 ) .
burkhardt d , de la motte i . big ' antlers ' are favoured : female choice in stalk - eyed flies ( diptera , insecta ) , field collected harems and laboratory experiments .
stalk - eyed flies ( family diopsidae ) are a model system for studying sexual selection due to the elongated and sexually dimorphic eye - stalks found in many species . more . . .
^ wilkinson g . s . , johns p . m . , kelleher e . s . , muscedere m . l . , lorsong a . , 2006 . fitness effects of x chromosome drive in the stalk - eyed fly , cyrtodiopsis dalmanni . journal of evolutionary biology . 19 : 1851–1860 .
is definitely the lateral head projection ( the “eye stalk” ) , which bore the eyes and the antenna . the eye stalk is derived from the posterior half of the eye - antennal disc
we have shown that the pattern of sperm usage is highly variable in the stalk - eyed fly , t . dalmanni . this greatly limits the utility of population - based estimates of p 2 as descriptors of sperm usage , and suggests that sperm precedence should be viewed as context - specific , rather than a general , constant , metric in stalk - eyed flies . the unexplained variance in male fertilization success found by this study requires further investigation in order to evaluate potential causes and consequences .
^ cotton s . , small j . , hashim r . , pomiankowski a . , 2010 . eyespan reflects reproductive quality in wild stalk - eyed flies . evolutionary ecology . 24 : 83–95 .
husak j . f . , ribak g . , wilkinson g . s . , swallow j . g . , 2011 . compensation for exaggerated eye stalks in stalk - eyed flies ( diopsidae ) .
1 . outer vertical bristle black twice as long as width of eye - stalk in the middle .
most times this just involves squaring up to each other like alcohol - infused humans . but sometimes the fights become more physical , with males head butting each other or having fist fights . yes , proper fly - on - fly boxing .
^ rogers d . w . , baker r . h . , chapman t . , denniff m . , pomiankowski a . , fowler k . , 2005 . direct and correlated responses to artificial selection on male mating frequency in the stalk - eyed fly cyrtodiopsis dalmanni . journal of evolutionary biology . 18 : 642 - 650 .
stalk - eyed flies of the family diopsidae exhibit a unique form of hypercephaly , which has evolved under both natural and sexual selection . male hypercephaly is used by female diopsids as an indicator of male quality . by choosing to mate with males expressing the most - exaggerated hypercephaly , females can benefit both from the enhanced fertility of these males and the transmission of other heritable advantages to their offspring . stalk - eyed flies are close relatives of the model organism , drosophila melanogaster . we have shown that similar genetic and cellular mechanisms regulate the initial development of the head capsule in fruitflies and diopsids . the great diversity of stalk - eyed fly species , exhibiting varying degrees of hypercephaly and sexual dimorphism , constitutes a major advantage for comparative studies of their development and evolution .
my first experience of stalk - eyed flies came while i was carrying out fieldwork in costa rica over 10 years ago and it can probably go down as one of my favourite fieldwork moments . so what happened ?
it is not just the richardiidae family that contain stalk - eyed species , in fact the stalk - eyed condition has evolved independently in at least eight families of flies . in these families , and generally only in males , the eyes have migrated away from the head and are now found on the end of stalks . called hypercephalization , this condition also includes examples where elaborate structures such as antlers and horns protrude from the head .
richard h . baker , robert i . s . ashwell , thomas a . richards , kevin fowler , tracey chapman , andrew pomiankowski ; effects of multiple mating and male eye span on female reproductive output in the stalk - eyed fly , cyrtodiopsis dalmanni , behavioral ecology , volume 12 , issue 6 , 1 november 2001 , pages 732–739 , urltoken
diopsinae is a taxonomic group of fly in which members posse the eye stalks . this group is nested within the diopsidae , the stalk - eyed flies . however , taxonomists do not agree with what constitute the diopsidae . the traditional definition of diopsidae includes the centrioncinae , in which members do not have the eye stalks , and the diopsinae . however , some taxonomists argue that the centrioncinae should not be included in the diopsidae , and should treat as a different group . [ 20 ] stalk - eyed flies in this page refers to the members of diopsidae that includes centrioncinae .
this group of flies contains some spectacular species including the stalk - eyed signal flies , belonging to the genus achias . true to their name , the males have eyes on long stalks extending from either side of their heads .
the signal fly family ( platystomatidae ) includes a great diversity of species found in australia , new guinea and other nearby tropical countries .
scientists working on this family estimate that there may be as many as 900 species of signal fly that occur in the australasian region .
kirschfeld k , franceschini n , minke b ( 1977 ) evidence for a sensitising pigment in fly photoreceptors . nature 269 : 386–390 .
the common horse fly may seem an ugly nuisance from far away , but in the macro perspective , its eyes draw you in .
the band - eyed drone fly is a bee mimic , and it takes the ruse to the limit , with stripes even on its eyes . but why ? scientists hypothesize that it affects what this insect sees — definitely a subject for further research .
land mf , eckert h ( 1985 ) maps of the acute zones of fly eyes . journal of comparative physiology a 156 : 525–538 .
this deer fly has a distinct pattern on its eyes . in fact , their genus , chrysops , translates to
gold eyes .
most of the described stalk - eyed flies species are found in the old world with the afrotropical region having the most species . there is only two species found in the new world and are found in temperate north america . the distribution of
this entry was posted in ecology , physiology and tagged compound eye , diptera , eyes , fly , insect , ommatidia . bookmark the permalink .
ribak g and swallow jg . 2007 . free flight maneuvers of stalk - eyed flies” do eye - stalks effect aerial turning behavior ? journal of comparative physiology a neuroethology , sensory , nuerual , and behavioral physiology 193 ( 10 ) : 1065 - 1079 .
till date , there is no proper studies in surveying the diversity of stalk - eyed flies in singapore , but they have been photographed by photography enthusiasts in matritchie reservoir park , dairy farm ( figure 16 ) , singapore botanic garden , bukit timah and nee soon swamp forest ( map ) . teleopsis dalmanni could be found in central catchement area . stalk - eyed flies are generally found at shady areas , near edge of forest streams , or in wet muddy area with rotten vegetation in proximate ( figure 17 ) .
imagine seeing through the extended eyes of the stalk - eyed fly ! these funky - looking bugs can extend their eyes outward by filling their heads with air as they make their final transformation from pupa to adult ( see this amazing process in this discovery
life
video ) . while it seems as though these outward - facing eyes might provide superior vision , the main purpose is to attract female flies .
cotton , a j földvári , m cotton , s and pomiankowski , a 2014 . male eyespan size is associated with meiotic drive in wild stalk - eyed flies ( teleopsis dalmanni ) . heredity , vol . 112 , issue . 4 , p . 363 .
wernet mf , desplan c ( 2004 ) building a retinal mosaic : cell - fate decision in the fly eye . trends in cell biology 14 : 576–584 .
update ( 22 jan . 13 ) : thanks to hans feijen for identifying the fly species , which turned out to be diasemopsis fasciata , and not diopsis .
fly , ( order diptera ) , any of a large number of insects characterized by the use of only one pair of wings for flight and the reduction of the second pair of wings to knobs ( called halteres ) used for balance . the term fly is commonly used for almost any small flying insect . however , in entomology…
bath , eleanor wigby , stuart vincent , claire tobias , joseph a . and seddon , nathalie 2015 . condition , not eyespan , predicts contest outcome in female stalk - eyed flies , teleopsis dalmanni . ecology and evolution , vol . 5 , issue . 9 , p . 1826 .
bonduriansky r , rowe l ( 2003 ) interactions among mechanisms of sexual selection on male body size and head shape in a sexually dimorphic fly . evolution 57 : 2046–2053 .
within the fruit fly family , tephritidae , there are two species that have stalked eyes . one of them , pelmatops ichneumonea , was once placed in a different genus and family . in fact , it is easy to see at first glance why this happened when you compare it with achias , the genus it was originally described as . achias is now within the platystomatidae family and contains arguably the most impressive of all of the stalk - eyed flies – achias rothschildii .
^ wilkinson , g . s . , amitin , e . g . and johns , p . m . , 2005 . sex - linked correlated responses in female reproductive traits to selection on male eye span in stalk - eyed flies . integrative and comparative biology . 45 : 500 - 510 .
shortly after eclosion , before their structures hardened , t . dalmanni ingest air thorough their oral cavity , and pump it to the tips to their stalk through a duct in the head , thereby elongating their stalk . watch the video to see an newly - emerged adult flies pumping their eye stalks !
3 . inner vertical bristle up to 4 . 5–5 times as long as width of eye - stalk in the middle ; on a small tubercle .
^ ribak g . , pitts m . l . , wilkinson g . s . , swallow g . s . , 2009 . wing shape , wing size , and sexual dimorphism in eye - span in stalk - eyed flies ( diopsidae ) . biological journal of the linnean society . 98 : 860–871 .
stalk - eye flies shared the same general life history pattern ( figure 18 ) , although fecundity , age of sexual maturity and longevity varies among species .
^ worthington a . m . , berns c . m . , and swallow j . g . , 2012 . size matters , but so does shape : quantifying complex shape changes in a sexually selected trait in stalk - eyed flies ( diptera : diopsidae ) . biological journal of the linnean society . 106 : 104–113 .
hardie rc ( 1985 ) functional organization of the fly retina . progress in sens physiol . berlin , heidelberg , new york , toronto : springer , vol . 5 . pp . 1–79 .
before the museum exhibition about colour and vision closes on 6 november , i thought i should write a piece about some of nature’s most amazing eyes ( their patterns and shapes ) . i’m talking of course about those belonging to flies – the most enigmatic of all species on the planet – and specifically all the species referred to as stalk - eyed flies .
within micropezidae there is only one species that exhibits the stalk - eyed condition : anaeropsis guttipennis . and within our collection there is only one , slightly damaged specimen that only has one stalk ! however beside this specimen is an essay – albeit the smallest of essays . unwrapped , this essay ( by ernest edward austin – a great dipterist and my forefather in curation terms ) describes how several authors have moved the taxonomic placement of this species around . ernest felt that any decisions to be made needed further proof and greater comparisons – wise words .
fascinating , erica ! do such structures occur in just flies ? is it possible that they fulfill functions other than visual superiority of some kind and sexual prowess ? i note the forward rake in some species , which half suggests equivalence to antennae in lepidoptera… that visual superiority , if it exists… i wonder if having the eyes on stalks enables fuller all - round vision – because the optically - facetted part of the stalk - side of each eye fills - in the blind spot in normal flies caused by the head . that might be an evolutionary driver for just thin - stalked eyes . but then why the species with thick - stalked eyed ? one also wonders if the stalked - eyes structure helps with orientation awareness , it’s inertia tending to keep it at a fixed orientation , in turn enabling the fly to sense changes in the orientation of its body . …related to the gyroscopic function of halteres . but then , non - stalk - eyed flies seem to manage well enough without… …so many questions… mike
the base stock was an outbred laboratory population of the stalk - eyed fly , c . dalmanni , collected from gombak , malaysia in 1993 . the stock was maintained in large cages at high population size ( typically more than 200 individuals per cage ) and with a 1 : 1 sex ratio . flies were fed ground corn medium and kept at 25°c on a 12 h / 12 h light / dark regime . the regime included a 15 - min
dawn
period in which the culture room was illuminated by a single 60 - w bulb . all observations of behaviour commenced at the start of this dawn period .
that is , of course , if the organism needs to see . eyes are usually the first things to disappear if you live in caves . or if you’re a fly maggot that invests its early days face first eating rotting corpses .
the distribution of second male sperm precedence ( p 2 ) among 22 dam - sire pair families in the stalk - eyed fly , teleopsis dalmanni . open bars depict the frequency of families with p 2 significantly different from 0 . 5 ( using 2 - tailed binomial tests ) ; black bars denote those with p 2 not significantly different from 0 . 5 . the grey bar denotes the sperm precedence of a single family where the observed p 2 of zero was not significantly different from p 2 = 0 . 5 . however , the brood size of this family was small ( n = 5 ) , so this result should be treated with caution .
internal reproductive organ size is an important determinant of male reproductive success . while the response of testis length to variation in the intensity of sperm competition is well documented across many taxa , few studies address the importance of testis size in determining other components of male reproductive success ( such as mating frequency ) or the significance of size variation in accessory reproductive organs . accessory gland length , but not testis length , is both phenotypically and genetically correlated with male mating frequency in the stalk - eyed fly cyrtodiopsis dalmanni . here we directly manipulate male mating status to investigate the effect of copulation on the size of both the testes and the accessory glands of c . dalmanni .
internal reproductive organ size is an important determinant of male reproductive success . while the response of testis length to variation in the intensity of sperm competition is well documented across many taxa , few studies address the importance of testis size in determining other components of male reproductive success ( such as mating frequency ) or the significance of size variation in accessory reproductive organs . accessory gland length , but not testis length , is both phenotypically and genetically correlated with male mating frequency in the stalk - eyed fly cyrtodiopsis dalmanni . here we directly manipulate male mating status to investigate the effect of copulation on the size of both the testes and the accessory glands of c . dalmanni .
since males with longer eye span gain benefits of having more mating opportunities , there is a strong sexual selection force for increasing longer eye stalks in males . however , there is a limit in the length of the eye stalks . the process of developing longer eye stalk could create developmental problems in the larvae stage , leading to high larvae mortality rate in the pupae stage . longer eye stalk also requires longer development time , which increase the exposure to toxic metabolites produced by conspecific larva , as well as to predators and parasites . therefore , the length of male eye stalk is maintain by natural selection opposing the force of sexual selection .
we thank t . chapman and g . hurst for comments , and a . hingle for help in rearing fly stocks . this work was supported by royal society university research fellowships ( k . f . and a . p . ) and the nerc .
there are about 150 , 000 described species of described true flies ( diptera ) with an estimated total number of fly species to be around 240 , 000 . so , this is going to be very generalized and does not at all encompass every organism .
male mating history is often associated with changes in male investment in current mating attempts , with concomitant effects on patterns of paternity [ 12 , 13 ] . however , all males were virgins at the start of the experiment and performed equal numbers of copulations , so variation in mating experience did not differ between first and second males , and hence is not an explanation of our data . in addition , all males were maintained on a high quality diet , so variation in environmental conditions was minimised between pairs of sires . these factors ( when variable ) may well be important in determining stalk - eyed fly paternity and deserve further investigation , but they cannot explain the results reported in the current study .
in some species , females choose mates possessing ornaments that predict offspring survival 1 , 2 , 3 , 4 , 5 . however , sexual selection by female preference for male genetic quality 6 , 7 , 8 remains controversial because conventional genetic mechanisms maintain insufficient variation in male quality to account for costly preference and ornament evolution 9 , 10 . here we show that females prefer ornaments that indicate genetic quality generated by transmission conflict between the sex chromosomes . by comparing sex - ratio distributions in stalk - eyed fly ( cyrtodiopsis ) progeny we found that female - biased sex ratios occur in species exhibiting eye - stalk sexual dimorphism 11 , 12 and female preferences for long eye span 13 , 14 . female - biased sex ratios result from meiotic drive 15 , the preferential transmission of a ‘selfish’ x - chromosome . artificial selection for 22 generations on male eye - stalk length in sexually dimorphic c . dalmanni produced longer eye - stalks and male - biased progeny sex ratios in replicate lines . because male - biased progeny sex ratios occur when a drive - resistant y chromosome pairs with a driving x chromosome 15 , long eye span is genetically linked to meiotic drive suppression . male eye span therefore signals genetic quality by influencing the reproductive value of offspring 16 .
when females mate with different males , competition for fertilizations occurs after insemination . such sperm competition is usually summarized at the level of the population or species by the parameter , p 2 , defined as the proportion of offspring sired by the second male in double mating trials . however , considerable variation in p 2 may occur within populations , and such variation limits the utility of population - wide or species p 2 estimates as descriptors of sperm usage . to fully understand the causes and consequences of sperm competition requires estimates of not only mean p 2 , but also intra - specific variation in p 2 . here we investigate within - population quantitative variation in p 2 using a controlled mating experiment and microsatellite profiling of progeny in the multiply mating stalk - eyed fly , teleopsis dalmanni .
colored eyes , like those of the soldier fly above , serve a specific purpose in insects : they narrow down the information being processed by each of the tiny lenses . some insects with metallic - colored eyes have layers and layers of lenses that reflect light , lending an iridescent quality .
the length of eye stalk strongly affects the outcome of the males’ competition over a female aggregation . males with larger eyespan , independent of body size , are more aggressive and win proportionately more aggressive interaction than males with smaller eye span [ 6 ]
micropezidae , diopsidae , ottidae , platystomatidae , tephritidae , richardiidae , periscelididae and drosophilidae all contain some species that exhibit some level of hypercephalization and , interestingly , the stalk - eyed condition has evolved independently more than once in platystomatidae . this is a very good example of convergent evolution – where species that are not related have a very similar form ( the hedgehog and the spiny anteater are another good example ) . it is also an example of recurrent evolution – where a similar feature evolves time and time again .
we used the stalk - eyed fly cyrtodiopsis dalmanni to examine predictions made by condition - dependent handicap models of sexual selection . condition was experimentally varied by manipulation of larval food availability . cyrtodiopsis dalmanni is a highly dimorphic species exhibiting strong sexual selection , and the male sexual ornament ( exaggerated eyespan ) showed strong condition - dependent expression relative to the homologous trait in females and nonsexual traits . male eyespan also showed a great increase in standardized variance under stress , unlike nonsexual traits . the inflated variance of the male ornament was primarily attributable to condition - dependent ( but body - size - independent ) increase in variance . thus , evaluation of male eyespan allows females to gain additional information about male condition over and above that given by body size . these findings accord well with condition - dependent handicap models of sexual selection .
under the handicap hypothesis of mate choice , female judge the quality of males through sexual ornamentation , such as the eye stalk , which are costly to develop . a high - quality male will be able to support longer eye stalks which are costly despite poor condition
males come head to head and primarily compare stalk length . if there is a difference the male with the smaller stalks leaves and the larger male retains control of his territory ( and often harem ) . however , when they judge themselves as equal they fight !
figure 1 : mapping of compound eyes ( red ) , antenna ( green ) and eye - antenna dics ( blue ) in the stalk - eyed flies ( diopsidae ) , showing a gradual evolution of eye - stalks from condition ' a ' through ' b ' and ' c ' to ' d ' : ( a ) centrioncus prodiopsis speiser ; ( b ) sphyracephala beccari ; ( c ) sphyracephala ( pseudodiopsis ) detrahens walker ; ( d ) diopsis thoracica westwood . ( extracted and modified from meier & baker , 2002 )
stalk - eyed flies ( diptera : diopsidae ) are increasingly important model organisms for studies of sexual selection [ 19 - 22 ] . they are characterised by the lateral extension of the eyes on elongate protuberances on the side of the head capsule , a trait common to both sexes in all species [ 19 , 23 ] . in many species the eyespan of males is greater than that of females , the result of sexual selection through female mate choice [ 24 - 29 ] and male - male competition [ 30 , 31 ] .
the authors thank matthew denniff for assistance with fly rearing and behavioural observations , and two anonymous referees for their helpful comments on the manuscript . this research was funded by awards from the natural environment research council ( tc , kf & ap ) , the royal society ( tc ) and the association of commonwealth universities ( dwr ) .
both male and female stalk - eyed flies mate frequently . in the current study , each male mated an average of 3 . 79 times ( up to a maximum of 12 ) during the 60 - minute observation period . only 23 . 9 % ( 37 out of 155 ) of males mated at least 6 times and therefore 76 . 1 % ( 118 out of 155 ) of males failed to mate with all 6 virgin females provided . as females housed with three males will mate an average of 5 . 51 times during the 60 minutes following artificial dawn [
flies that rely heavily on their vision , for the most part , have cashed in on all of the options . house flies ( musca domestica ) have a moderate amount of ommatidia , but males have more ommatidia ( ~ 3 , 500 ) than females ( ~ 3 , 400 ) and bigger eyes , suggesting that vision plays an important role in mate determination . in fact , this pattern is readily seen in two other families of flies , the flesh flies ( sarcophagidae ) and the blow flies ( calliophoridae ) . house flies were at the lower end of the spectrum with some blow files coming in closer to the ~ 5 , 500 ommatidia honey bees have . while this isn’t the 30 , 000 that dragonflies have , each ommatidium of a house fly , flesh fly , or blow fly is bigger than that of a dragonfly . plus they still have a lot more than the 2 , 000 american cockroaches have and the 800 that drosophila have . so we’ll call it a nice middle ground .
when fighting face to face , rival males gauge their opponents ' size and strength from their head width . small males ( which may have begun life as under - nourished or disadvantaged larvae ) have very short or no eye - stalks . the clear difference in head - width allows the weaker fly to back off early and avoid possible injury .
many little gnats and mosquitoes are crepuscular meaning they fly at dawn and at dusk but flies generally have eye modifications for frolicking around in the sunlight . normally they’d be out of luck seeing once the sun dipped below the horizon , but flies have that 7 way split rhabdom . this helps them fly in low light conditions because the 7 parts of the rhabdom are separated and act in a similiar manner to the rhabdoms separated from the ommatidia by the clear zone . specifically for gnats and mosquitoes , it gives them an extra 15 or so minutes before dawn and after sunset . this doesn’t seem like much for us , but it gives them a small window to swarm , mate , and feed without being incredibly visible to predators .
i was walking along a path in the forest near the tropical field station la suerte ( where i was based for a month or so ) when i saw this strange creature – a hammerhead fly – scuttling over the bark of a tree . i chased it round and round the tree and finally managed to catch it in my hands ! it turned out to be this little beast –
handicap models of sexual selection predict that male sexual ornaments have strong condition - dependent expression and this allows females to evaluate male genetic quality 1 , 2 , 3 , 4 , 5 . a number of previous experiments have demonstrated heightened condition - dependence of sexual ornaments in response to environmental stress 6 , 7 , 8 , 9 . here we show that genetic variation underlies the response to environmental stress ( variable food quality ) of a sexual ornament ( male eye span ) in the stalk - eyed fly cyrtodiopsis dalmanni . some male genotypes develop large eye span under all conditions , whereas other genotypes progressively reduce eye span as conditions deteriorate . several non - sexual traits ( female eye span , male and female wing length ) also show genetic variation in condition - dependent expression , but their genetic response is entirely explained by scaling with body size . in contrast , the male sexual ornament still reveals genetic variation in the response to environmental stress after accounting for differences in body size . these results strongly support the hypothesis that female mate choice yields genetic benefits for offspring .
females of the stalk - eyed fly , cyrtodiopsis dalmanni , mate repeatedly during their lifetime and exhibit mating preference for males with large eye span . how these mating decisions affect female fitness is not fully understood . in this study , we examined the effects of multiple mating and male eye span on short - term reproductive output in this species . experiments that manipulated the number of copulations and partners a female received suggested that obtaining a sufficient sperm supply is an important benefit associated with multiple mating . the average percentage of fertile eggs laid by females increased as a function of mating frequency and ranged from 40 % for females mated once , to 80 % for females mated continuously . in addition , a high proportion of copulations in this species appeared to be unsuccessful . one - third of all females mated once laid less than 10 % fertile eggs . there was no significant difference in reproductive performance between females mated to multiple partners and females mated to a single partner . there was also no indication that females received any short - term reproductive benefits from mating with males with large eye span . in fact , females mated to males with short eye span laid a higher percentage of fertile eggs than females mated to large eye span males .
the malaysian stalk - eyed fly , teleopsis dalmanni ( formerly known as cyrtodiopsis dalmanni ; [ 32 ] ) , exhibits extreme sexual dimorphism for eyespan resulting from strong intra - and inter - sexual selection on the trait in males ( ibid . ) . females form large harems on root hairs overhanging the eroded banks of streams and males compete to control these harems [ 19 , 33 ] . both sexes are highly promiscuous and mate at high frequencies ( ~ 10 and 6 times per hour in the laboratory , for males and females respectively ; [ 34 , 35 ] ) . there is also some evidence that males strategically allocate more ejaculate to larger , more productive females through the production and delivery of larger spermatophores [ 36 ] . however , t . dalmanni spermatophores are small [ 37 ] and females store few sperm following a single mating ( ~ 140 ; [ 36 ] ) . females are therefore sperm - limited and must copulate repeatedly to attain maximal fertility [ 38 ] . this problem is exacerbated in large , highly fecund females despite being allocated more ejaculate , as they lay a lower proportion of fertilised eggs following a single copulation in comparison with less productive females [ 38 ] . without measures of paternity however , the value of both mate choice and multiple mating can only be inferred indirectly .
with eyes at the tip of their eye stalk , teleopsis dalmanni and members of diopsinae definitely make into the list of most bizarre - looking animals on earth . this group of flies has sparks research interest in the physiology , development and evolutionary mechanism leading to the eye stalks . t . dalmanni is probably the most widely studied particularly in the field of sexual selection leading to sexual dimorphism in this species , where males and females differ in appearance .
the fly , c . dalmanni , is highly promiscuous ( wilkinson et al . , 1998 ) thereby creating ample opportunity for intense postcopulatory sexual selection by sperm competition or selection . comparative studies across stalk - eyed flies reveal that sperm length , spermathecal duct length and ventral receptacle size have coevolved between males and females ( presgraves et al . , 1999 ) . the size and shape of female sperm storage organs have the potential to influence which sperm are used for fertilization . thus , correlated evolution between male and female reproductive traits may be due , in part , to selection on sperm size mediated by the size of the female organs . however , correlated evolution between male and female reproductive traits may also be the result of linkage disequilibrium between pre - and postcopulatory traits . among lines selected for long or short male eye span , wilkinson et al . ( 2005 ) found a strong correlation between eye span and ventral receptacle size due , in part , to x - linked effects . eye span is , therefore , partly indicative of both male and female postcopulatory features . consequently , genetic linkage among x - linked genes that influence eye span , sperm length and ventral receptacle size provides a mechanism by which female mate choice can potentially influence male external morphology and sperm length as well as result in coevolution between sperm length and female sperm storage organ size . furthermore , the genetic linkage between eye span and sperm length allows both pre - and postcopulatory sexual selection to act against the transmission bias caused by the sex - ratio chromosome .
lay on average 4 - 6 eggs per day and may continue laying eggs for many months . the eggs require 1 - 3 days to hatch . larvae stage range from 10 days to several weeks , depending on temperature of environment and food quality . late - instar larvae empty their gut content in a process called gut - purge , and enter pupation state . in food - depriving environment , pupation might occur at smaller larvae size . they typically pupate on emergent vegetation . pupation stage last for almost a week , after which an adult fly will emerge , completing the life cycle of
as yet , there has been no effort to identify within - population quantitative variation in p 2 in stalk - eyed flies . in this paper we investigated intra - specific , within population variation of p 2 in t . dalmanni using controlled mating experiments and microsatellite profiling of progeny . we standardized male mating history , diet and body size in order to limit variation in male reproductive organ size , as we have shown that testes and accessory glands are reduced by mating [ 41 ] and scale positively with diet quality and body size [ rogers et al . in prep ; cotton & pomiankowski unpublished ] . we also chose sires with similar eyespan to each other to limit the potential effects on p 2 of female mate choice based on male ornament size [ 25 , 27 , 28 ] . we show that even in the absence of these factors there is significant variation in sperm precedence in t . dalmanni , with familial p 2 values ranging from zero to one . we discuss the likely causes of this variation .
we know that sperm precedence in c . dalmanni is highly variable and exhibits a trimodal distribution ( corley et al . , 2006 ) . furthermore , when females are double - mated to st and sr males , st males are twice as likely to fertilize offspring ( wilkinson et al . , 2006 ) . the greater sperm length of st males may facilitate fertilization success . in other insect species , longer sperm do not necessarily have a fertilization advantage over shorter sperm ( e . g . , gage and morrow , 2003 ) , but can in species where females mediate fertilization through organs like sperm receptacles ( reviewed in snook , 2005 ) . we do not know the exact mechanisms of sperm competition in c . dalmanni ; there may be a trade - off between producing many small or few long sperm , although very few sperm per ejaculate succeed in getting stored in female spermathecae ( approximately 35 in cyrtodiopsis whitei ; fry and wilkinson , 2004 ) . male and female postcopulatory traits have coevolved across stalk - eyed fly taxa ( presgraves et al . , 1999 ) and are the fastest evolving traits measured among populations within c . dalmanni ( eg amitin and gs wilkinson , unpublished data ) , which are patterns consistent with sperm competition mediated by ventral receptacles . double - mating experiments using flies that differ only in sperm length , but not presence of drive , are needed to determine how sperm length influences sperm competitive ability . by simultaneously allowing males to differ in eye span it may be possible to determine whether pre - and postcopulatory selection operates in concert or independently on male pre - and postcopulatory sexually selected traits .
templeton ( 26 ) proposed that sexual selection is not important in speciation in the hawaiian drosophila because mate choice is stabilizing within species . the sexually selected trait that we examined is not important in behavioral isolation , but the reason is not consistent with templeton’s proposal because we found directional sexual selection through female mate choice . our detection of directional selection is not due to sampling a smaller size range than templeton did because the range of head widths in our study was 2 . 4–3 . 1 mm , nearly the same as the range studied by templeton ( 17 , 26 ) . statistical analyses of selection are essentially correlational ( 25 , 29 ) , and the cause of selection needs to be confirmed with an experimental approach , such as increasing the male eye span through artificial selection ( 30 ) . female preferences for broad heads in distantly related stalk - eyed flies ( diopsidae ) also appear to be directional ; females preferred to perch near males with heads that had been widened experimentally well beyond the natural range of the species ( 31 ) .


kerria lacca is a species of insect in the family kerriidae , the lac insects .

these are in the superfamily coccoidea , the scale insects .

this species is perhaps the most commercially important lac insect , being a main source of lac , a resin which can be refined into shellac and other products .

this insect is native to asia .

kerria lacca

general remarks : takahashi ( 1949 ) regarded this species as a form or subspecies of kerria lacca . varshney ( 1976 ) gave a table of taxonomic characters to distinguish between kerria ( kerria ) lacca lacca , kerria ( kerria ) lacca ambigua and kerria ( kerria ) lacca mysorensis .
kerria ( kerria ) lacca ambigua ( misra ) ; varshney 1984b : 368 . change of combination
kerria lacca ambigua ( misra ) ; varshney 1977 : 43 . change of combination
raising kerria lacca keer entail heavy production costs and is not as difficult as farm work . locals just have to rezone forest land to raise kerria lacca keer and harvest shellac .
people in huoi leng commune tend palm trees to raise kerria lacca keer ( photo : baodienbienphu . com . vn )
studies on the biology of lac insect , laccifer lacca ( kerr ) targ .
huoi leng’s soil and weather are favorable for raising kerria lacca keer , a kind of insect that lives on palm trees and produces a shellac used in fine arts and medicine .
studies on the biology of lac insect , laccifer lacca ( kerr ) targ .
krishan sharma k , ramani r ( 2001 ) parasites effected reduction in fecundity and resin yield of two strains of indian lac insect , kerria lacca . indian j ent 63 : 456–459 .
studies on the biology of lac insect , laccifer lacca ( kerr ) targ . [ 1984 ]
since they began raising kerria lacca keer , h’mong people no longer burn the forests to create terraced fields . bare hills have been recovered with palm trees . kerria lacca keer raising areas have expanded to 300 hectares . traders come to huoi leng to buy shellac at high prices , sometimes as high as 150 usd per kg . hang say dua , chairman of the huoi leng people’s committee , says : “ about 200 h’mong households have become better - off from raising kerria lacca keer . h’mong people now have access to education and mass media and their knowledge has improved . their lives have improved remarkably . ”
genetic diversity in lac resin - secreting insects belonging to kerria spp . , as revealed through issr markers
genetic diversity in lac resin - secreting insects belonging to kerria spp . , as revealed through issr markers - pubag
lac is a natural resin secreted by a tiny insect – kerria lacca ( kerr ) for its own protection . this was an important resin because it was commercially used in various industries such as food , pharmaceuticals , cosmetics , varnishes , sealing wax , lubricants and insulating materials .
ho thi dinh says her family’s average annual income is 2 , 500 usd and their lives are now much better . “ we sell shellac and have enough money to afford to send my children to school and buy a motorbike . last season , i had enough money to buy 3 horses and 5 buffaloes . growing rice earns less money than raising kerria lacca keer . ” dinh said .
the purpose of this study is to clarify the taxonomic status of two species of kerria in china . fresh insects were collected from their host in the locations according the first record in publication . the two species were compared morphologically at the cellular level by studying their karyotypes , rapd reactions were performed , they were sequenced with ef1α genes and hybridization to establish their identifications , and their relationship with other species in genus kerria was analyzed .
citation : chen y , lu z , li q , hoffmann bd , zhang w ( 2014 ) multiple ant species tending lac insect kerria yunnanensis ( hemiptera : kerriidae ) provide asymmetric protection against parasitoids . plos one 9 ( 6 ) : e98975 . urltoken
the technique of random amplified polymorphic dna ( rapd ) was used to study the relationships of 12 populations from 7 species of kerria . the genetic distance and identity among species were generated by popgene32 ( yeh and boyle 1997 ) . the molecular dendrogram was constructed based on nei ' s genetic distance by meg a3 using the upgma method ( kumar et al . 2004 ) .
morphology , cytology , and molecular biology evidence consistently indicated k . yunnanensis and k . ruralis , the two chinese endemic species , had significant differences with the other five species examined in this study . the relationships of the seven species were basically consistent , with a few phylogenetic positions being incomplete . however , with no matter which methods , k . lacca and k . sindica , and k . pusana and k . nepalensis , always clustered together and formed two sister groups indicating a close genetic relationship between them . with the exception of rapd , k . nepalensis and k . pusana always clustered together , indicating a close genetic relationship . k . chinensis was rather special and always stood alone in a separate branch .
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copyright : © 2014 chen 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 .
funding : this research was partly supported from grant 201204602 provided by special fund for forestry research in the public interest and national natural science foundation ( nsf ) grant 31270561 . the funders had no role in study design , data collection and analysis , decision to publish , or preparation of the manuscript .
lac cultivation is unique among most agricultural practices in that it is obliged to be organic , because chemicals that can be used to kill predators and parasitoids of lac insects would also kill the lac insects . many ant species tend k . yunnanensis for honeydew [ 21 ] , and the presence of at least one species in china , crematogaster macaoensis , was recently found to improve key features of fitness of k . yunnanensis , including survival rate , number of females and number of offspring [ 22 ] , presumably by providing protection from predators and parasitoids . therefore ants have a clear role in the management of lac cultivation .
despite the economic importance of this agricultural system , and a clear reliance on ants for lac production , very little is known about lac insect - ant - parasitoid relationships . all other work conducted to date has been limited to the identification and basic biology of parasitoids [ 23 ] , [ 24 ] , [ 25 ] , [ 26 ] , [ 27 ] , and demonstrating the impacts of parasitoids on lac insects [ 22 ] , [ 28 ] , [ 29 ] . here , for the first time , we investigate the role of ants in protecting lac insects from parasitoids within the typical conditions of a lac farming system . specifically we address two hypotheses : 1 ) that different ant species with different abundance levels provide different levels of protection to k . yunnanensis ; and 2 ) that these relationships differ with the different life stages of k . yunnanensis .
the study was carried out on private land ; we confirmed that the owner of the land gave permission to conduct the study on this site .
at experimental commencement in october 2009 , one third of the trees throughout the plantation were inoculated with k . yunnanensis , typical of normal lac farming practices . inoculation involved placing brood lac , containing k . yunnanensis larvae , on branches . the larvae move from the brood lac onto the branches and aggregate .
we selected 174 d . obtusifolia that were eight years old , 2 . 5 m to 2 . 8 m high , with a trunk diameter of 5 cm to 7 cm , and spaced at least 20 m apart . for the ant exclusion treatment , prior to k . yunnanensis inoculation , all ants visiting or living on 60 trees were removed , and a ring barrier of insect glue was applied to the main branch 0 . 5 m above the ground to prevent ant access . all other vegetation touching the tree were also cleared . every second week the insect glue was replaced and the surrounding vegetation was cleared to maintain ant exclusion .
if our two treatments merely varied the abundance of a single ant species we would anticipate having a single relationship between parasitoids and the ants of the two ant attendance treatments . instead , because there are different ant species within the two ant attendance treatments , we would anticipate multiple relationships . this more complicated design reflects the on - ground reality of the farming system and therefore has greater practical application .
except when stated otherwise , analyses were conducted using statistica 11 . not all k . yunnanensis aggregations were parasitized and there was no pattern of lack of parasitism with treatment ( e . g . all non - parasitized aggregations being in the high ant attendance treatment , or increasing lack of parasitism with decreasing ant attendance ) , so we excluded such trees from statistical analyses , leaving 22 , 20 and 21 samples ( february ) and 28 , 29 and 22 samples ( april ) in the exclusion to high ant attendance treatments respectively .
mean ( ±se ) lac crust area sampled in the three treatments and two sample times .
to compensate for the slight non - homogeneity of sample size , we initially standardized the ant and parasitoid data per crust area , however , we found no relationship between ant abundance or parasitoid abundance with crust area . instead it appears that ants merely respond to the presence / absence of crust , and the effects on parasitoids are dependent on the type and number of ants present , irrespective of the size of the crust . additionally , the patterns of parasitoid abundance per crust and their statistical separation were identical when using non - standardized and standardized data . accordingly we used unadjusted abundance data for both ants and parasitoids in all analyses , and acknowledge that sample surface area of crusts between sample times are not fully homogeneous .
non - metric multidimensional scaling was used to compare homogeneity of ant community structure of the low ant attendance treatment between the two sample times , and parasitoid community structure among the three treatments in the two sample times . the association matrices were based on a bray - curtis association of species - level abundance data per crust of ants and parasitoids respectively . differences among categories were tested using analysis of similarity ( anosim ) for the ant data , and permanova for the parasitoid data . all multivariate analyses were conducted using primer 6 .
because parasitoid abundance and species varied so greatly between the two sample times , different statistical tests were used for different levels of analysis . total parasitoid abundance data were compared among treatments and between sample times using 2 - way anova , and tukey ' s hsd test for the post - hoc comparison of means . the abundance data was log ( x + 1 ) transformed so that residuals met parametric test assumptions , with normality and homogeneity of variance being assessed using graphical observations and levene ' s test respectively .
abundance data of individual species could only be assessed for the three most common species , but not always for both sample times . additionally these data could not satisfy assumptions for parametric tests . for consistency of tests , where there were enough data to perform a test , we analysed sample times separately using non - parametric kruskal - wallis anova .
parasitoid species richness per crust varied little and was heavily skewed , so we compared parasitoid species richness among treatments and between sample times using a poisson - log generalized linear model .
within the low ant attendance treatment we found 11 species tending crusts , six in february , and eight in april ( table s1 ) . the two most abundant species were crematogaster ferrari and c . macaoensis , contributing 50 . 8 and 30 . 5 % of individuals respectively . ordination of the species - level ant abundance data showed that the species and their abundances differed significantly between the two sample times ( figure 2 ; anosim global r = 0 . 133 , p = 0 . 005 ) . this is largely due to the february sample having approximately half ( 5 ) the number of ants per crust compared to the april sample ( 10 ) , and the greater presence of the two most common ants , c . ferrari and c . macaoensis in april compared to february ( present on 86 % of crusts vs 65 % respectively ) .
data are for low attendance treatment only , sampled in february ( closed symbols ) and april ( open symbols ) . 2d stress = 0 . 1
a total of 2574 individuals of parasitoids belonging to five species were collected ( table 1 ) . the two most abundant species were tetrastichus purpureus and tachardiaephagus tachardiae , contributing 82 . 7 % and 13 . 2 % of individuals respectively . total parasitoid abundance varied greatly with treatment ( two - way anova ; f = 4 . 4 , p = 0 . 014 ) and sample time ( f = 52 , p < 0 . 0001 ) , with the two factors having a strong interaction effect ( f = 33 , p < 0 . 0001 ) . total parasitoid abundance was lowest in the february sample when k . yunnanensis was in its younger life stage , and interestingly was significantly lower in the ant exclusion treatment ( figure 3 ) . in april , all three treatments had significantly different parasitoid abundances ( figure 3 ) , with abundance being highest in the ant exclusion treatment and the lowest in the high ant attendance treatment .
mean ( ±se ) parasitoid abundance in the three treatments and two sample times .
mean ( ±se ) parasitoid species abundance data within the three ant attendance treatments from the two sample times .
when ants were present , there were strong negative relationships between total parasitoid abundance and ant abundance , with the relationships being dependent upon the ant species composition and abundance ( figure 4 ) . for the single species c . macaoensis , its relationship with parasitoid abundance was consistent between the two sample times . however , the ant assemblages of the low ant attendance treatment that differed between the two sample times ( figure 2 ) displayed distinctly different relationships with parasitoid abundance ( figure 4 ) . the patterns of total parasitoid abundance were clearly driven by the two most abundant parasitoid species ( figure 5a , b ) . no relationship was evident for marietta javensis , the only other species with enough data to analyse ( figure 5c ) .
treatments and sample times are low ( triangles ) and high ( circles ) ant attendance treatments sampled in february ( closed symbols ) and april ( open symbols ) . relationship metrics are : low ant attendance february sample : y = 22 . 627e −0 . 459x , r 2 = 0 . 9257 ; low ant attendance april sample : y = 831 . 1e −0 . 439x , r 2 = 0 . 914 ; high ant attendance : y = 39371x −2 . 837 , r 2 = 0 . 8571 .
treatments and sample times are low ( triangles ) and high ( circles ) ant attendance treatments sampled in february ( closed symbols ) and april ( open symbols ) . relationship metrics are : graph a ) low ant attendance february sample : y = 70277e −0 . 301x , r 2 = 0 . 464 ; low ant attendance april sample : y = 839 . 3e −0 . 454x , r 2 = 0 . 827 ; high ant attendance : y = 2930 . 6x −2 . 079 , r 2 = 0 . 753 ; graph b ) low ant attendance february sample : y = 10 . 316x −1 . 128 , r 2 = 0 . 496 ; high ant attendance : y = 523271x −3 . 87 , r 2 = 0 . 638 .
treatments and sample times are low ( triangles ) and high ( circles ) ant attendance , and ant exclusion ( squares ) , sampled in february ( closed symbols ) and april ( open symbols ) . 2d stress = 0 . 1 .
our results clearly showed strong negative relationships between the abundance of tending ants and parasitoids , with the relationships being dependent upon the ant species present , ant abundance , and the k . yunnanensis developmental stage . greatest parasitism rates occurred at the oldest developmental stage measured , with the rate being least where crematogaster macoensis were present in high abundance .
asymmetric protection from enemies provided to symbionts by different ant species is well documented globally [ 7 ] , [ 10 ] , [ 33 ] , [ 34 ] with greater protection being provided by ants with greater abundance and greater aggression . here , c . macaoensis was the most abundant , and the most aggressive species , and it provided the greatest protection from parasitoids possibly because they build structures over the lac crusts that provide additional protection against parasitoids . so from the perspective of both k . yunnanensis , and the farmer , c . macaoensis is best to protect lac insects within this agricultural system .
more broadly for theoretical ecology , several hypotheses have been offered to explain mutually beneficial interactions between ants and honeydew - producing insects . our results supported the predictable rewards hypothesis [ 39 ] , [ 40 ] . for ants , the honeydew secreted by k . yunnanensis was predictable in space and time , which elicited repeated visits by ant foragers , and incidentally increased the likelihood of encounters with parasitoids without requiring an increase in ant activity or aggressiveness .
parasitoid trap , consisting of a 25 cm long×10 cm diameter polyester mesh ( 1 mm 2 ) , was placed on the lac crust to collect parasitoids .
parasitoid species and abundance captured from certain area of lac crust within the three ant attendance treatments from the two sample times , which were used in the analyses presented in the paper .
we thank the property owner of the lac - producing farm in yunnan province , china for access to the land , and dr . zhu chaodong for specimen identification .
conceived and designed the experiments : yc ql . performed the experiments : zl wz . analyzed the data : yc zl bh . contributed reagents / materials / analysis tools : yc zl . wrote the paper : yc bh .
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( vovworld ) – huoi leng is a disadvantaged mountain commune in muong cha district , dien bien province . in recent years , the commune has strived to fulfill its socio - economic targets and 10 % of its 200 poor households have escaped poverty . the improvement is due to changes in people’s production habits and views .
huoi leng covers an area of 10 , 000 hectares and has a population of 2 , 500 people , mainly of the h’mong ethnic group . they grow rice and corn on hillsides and their productivity is not very high .
local authorities have encouraged people to change animal raising on mountains to centralized husbandry , which ensures better animal health and higher profits . giang chu nu is a villager . “ previously , we farmed on terraced fields and grazed cattle on mountains but we didn’t have techniques and the yield was not high . now we are trained in modern production methods and production has improved . we earn higher incomes and our lives have changed positively . ”
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laccifer ambigua misra 1930 : 163 . type data : india : uttar pradesh , guna , jhansi , on
jheolia
[ a botanical or vernacular name of unknown plant ( kapur , 1958 ) ] . . syntypes , female and first instar , accepted valid name notes : varshney ( 1976 : 30 ) reported that no type material of the species described by misra , was found at banaras hindu university , varanasi , where dr . a . b . misra has been working . illustr .
biology : misra ( 1930 ) reported this species from
jheolia
, a vernacular name that was not verified by kapur ( 1958 ) and by varshney ( 1976 ) .
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wang , y . d . ( chinese academy of forest sciences , beijing ( china ) . inst . of lac research )
the following is a representative barcode sequence , the centroid of all available sequences for this species . no available public dna sequences . download fasta file
wouters , jan , and verhecken , andré ( 1989 ) .
the coccid insect dyes : hplc and computerized diode - array analysis of dyed yarns
. studies in conservation 34 ( 4 ) : 189–200 . doi : 10 . 1179 / sic . 1989 . 34 . 4 . 189 .
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was identified by wang ziqing . both of them were collected in yunnan , china .
were collected in burma . the three species were identified by authors after a discussion with dr . xie yinping , who is an expert of scale insect in shanxi university , china .
microphotographs of chromosomes in different species were taken with a nikon e800 optical system using the air - dried method ( chen et al . 2007 ) and karyotypic parameters were measured by im50 software ( leica ltd . 1992 ) . karyotype analysis was performed according to the standard method ( leven 1964 ; stebbins 1971 ; guo 1972 ) . phylogenetic relationship of lac insects were studied to built the dendrogram clustered using upgma by applying karyotype resemblance - near coefficients ( λ ) and the evolution distance with specific software ( li et al . 2005 ) .
total genomic dna was isolated from whole insect body using a standard proteinase k , phenol / chloroform extraction technique ( marchant 1988 ; tian 1999 ) . ef1α genes were amplified by polymerase chain reaction ( pcr ) using primer pair forward ( 5′ - atgtgagcagtgtggcaatccaa - 3≈ ) and reverse ( 5′ - gaacgtgaacgtggtatcac - 3′ ) ( palumbi 1996 ) .
dna amplifications were carried out in the bio - rad mycycler thermal cycler . amplification cycles were as follows : 95° c for 4 min as initial denaturation step ; 35 cycles of 64° c denaturation for 1 min , 72° c annealing for 2 min , 72° c extension for 7 min , and ended by cooling at 4° c . the resulting sequences were assembled using bioedit version 7 . 0 . 5 . 3 (
all male insects were manually removed from twigs harboring second instar larvae and the remaining females were covered by a synthetic net sleeve ( 80 mesh ) , which protected the insects from attack from parasitoids and predators . when the females in the sleeves developed into adults , they were copulated with males chosen from other species . the female insects not copulating were treated as controls .
adult female : 1 . 04–1 . 9 mm long , 0 . 69–1 . 38 mm wide , globe - like body , dark reddish brown . anal tubercle heavily sclerotized with 0 . 08–0 . 38 mm long and 0 . 06–0 . 36 mm wide , nearly quadrate , apparently two - segmented and harboring 6–13 anal ring setae about 0 . 18–0 . 27 mm long . branchial tube less than 0 . 09 mm high , brachial plate with crater about 0 . 10–0 . 15 mm long , 0 . 08–0 . 14 mm wide , and 0 . 03–0 . 05 mm 2 in the center . dimples in crater are formed by the brachial pores , numbers vary from 8 to 15 . anterior spiracles are situated 0 . 02–0 . 11 mm to brachial plates with 0 . 18–0 . 3 lmm in length and 0 . 10–0 . 15mm in width , inside the keratinization trail is inconspicuous and less than 0 . 22 mm . dorsal spines are found between the brachia and anal tubercle , and have two parts : a stout pedicel about 0 . 01–0 . 11 mm long and 0 . 04–0 . 05 mm wide and a conspicuous scletotized spine averaged 0 . 10–0 . 23 mm in length . perivulvar pore clusters originate circularly near the anal tubercle . mouthparts have a labium about 0 . 31–1 . 40 mm long , 0 . 11–0 . 21 mm wide with inconspicuous segmentation and a pair of post oral lobes 0 . 03–0 . 12 mm wide just behind the mouth .
holotype : ♀ , paratypes , 8 ♀♀ , 4 may 1987 , yunnan , p . r . china ( ou and hong 1990 ) .
biological characteristics : bi - voltine , summer ( may–october ) and winter crops ( october - the next may ) . life history of females and males are list in
distribution : subtropical areas of pu ' er and lincang of yunnan province , p . r . china .
adult female : length on slide 1 . 02–2 . 3 mm and 0 . 58–1 . 27 mm wide , globe - like body , two body color types , i . e . dark reddish brown or yellow . anal tubercle heavily sclerotized , 0 . 06–0 . 36 mm long and 0 . 21–0 . 37mm wide , apparently two - segmented and consist 1–13 anal ring setae about 0 . 17–0 . 31 mm long . brachial tube less than 0 . 09 mm long , brachial plate with a crater about 0 . 08–0 . 13 mm long , 0 . 06–0 . 11 mm wide and 0 . 03–0 . 04 mm 2 in the center . numbers of pores in crater vary from 4 to 11 . anterior spiracles are situated 0 . 02–0 . 11 mm to brachia 0 . 18–0 . 26 mm long and 0 . 10–0 . 15 mm wide , inside the inconspicuous keratinization trail is less than 0 . 12 mm . dorsal spine heavily sclerotized 0 . 10–0 . 23 mm long ; pedicel of dorsal spine 0 . 02–0 . 16 mm long , 0 . 04–0 . 13 mm wide . perivulvar pore clusters circular , present near anal tubercle . mouthparts with a labium about 0 . 21–0 . 82 mm long , 0 . 12–0 . 19 mm wide , with inconspicuous segmentation and a pair of post oral lobes , each 0 . 06–0 . 14 mm wide , behind the mouthparts .
holotype : ♀ , paratypes , 7 ♀♀ , 10 june 1969 , yunnan , p . r . china ( wang et al . 1982 )
biological characteristics : bi - voltine , summer ( march–july ) and winter crops ( august - the next march ) . life history of female and male were list in
, respectively . two body color types , i . e . red and yellow . the ratio of red : yellow is about 12 : 1 .
distribution : tropical and subtropical areas of pu ' er and xishuangbanna of yunnan province , china .
shows the results . these specimens and permanent slides were deposited in research institute of resource insect of china .
phylogenetic hypotheses among seven species of lac insects based on the morphological characters by the mp method ( the numbers above branches are bootstrap values , chen et al . , 2008 ) . the aphid stomaphis japonica ( hemiptera : aphididae ) is the outgroup . high quality figures are available online .
are made of six metacentric ( or sub - metacentric ) and ten telocentric chromosomes . and
were formed with eight metacentric and ten telocentric chromosomes . both differ from other species and have a certain degree of uniqueness . differences of interspecific relationship were reflected in the centromere position of chromosomes . the cluster analysis method of karyotype resemblance - near coefficient indicated
had the highest identity in karyotype ( 0 . 9688 ) and nearest distance in evolution ( 0 . 0317 ) , which showed they were the latest species of the seven grouped in the dendrogram (
dendrogram of seven species of lac insect based on the karyotype resemblance - near coefficients and evolutionary distance . rnc = resemblance - near coefficients . ed = evolutionary distance . high quality figures are available online .
the results of rapd analysis showed the genetic distance inter - species were 0 . 1854–0 . 7917 , in which the average genetic distance among species was 0 . 4430 (
were 0 . 6297 and 0 . 5789 , respectively ; obviously for different categories . the results of upgma showed that the seven species could be divided into two natural groups (
was the earliest diverging member of this group and is placed as the base branch in the group .
dendrogram of lac insect among seven species based on genetic distances using the method of upgma . high quality figures are available online .
( 57 % ) , in which mp tree length = 377 , ci = 0 . 976 , ri = 0 . 813 , and rc = 0 . 793 using the maximum likelihood ( ml ) to build the phylogenetic tree , the gtr + g model was selected as the best model for phylogenetic analysis in accordance with the hlrt test ( - inl = 2913 . 3172 ) . the support value of all branches was in excess of 70 % . bayesian analysis results were consistent with the systematic relationships of mp and ml trees . except the branch of
, which was 92 % , other branches all received high posterior probability ( pp > 95 % ) .
majority - rule consensus tree resulting from bayesian analysis of ef1α gene ( model = gtr + g ) from seven species of lac insects and the aphid stomaphis japonica ( hemiptera : aphididae ) as outgroup . branches represented are based on the maximum likelihood topology . numbers above internodes indicate bayesian posterior probabilities ; numbers below internodes indicate nonparametric bootstrap proportions for the parsimony analysis ( left ) and likelihood analysis ( right ) . thickened branches indicate bayesian posterior probabilities ≥95 % . high quality figures are available online .
( bayes , 99 % ; mp , 100 % ; ml , 100 % ) and the other containing the remain five species ( bayes , 92 % ; mp , 92 % ; ml , 71 % ) . in group 1 ,
formed a sister branch , showing a close relationship , and indicated that they were the most recently evolved species of the seven .
was the earliest diverging member of group 2 , and has a distant relationship with the others .
were the most recently evolved species of the taxon , forming a close sister group with high support value ( bayes , 100 % ; mp , 83 % ; ml , 78 % ) .
produced first filial generation , indicating the close genetic relationship of the two species . however , crossbreeding between
) , which indicated that three populations belong to separate species with the existence of reproductive isolation among them .
should belong to different categories , with obvious differences from other five species on the morphological characters . the results were in accord with the study of morphological characteristics by using scanning electron microscopy , which confirmed lac commercial species in china was a new species with clear differences from
) , both higher than the average value of inter - species , which proved the three individuals should be classified into different species . this was in contrast to the idea that lac production species in china were the same as
the phylogenetic analysis also found that species distributed in similar ecological environments usually clustered indicating a near relationship .
was originally found ( india , pakistan , nepal , bangladesh , and sri lanka ) .
had the closest relationship and were sister taxa with high support values ( bayes , 99 % ; mp , 100 % ; ml , 92 % ) , in the most basal branch of seven species .
were the most recent of the genus , forming a close sister group with high support values ( bayes , 100 % ; mp , 83 % ; ml , 78 % ) . these four more advanced species were mainly distributed in edge of south subtropical and the north tropical regions .
was found in semi - arid and semi - humid area of southern subtropical zone , with 600–1500 m elevation and 18–20° c average annual temperature .
occured in xishuangbanna , yunnan province in the type humid subtropical climate , close to tropical north border , where the average annual temperature was 19–21° c and the annual precipitation was 1200–1700 mm .
distributed in 800–1400 m altitude area of taunggyi , lashio , meimiao , which belonged to subtropical climate with average annual temperature of 19–20° c and 1200–1500 mm average annual rainfall . while
was located at low elevations in south mandalay , about 200 m above sea level , in a tropical monsoon climate with annual average temperature of 23–29° c and an average of 800–1000 mm annual precipitation .
the authors thank dr . lei shi , youqing chen and engineer shoude ye for their kind help in collecting some lac insect specimens . this research work was supported by the national natural sciences foundation of china ( no . 30800105 ) and the national key technology support program ( 2006bad06b07 ) . the authors also offer special thanks to the anonymous reviewers .
paper copies of this article will be deposited in the following libraries . universitaetsbibliothek johann christian senckenberg , frankfurt germany ; national museum of natural history , paris , france ; field museum of natural history , chicago , illinois usa ; university of wisconsin , madison , usa ; university of arizona , tucson , arizona usa ; smithsonian institution libraries , washington d . c . usa ; the linnean society , london , england . the date of publication is given in ‘about the journal’ on the jis website .
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lakra , r . k . ( haryana agricultural univ . , hisar ( india ) . dept . of entomology )
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scientists from icar - iinrg – vaibhav d . lohot and a . mohanasundaram discovered lac insects in the peripheral areas of jawadhu hills , gandhi nagar , cmc and vellore fort on rain tree and pipal tree during a survey . — photo : v . m . maninathan
two scientists from the indian council of agricultural research ( icar ) - indian institute of natural resins and gums ( iinrg ) , ranchi , have discovered lac insect on rain tree and pipal tree in the peripheral areas of jawadhu hills , gandhi nagar in katpadi and vellore fort .
in fact , the resin was a major source of livelihood for tribals in jharkhand , chhattisgarh , odisha and west bengal as it was traditionally used in making jewellery , according to the scientists .
the scientists from icar - iinrg – vaibhav d . lohot ( plant physiology ) and a . mohanasundaram ( entomology ) - came across the lac insect and their host plants during a survey across various parts of vellore district including yelagiri hill , alangayam , tirupattur , vaniyambadi and gudiyatham , and kancheepuram and tiruvannamalai districts , from october 28 to 31 .
“we have conducted surveys across tamil nadu . we had carried out surveys in madurai and theni in 2011 and in salem in 2014 and found the insect on rain tree , known as ‘thoongu moonchi maram’ in tamil , ” mr . mohanasundaram said .
now , the two scientists , said , for the first time , they have found the lac insect and their host plants in the peripheral areas of jawadhu hills that is venkatesapuram , gandhi nagar in katpadi and vellore fort . it was found on rain tree and pipal tree ( ‘arasa maram’ ) , he added . the insect’s host plants were also observed during the survey .
he said the resin had medicinal value and was used in creams for treating cracked heels , tablet coating and also in ayurveda .
lac insect was reported in the state from the 1930s . however , over the years , lac insect cultivation has lost importance in the state , and people were not aware of it now , leading to disappearance of the insect from southern parts of india , the scientists said .
it is here that icar - iinrg has been playing a pivotal role . with india being a leading producer and exporter of lac resin in the world , icar - iinrg that was established in 1924 is exclusively dedicated to lac insect cultivation . the institute has been taking up exploration , collection and conservation of lac insect and host plants throughout the country on a regular basis .
“this exploration activity was mainly taken up to conserve lac insect , its host plant biodiversity from extinction , ” he said .
with this discovery , the scientists will be taking samples of the insect for morphology and molecular study . “the insect is usually found in areas where the temperature is between 36 and 37 degree celsius . we will also study how it exists in a place like vellore where temperature levels are high , ” he added .
this discovery , they say , has brightened the scope of lac cultivation in the state and will provide income source to many people .