the keeled indian mabuya ( eutropis carinata ) , also called many-keeled grass skink or ( ambiguously ) " golden skink " , is a species of skink found in south asia .

eutropis carinata

valter jacinto marked
file : eutropis carinata lankae common skink සුලබ හිකනලා1 . jpg
as trusted on the
eutropis carinata ( schneider , 1801 )
page .
this species has two subspecies : e . carinata carinata ( schneider , 1801 ) e . carinata lankae ( deraniyagala , 1953 ) .
scincus carinatus schneider 1801 : 183 ( part . ) tiliqua carinata — gray 1827 euprepes merremi duméril & bibron 1839 : 671 euprepes merremii — gravenhorst 1851 : 317 euprepes carinatus — gravenhorst 1851 : 339 euprepis carinatus — peters 1864 : 50 euprepes ( tiliqua ) carinatus — blanford 1870 tiliqua carinata — stoliczka 1870 : 169 mabuia carinata — boulenger 1887 : 181 tiliqua rubriventris hardwicke & gray 1829 tiliqua rubriventris — gray 1846 : 430 euprepes sebae duméril & bibron 1839 : 692 ( part . ) euprepes rufescens günther 1864 : 79 ( part . ) mabuya carinata — smith 1935 : 266 mabuya carinata — greer et al . 2000 eutropis carinata — mausfeld et al . 2002 mabuya carinata — murthy 2010 eutropis carinata lankae ( deraniyagala 1953 ) mabuya carinata lankae deraniyagala 1953 mabuya carinata lankae — manthey 1981 mabuya carinata lankae — ziesmann et al . 2007 eutropis carinata lankae — somaweera & somaweera 2009 eutropis carinata lankae — karunarathna & amarasinghe 2011
once in abundance in and around gardens , these skinks ( eutropis carinata or mabuya carinata or scincus carinatus ) are now getting rarer and very hard to find one . urltoken
testis of the lizard mabuya carinata : a light microscopic and ultrastructural seasonal study .
distribution : see map in deepak et al . 2016 : 80 ( fig . 8 ) . habitat . coastal sand dunes , grassland habitats , prosopis juliflora dominated areas , and grassland plains in the southern tamil nadu . sympatry : eutropis carinata , eutropis bibroni , and calotes versicolor .
pratihar , s . 2012 . ecological plasticity of eutropis carinata ( schneider 1801 ) , in west bengal , india . sonoran herpetologist 25 ( 1 ) : 2 - 3 . - get paper here
testis of the lizard mabuya carinata : a light microscopic and ultrastructural seasonal study . - pubmed - ncbi
chandavar , vidya r . and prakash r . naik . ( 2011 ) localization of glucagon and insulin cells and its variation with respect to physiological events in eutropis carinata . : acta herpetologica 6 ( 1 ) : 47 - 57
we report the first record of eutropis innotata ( blanford , 1870 ) based on a specimen collected in 2 . . .
chandavar , vidya r . and prakash r . naik . 2011 . localization of glucagon and insulin cells and its variation with respect to physiological events in eutropis carinata . acta herpetologica 6 ( 1 ) : 47 - 57 - get paper here
karunarathna , d . m . s . s . & a . a . t . amarasinghe ( 2009 ) common skink eutropis carinata ( reptilia : scincidae ) feed on ratnadvipia irradians ( limacoidea : ariophantidae ) . : taprobanica 1 ( 2 ) : 135 - 136
karunarathna , d . m . s . s . & a . a . t . amarasinghe 2009 . common skink eutropis carinata ( reptilia : scincidae ) feed on ratnadvipia irradians ( limacoidea : ariophantidae ) . taprobanica 1 ( 2 ) : 135 - 136 - get paper here
walikanna , t . p . & d . m . s . s . karunarathna 2009 . notes on ex - situ incubation and hatchlings of eutropis carinata ( schneider , 1801 ) ( reptilia : scincidae ) from sri lanka . taprobanica 1 ( 1 ) : 63 - 65
{ author1 , author2 . . . } , ( n . d . ) . eutropis carinata ( schneider , 1801 ) . [ online ] india biodiversity portal , species page : { name of species field } available at : urltoken [ accessed date jul 10 , 2018 ] .
first report of < i > eutropis innovate < / i > ( blanford , 1870 ) ( reptilia : scincidae ) from nallamalai hills , . . .
justification : eutropis carinata has been listed as least concern because it has a widespread distribution across india , bangladesh , the maldives , and sri lanka . no major threats have been reported and this species is not undergoing population declines . further research into the specific habitat preferences of this species should be carried out .
a . v . ramachandran , m . s . swamy and r . v . shah . 1984 . tail regeneration in the scincid lizard , mabuya carinata , related to breeding seasons and thyroid activity . amphibia - reptilia 5 ( 2 ) : 135 - 144 - get paper here
histomorphology and ultrastructure of the testis during breeding and nonbreeding phases of the reproductive cycle of the lizard mabuya carinata are studied . observations of the ultrastructural features of the testis during breeding and nonbreeding phases of the reproductive cycle reveal a prenuptial type of spermatogenesis and a clearcut discontinuous spermatogenic cycle . seminiferous tubules are enlarged and there is active spermatogenesis as shown by the presence of all the stages of spermatogenesis ( spermatogonia to spermatids ) and spermatozoa during the breeding phase ( november ) . during the nonbreeding phase ( april ) only spermatogonia and sertoli cells are seen in the shrunken seminiferous tubules . leydig cells and sertoli cells show distinct changes in the morphological appearance with hypertrophy of the cells in breeding phase and atrophy of the cells in the nonbreeding phase of the reproductive cycle . the present study suggests that sertoli cells and leydig cells functions are synchronous in the lizard m . carinata .
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de silva , r . , milligan , h . t . , wearn , o . r . , wren , s . , zamin , t . , sears , j . , wilson , p . , lewis , s . , lintott , p . & powney , g .
this species inhabits all of india , including the northwest ( r . vyas pers . comm . 2010 ) . it is also found in nepal ( r . vyas pers . comm . ) , bangladesh , the maldives , and sri lanka ( shrestha 2001 ) .
this species is ground - dwelling ( horton 1973 ) , and is thought to inhabit shrubland and forests of the region . it is oviparous ( smith 1935 ) .
there are no known species - specific conservation measures in place for this species . further research into the specific habitat preferences of this species should be carried out . population monitoring is also recommended .
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research & database prepared by snake research organization , ujjain , m . p .
body robust ; lower eyelids scaly ; vertebral scales smooth . brown to olive or bronzy above , uniform or with dark - brown or black spots , or longitudinal streaks along the lateral margins of the scales . sides are darke brown or chestnut , with or without light spots . a light dorso - lateral line starting from above the eye and continued to the base of the tail . lower parts whitish or yellowish . svl : 160mm .
distribution : madhya pradesh : throughout . elsewhere in india : throughout , except north - west india and west bengal . elsewhere outside : sri lanka , myanmar , bangladesh and nepal .
copyright © 2013 - 2014 , snake research organization ujjain & urltoken 26 , kisan bhawan , 1st floor , arera hills bhopal ( m . p . ) - 462011
india ( gujarat , maharashtra , madhya pradesh , kerala , himachal pradesh , tamil nadu , mizoram , etc . ) , nepal , bangladesh , maldives , sri lanka
note : tdwg regions are generated automatically from the text in the distribution field and not in every cases it works well . we are working on it .
synonymy partly after smith 1935 . distribution : possibly in bhutan ( lenz 2012 ) .
al - razi , hassan ; shayer mahmood ibney alam , md . muzammel hossain , amit kumer neogi , and mohammad abdul baki 2017 . four new herpetofaunal records from saint martin’s island , cox’s bazar , bangladesh ircf reptiles & amphibians 24 ( 3 ) : 162–167 - get paper here
ardesana , r . , r . jhala & m . bharad 2018 . a preliminary report on reptiles of khirasara vidi , rajkot district , gujarat , india . reptile rap # 180 . in : zoo’s print 33 ( 2 ) : 17 : 22 - get paper here
bauer , aaron m . 1998 . south asian herpetological specimens of historical note in the zoological museum , berlin . hamadryad 23 ( 2 ) : 133 - 149 [ 1999 ]
bhattarai , santosh ; chiranjibi p . pokheral , baburam lamichhane , , and naresh subedi 2017 . herpetofauna of a ramsar site : beeshazar and associated lakes , chitwan national park , nepal . ircf reptiles & amphibians 24 ( 1 ) : 17 - 29 - get paper here
bhupathy , subramanian & n . sathishkumar 2013 . status of reptiles in meghamalai and its environs , western ghats , tamil nadu , india . journal of threatened taxa 5 ( 15 ) : 4953 - 4961 - get paper here
blackburn , d . g . 1993 . standardized criteria for the recognition of reproductive modes in squamate reptiles . herpetologica 49 ( 1 ) : 118 - 132 - get paper here
botejue , w . madhava s . ; jayantha wattavidanage 2012 . herpetofaunal diversity and distribution in kalugala proposed forest reserve , western province of sri lanka . amphibian & reptile conservation 5 ( 2 ) : 65 - 80 ( e38 ) . - get paper here
boulenger , g . a . 1887 . catalogue of the lizards in the british museum ( nat . hist . ) iii . lacertidae , gerrhosauridae , scincidae , anelytropsidae , dibamidae , chamaeleontidae . london : 575pp . - get paper here
boulenger , george a . 1890 . the fauna of british india , including ceylon and burma . reptilia and batrachia . taylor & francis , london , xviii , 541 pp . - get paper here
dreckmann , f . 1908 . breeding habits of some snakes and lizards . j . bombay nat . hist . soc . 18 : 434 - get paper here
duméril , a . m . c . and g . bibron . 1839 . erpétologie générale on histoire naturelle complète des reptiles . vol . 5 . roret / fain et thunot , paris , 871 pp . - get paper here
ganesh , s . r . ; m . arumugam 2016 . species richness of montane herpetofauna of southern eastern ghats , india : a historical resume and a descriptive checklist russ . j . herpetol . 23 ( 1 ) : 7 - 24
gravenhorst , j . l . c . 1851 . über die im zoologischen museum der universität breslau befindlichen wirtelschleichen ( pseudosaura ) , krüppelfüssler ( brachypoda ) , und einige andere , denselben verwandte reptilien aus den zünften der schleichen und dickzüngler [ euprepes striolatus ] . acta acad . caes . leop . carol . nat . cur . 23 ( 1 ) : 291 - 394 [ 1847 ] - get paper here
gray , j . e . 1846 . descriptions of some new species of indian lizards . ann . mag . nat . hist . ( 1 ) 18 : 429 - 430 - get paper here
greer , a . e . , arnold , c . & arnold , e . n . 2000 . the systematic significance of the number of presacral vertebrae in the scincid lizard genus mabuya . amphibia - reptilia 21 ( 1 ) : 121 - 126 - get paper here
günther , a . 1864 . the reptiles of british india . london ( taylor & francis ) , xxvii + 452 pp . - get paper here
inger , robert f . ; shaffer , h . bradley ; koshy , mammen ; bakde , ramesh 1984 . a report on a collection of amphibians and reptiles from the ponmudi , kerala , south india . j . bombay nat . hist . soc . 81 ( 3 ) : 551 - 570 - get paper here
jayaneththi , hareschandra bandula 2015 . vertebrate fauna of morankanda - mukalana secondary forest patch in sri lanka : a checklist reported from 2004 - 2008 survey . ruhuna journal of science 6 : 21 - 41
kandamby , dharmasri 1997 . herpetological types reposed in the national museum colombo , sri lanka . lyriocephalus 3 ( 1 ) : 31 - 33
karunarathna , suranjan ; d . m . s . and a . a . thasun amarasinghe 2011 . a preliminary survey of the reptile fauna in nilgala forest and its vicinity , monaragala district , sri lanka . taprobanica 3 ( 02 ) : 69 - 76
kästle , w . , rai , k . & schleich , h . h . 2013 . field guide to amphibians and reptiles of nepal . arco - nepal e . v . , 625 pp . - get paper here
lalremsanga , h . t . , l . khawlhring & lalrotluanga 2010 . three additional lizard ( squamata : sauria ) records for mizoram , india . journal of threatened taxa 2 ( 2 ) : 718 - 720
lenz , norbert 2012 . von schmetterlingen und donnerdrachen - natur und kultur in bhutan . karlsruher naturhefte 4 , naturkundemuseum karlsruhe , 124 pp .
manthey , u . 1981 . die echsen des ceylonischen regenwaldes und seiner randgebiete . sauria 3 ( 2 ) : 25 - 35 - get paper here
murthy , t . s . n . 2010 . the reptile fauna of india . b . r . publishing , new delhi , 332 pp .
palot , m . j . 2015 . a checklist of reptiles of kerala , india . journal of threatened taxa 7 ( 13 ) : 8010–8022 - get paper here
pandharikar s . d . ; joshi p . s . & v . t . tantarpale 2015 . diversity of saurian fauna from jalgaon ( jamod ) territory , buldhana district ( m . s . ) , india . reviews of progress 3 ( 25 ) : 1 - 4 - get paper here
peters , w . c . h . 1864 . die eidechsenfamilie der scincoiden , insbesondere über die schneider ' schen , wiegmann ' schen und neue arten des zoologischen museums . monatsber . k . akad . wiss . berlin , 1864 : 44 - 58 - get paper here
saikia , u . ; sharma , d . k . & sharma , r . m . 2007 . checklist of the reptilian fauna of himachal pradesh , india . reptile rap ( 8 ) : 6 - 9 - get paper here
schneider , johann gottlob 1801 . historiae amphibiorum naturalis et literariae . fasciculus secundus continens crocodilos , scincos , chamaesauras , boas . pseudoboas , elapes , angues . amphisbaenas et caecilias . frommanni , jena . 374 pp . - get paper here
shrestha , biraj 2017 . reptile rap : herpetofauna of korak village , northern chitwan , nepal . zoo’s print 32 ( 9 ) : 23 - 30 - get paper here
smith , m . a . 1935 . the fauna of british india , including ceylon and burma . reptiles and amphibia , vol . ii . sauria . taylor and francis , london , 440 pp .
somaweera , r . & somaweera , n . 2009 . lizards of sri lanka : a colour guide with field keys . chimaira , frankfurt , 304 pp .
stoliczka , f . 1870 . observations on some indian and malayan amphibia and reptilia . j . asiat . soc . bengal 39 : 134 - 228 . - get paper here
swan , l . w . , & leviton , a . e . 1962 . the herpetology of nepal : a history , check list , and zoogeographical analysis of the herpetofauna . proc . cal . acad . sci . 32 ( 6 ) ( 4 . s . ) : 103 - 147 . - get paper here
taylor , e . h . 1953 . a review of the lizards of ceylon . univ . kansas sci . bull . 35 ( 12 ) : 1525 - 1585 - get paper here
tsetan , chime & r . ramanibai 2011 . reptilian fauna of agricultural landscapes of chembarambakkam lake , chennai , tamil nadu . reptile rap ( 13 ) : 2 - 8 - get paper here
upadhye , madhav v . ; vinayak v . puranik , prasad dabholkar & ujwala jadhav 2012 . herpetofauna of the vidyanagari campus of the university of mumbai , maharashtra . reptile rap ( 14 ) : 15 - 20 - get paper here
venugopal , p . d . 2010 . an updated and annotated list of indian lizards ( reptilia : sauria ) based on a review of distribution records and checklists of indian reptiles . journal of threatened taxa 2 ( 3 ) : 725 - 738 . - get paper here
vyas , raju 2007 . herptofauna of puma wildlife sanctuary , gujarat , india . reptile rap ( 8 ) : 10 - 15 - get paper here
vyas , raju 2011 . priliminary survey on reptiles of jassore wildlife sanctuary , gujarat state , india . russ . j . herpetol . 18 ( 3 ) : 210 - 214 - get paper here
vyas , raju 2011 . reptilian diversity in and around the shoolpaneshwar wildlife sanctuary , gujarat , india . reptile rap ( 11 ) : 5 - 15 - get paper here
ziesmann , s . ; klaas , p . & janzen , p . 2007 . von skinken und anderen echsen [ sri lankas ] . draco 7 ( 30 ) : 18 - 23 - get paper here
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< a target = ' _ blank ' href = ' urltoken ' > iucn 2011 . iucn red list of threatened species . version 2011 . 2 . exported on 12 january 2012 < / a >
a general description , with any kind of information about the taxon . its main goal is summarize the most relevant or attractive characteristics of this taxon to the general public .
lower eyelid scaly . ear opening has short pointed lobes anteriorly , no postnasal present . supralabials : 7 ; infralabials : 5 - 8 . dorsal scales have 3 , 5 or 7 keels . 30 - 34 scales round midbody ; lamellae under 1st toe : 6 - 10 ; lamellae under 4th toe : 14 - 18 . dorsum brown , bronze or olive with black spots . dorsolaterally dark brown with a light dorsolateral line travelling from eyes to vent . venter white .
describes reproductive physiology and behavior , including mating and life history variables . includes cues , strategies , restraints , rates .
describes average size , max , range ; type of size ( perimeter , length , volume , weight . . . ) .
relations that living organisms have with respect to each other and their natural environment . variables of interest to ecologists include the composition , distribution , amount ( biomass ) , number , and changing states of organisms within and among ecosystems .
general description of the sites where the species is found ( ecosystem , forest , environment or microhabitat ) . includes realm ( e . g terrestrial etc ) and climatic information ( e . g boreal ) ; also includes requirements and tolerances ; horizontal and vertical ( altitudinal ) distribution . also includes information referring to territorial extension of the individual or group in terms of its activities ( feeding , mating , etc . ) , associated mostly to vertebrates .
gujarat , maharashtra , madhya pradesh , kerala , himachal pradesh , tamil nadu , mizoram , etc . .
enumerates geographic entities where the taxon lives . covers ranges , e . g . , a global range , or a narrower one ; may be biogeographical , political or other ( e . g . , managed areas like conservencies ) ; endemism ; native or exotic . does not include altitudinal distribution , which is covered under habitat .
includes abundance information ( population size , density ) and demographics ( e . g . age stratification ) .
describes the likelihood of the species becoming extinct in the present day or in the near future . population size is treated under population biology , and trends in population sizes are treated under trends . however , this is the preferred element if an object includes all of these things and details about conservation listings .
boulenger , g . a . 1890 . fauna of british india . reptilia and batrachia .
satara district of maharashtra state is a part of northern western ghats and deccan plateau biogeog . . .
the construction of a road , directly and indirectly , impacts on the ecosystems where the road is bu . . .
a checklist of reptiles of kerala state is presented , along with their scientific , english and mala . . .
this research provides an updated checklist of vertebrate fauna of the chambal river basin in north . . .
cepf western ghats special series : status of reptiles in meghamalai and its environs , western ghats , t . . .
we update the reptile fauna of meghamalai area , western ghats based on a literature review and a re . . .
a herpetofaunal inventory based on field surveys , literature records and photographic records is pr . . .
taxonomic status of the arboreal skink lizard < i > dasia halianus < / i > ( haly & nevill , 1887 ) in sri lanka . . .
a comparative study of dasia halianus in india and sri lanka is discussed . indian specimens of “d . . . .
we collected three additional species - calotes maria , calotes mystaceus and ophisaurus gracilis wh . . .
an updated and annotated list of indian lizards ( reptilia : sauria ) based on a review of distribution r . . .
over the past two decades many checklists of reptiles of india and adjacent countries have been pub . . .
road kill of animals by highway traffic in the tropical forests of mudumalai tiger reserve , southern i . . .
highways passing through natural reserves have adverse impact on wild animals . we evaluated the roa . . .
habitat suitability , threats and conservation strategies of hump - nosed pit viper < i > hypnale hymnal < / i > . . .
recent studies indicate that most species are best conserved in their natural community , which resu . . .
selection of egg attachment sites by the indian golden gecko < i > calodactylodes aureus < / i > ( beddome , 18 . . .
some geckos lay eggs at communal egg deposition sites with as many as 300 eggs per site . selection . . .
presentation on how to observe and identify common lizards by hari krishnan surendran at the ind . . .
technical report kanagavel , a . and r . pandya 2013 . establishing community conservation reserves in the . . .
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bauer , aaron m . ( 1998 ) south asian herpetological specimens of historical note in the zoological museum , berlin . : hamadryad 23 ( 2 ) : 133 - 149 [ 1999 ]
bhupathy , subramanian & n . sathishkumar ( 2013 ) status of reptiles in meghamalai and its environs , western ghats , tamil nadu , india . : journal of threatened taxa 5 ( 15 ) : 4953 - 4961
blackburn , d . g . ( 1993 ) standardized criteria for the recognition of reproductive modes in squamate reptiles . : herpetologica 49 ( 1 ) : 118 - 132
botejue , w . madhava s . ; jayantha wattavidanage ( 2012 ) herpetofaunal diversity and distribution in kalugala proposed forest reserve , western province of sri lanka . : amphibian & reptile conservation 5 ( 2 ) : 65 - 80 ( e38 ) .
boulenger , g . a . ( 1887 ) catalogue of the lizards in the british museum ( nat . hist . ) iii . lacertidae , gerrhosauridae , scincidae , anelytropsidae , dibamidae , chamaeleontidae . : london : 575pp .
boulenger , george a . ( 1890 ) the fauna of british india , including ceylon and burma . reptilia and batrachia . : taylor & francis , london , xviii , 541 pp .
dreckmann , f . ( 1908 ) breeding habits of some snakes and lizards . : j . bombay nat . hist . soc . 18 : 434
duméril , a . m . c . and g . bibron . ( 1839 ) erpétologie générale on histoire naturelle complète des reptiles . vol . 5 . : roret / fain et thunot , paris , 871 pp .
frank , norman & ramus , erica ( 1995 ) a complete guide to scientific and common names of reptiles and amphibians of the world . : pottsville : n g publishing inc . , 377 pp .
gravenhorst , j . l . c . ( 1851 ) über die im zoologischen museum der universität breslau befindlichen wirtelschleichen ( pseudosaura ) , krüppelfüssler ( brachypoda ) , und einige andere , denselben verwandte reptilien aus den zünften der schleichen und dickzüngler [ euprepes striolatus ] . : acta acad . caes . leop . carol . nat . cur . 23 ( 1 ) : 291 - 394 [ 1847 ]
gray , j . e . ( 1846 ) descriptions of some new species of indian lizards . : ann . mag . nat . hist . ( 1 ) 18 : 429 - 430
greer , a . e . , arnold , c . & arnold , e . n . ( 2000 ) the systematic significance of the number of presacral vertebrae in the scincid lizard genus mabuya . : amphibia - reptilia 21 ( 1 ) : 121 - 126
günther , a . ( 1864 ) the reptiles of british india . : london ( taylor & francis ) , xxvii + 452 pp .
hardwicke & gray ( 1829 ) ill . ind . zool . 2 : 75
inger , robert f . ; shaffer , h . bradley ; koshy , mammen ; bakde , ramesh ( 1984 ) a report on a collection of amphibians and reptiles from the ponmudi , kerala , south india . : j . bombay nat . hist . soc . 81 ( 3 ) : 551 - 570
jayaneththi , hareschandra bandula ( 2015 ) vertebrate fauna of morankanda - mukalana secondary forest patch in sri lanka : a checklist reported from 2004 - 2008 survey . : ruhuna journal of science vol 6 : 21 - 41
kandamby , dharmasri ( 1997 ) herpetological types reposed in the national museum colombo , sri lanka . : lyriocephalus 3 ( 1 ) : 31 - 33
karunarathna , suranjan ; d . m . s . and a . a . thasun amarasinghe ( 2011 ) a preliminary survey of the reptile fauna in nilgala forest and its vicinity , monaragala district , sri lanka . : taprobanica 3 ( 02 ) : 69 - 76
we are extremely grateful to the head of the department of zoology , utkal university , bhubaneswar , india for providing the necessary facilities to carry out this work .
acutia ml ( 1974 ) the hematology of the tropical lizard . iguana iguana linnaeus : ii . seasonal variations . herpetol 30 : 229–303
( reptilia : teiidae ) naturally infected with hemogregarine : confirmation of monocytosis . vet par 171 : 146–150
campbell tw ( 1998 ) interpretation of the reptilian blood profile . exotic pet pract 3 : 33–36
canfield pj , shea gm ( 1988 ) morphological observations on the erythrocytes , leukocytes and thrombocytesof blue tongue lizards ( lacertilia : scincidae , tiliqua ) . anat histol embryol 17 : 328–342
: a bright - field and phase - contrast study . anat histol embryol 25 : 11–14
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de pienaar uv ( 1962 ) hematology of some south african reptiles . witwatersrand univ , press , johannesburg
duguy r ( 1970 ) numbers of blood cells and their variations . in : gans c , parsons ts ( eds ) biology of the reptilia , vol 3 , morphology c . academic , london , pp 93–104
frye fl ( 1991 ) hematology as applied to clinical reptile medicine . in : reptile care . an atlas of diseases and treatment . tfh , neptune city , pp 211–277
harr ke , alleman ar , dennis pm , maxwell lk , lock ba , bennett ra , jacobson er ( 2001 ) morphologic and cytochemical characteristics of blood cells and hematologic and plasma biochemical reference ranges in green iguanas . j am vet med ass 218 : 915–921
hutchison vh , szarski h ( 1965 ) number of erythrocytes in some amphibians and reptiles . copeia 1965 : 371–375
ismail hu , sevinc ms , yildirimhan hs ( 2003 ) erythrocyte size and morphology of some tortoises and turtles from turkey . zool studies 42 : 173–178
lillywhite hb , smits aw ( 1984 ) liability of blood volume in snakes and its relation to activity and hypertension . j exp biol 110 : 267–274
lowry oh , rosebrough nj , farr al , randall rj ( 1951 ) protein measurement with the folin phenol reagent . j biol chem 193 : 265–275
mclean gs , lee ak , withers pc ( 1975 ) haematological adjustments with diurnal changes in body temperature in a lizard and a mouse . comp biochem physiol sla : 241 - 249
murthy tsn ( 2010 ) the reptile fauna of india . b . r . publishing , new delhi , p 332
) in ibadan metropolis , nigeria . vet res 4 ( 3 ) : 77–80
rosenthal hl , pfluke ml , buscaglia s ( 1960 ) a stable iron reagent for determination of cholesterol . j lab clin med 50 : 318
saad ah , elrido r , zada s , badir n ( 1984 ) effect of hydrocortisone on immune system of the lizard . chalcides ocellatus . i . response of lymphoid tissues and cells to in vitro hydrocortisone . dev comp immunol 8 : 121–130
saint girons m ( 1970 ) morphology of the circulating blood cells . in : gans c , parsons ts ( eds ) biology of the reptilia , vol 3 , morphology c . academic , new york , pp 73–91
szarski h , czopek g ( 1984 ) erythrocyte diameter in some amphibians and reptiles . bull acad sci pol ci iiser biol 14 : 443–447
wagner ra , wetzel r ( 1999 ) tissue and plasma enzyme activities in juvenile green iguanas . am j vet res 60 : 201–2003
parida , s . p . , dutta , s . k . & pal , a . comp clin pathol ( 2013 ) 22 : 869 . urltoken
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citation title :
sun skink diversification across the indiansoutheast asian biogeographical interface
.
study name :
sun skink diversification across the indiansoutheast asian biogeographical interface
.
we have kept jott a free peer - reviewed scientific journal to promote conservation . we have not put up a paywall to readers , and we do not charge for publishing . but running a monthly journal costs a lot . while we do have some partners , we still require support to keep the journal alive . if our readers help fund it , our future will be more secure .
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, ifs , inspector general of forests , ministry of environment and forests for funding support and encouragements . smmj expresses gratitude to
ravi singh , sg & ceo , wwf - india , new delhi and ms .
, state director and national coordinator , epcg , wwf - india , ap state office , hyderabad for constant support and encouragement . cs acknowledges csir , new delhi for funding and head , department of zoology ,
and dr . aaron m . bauer , villanova university , usa for helping in identification and constructive suggestions .
( anderson , 1871 ) are known from india to possess , more or less , transparent disc on the lower eye lid .
hills , eastern ghats , andhra pradesh and put on record range extension of this species further south in peninsular india .
) ( image 13 & 13a ) ; 33 scales round the mid body , dorsal scales the largest ; digits long , with feebly keeled lamellae and the fourth toe with 17 lamellae ( image 12 ) .
the specimen measured 46mm in snout - vent and 69mm in tail length ( image 2 & 3 ) .
of the skink was light golden brown , dorsal side without vertebral markings or streaks .
the present specimen was collected from dry leaf litter , and the area had mixed dry deciduous forest with patches of scrubs ( champion & seth 1968 ) .
c in december , and the locality receives about 900 - 1000 mm rainfall annually .
( 1890 ) , smith ( 1935 ) , and sharma ( 2002 ) .
with 2 or 3 strong keels …… . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
with 7 to 11 keels … . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . …… .
project ( bcpp ) india , endangered species project – conservation assessment and management plan ( c . a . m . p . ) workshops .
hills , eastern ghats , andhra pradesh , with eleven new records from the region including ten new records for the state .
fauna of andhra pradesh , part - i . zoological survey of india , calcutta , 1 - 63pp .
hills , eastern ghats , india : an annotated checklist , with remarks on nomenclature , taxonomy , habitat use , adaptive types and biogeography .
copyright ( c ) 2010 k . t . rao , s . m . m . javed , c . srinivasulu
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india ( tamil nadu ) type locality : tuticorin , georeferenced latitude n 8 . 784993° , longitude e 78 . 156341° , thoothukudi , tamil nadu state .
oviparous . breeding males were observed during the month of september and october . hatchlings were recorded during the month of january .
holotype . ncbs - aq054 , adult male , collected by v . deepak and m . kannan on 17 october 2013 . paratypes . ces 141163 , adult male , bnhs 2310 adult female , tuticorin , georeferenced latitude n8 . 784993° , longitude e78 . 156341° , thoothukudi , tamil nadu state , collected by v . deepak and m . kannan on 17 october 2013 .
the species epithet is derived from the regional tamil language for a palm leaf fan , a hand crafted fan made from palm leaf . the dewlap of this species resembles a local palm leaf fan .
deepak , v . ; akshay khandekar , sandeep varma , r . chaitanya 2016 . description of a new species of sitana cuvier , 1829 from southern india . zootaxa 4139 ( 2 ) : 167–182 - get paper here
deepak , v . ; varad b . giri , mohammad asif , sushil kumar dutta , raju vyas , amod m . zambre , harshal bhosale , k . praveen karanth 2016 . systematics and phylogeny of sitana ( reptilia : agamidae ) of peninsular india , with the description of one new genus and five new species contributions to zoology 85 ( 1 ) : 67 - 111 - get paper here
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new records of diplomys labilis ( bangs , 1901 ) ( mammalia , rodentia , . . .
first record of boana maculateralis ( caminer & ron , 2014 ) . . .
first record of the bignose unicornfish , naso vlamingii ( perciformes , . . .
ambidexter symmetricus manning & chace , 1971 ( decapoda , processidae ) : . . .
check list is a peer - reviewed , open access , on - line journal devoted to publishing annotated list of species , notes on geographic distribution of one or a few species , and distribution summary of a taxonomic group . these data are essential for studies on biogeography and provide a baseline for the conservation of biodiversity as a whole . the first step to undertaking effective conservation action is to understand species’ geographic distribution . check list was established to cater to this need by publishing papers on the geographic distribution of species and higher taxonomic groups .
doaj , scopus , zoological abstracts , ebsco host , and index copernicus . member journal of the brazilian association of science editors ( abec ) and of the committee on publication ethics ( cope ) .
hatching of this species at colombo district in sri lanka . in total , 25 eggs with two adults of
the mound . of these skink eggs , 20 were successfully incubated . the incubation period was 30 days from
the date eggs were collected . the average incubation temperature varied from 26–28°c . mean egg length
was 15 . 6 mm and mean egg width 10 . 4 mm . mean hatchling svl was 28 . 5 mm and mean tl is 41 . 3 mm .
the mean weight of hatchlings was ~ 3 g and the mean total length is 69 . 8 mm . after about five days ,
1 , 000 m a . s . l . ( das & de silva , 2005 ) . adults of
© taprobanica nature conservation society , 146 , kendalanda , homagama , sri lanka .
mean tl was 41 . 3 mm ( range 36 . 5–46 . 3 mm ) ( fig .
mean total length was 69 . 8 mm ( range 62 . 7–76 . 5
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body cylindrical , dorsal scales with 5 - 8 keels , ventral scales smooth ; 28 - 30 scales round the body . deep - brown , olive or bronze - brown in colour ; a pair of light or yellow dorso lateral bands starts from above the eye till the base of the tail ; sometimes with black spots on the base of the tail . breeding males have orange colour on the lateral side of the body .
diurnal ; terrestrial ; mostly seen on forest floor , dry forest , rain forest near human dwellings ; under leaf litter , rock boulders .
distribution : madhya pradesh : throughout . elsewhere in india : jharkhand , chhattisgarh , bihar , utter pradesh , west bengal , odisha and assam . elsewhere outside : pakistan , myanmar , thailand , cambodia , vietnam and the malay peninsula
head rather large and distinct ; laterally compressed body with a fairly long tail ; scales on the body pointing backwards and upwards ; two separate spines above tympanum . coloration variable from light - brown or greenish - brown or greyish above , uniform or with more or less instinct dark brown transverse spots or bars upon the back and sides ; or variegated with dark brown ; tail with light and dark annuli . head bright - red , a black patch on the throat . svl : 450mm .
diurnal ; arboreal and most abundant and widespread lizard found in parks , trees , shrubs etc .
distribution : madhya pradesh : throughout . elsewhere in india : throughout . elsewhere outside : pakistan , bangladesh , nepal , sri lanka , bhutan , china , afghanistan , the malay peninsula , hong kong , thailand , myanmar and vietnam .
body flattened , tail widens strikingly behind the narrow base , without denticulate edge but has ventro - lateral row of irregular tubercles ; head oval with a broad snout . dorsum pale - grey at night to olive ; by day with / without 5 dark irregularly undulated transverse bands ; belly light - yellow . svl : 120mm .
nocturnal ; arboreal , but can be seen during day time also . inhabits in the buildings , most agile , and a marked climber
insectivorous and generally feeds on flies , bugs , mole cricket , beetles , termites , spiders and moths .
distribution : madhya pradesh : throughout . elsewhere in india : throughout . elsewhere outside : pakistan , sri lanka , nepal , bangladesh , myanmar , southern china and southeast asia .
body robust ; dorsoventrally flattened ; snout elongated ; nostrils open as an oblique slit , more close to the eye ; neck elongated ; tail long whip like and laterally compressed ; tongue is very long , forked and protrusible . juveniles olive - green or dark - greyish with yellow transverse bands . adults grey or brown with light transverse bands on body and tail ; head with light marking , belly yellowish or brownish . svl : 1800mm .
diurnal ; burrowing habits ; found in scrub forests to evergreen forests . mostly in plantation area , inside caves , termite mounds , burrows , hollow old trees , nallah , under stones and boulders in dense vegetation bordering marshes , ponds , canals and tanks
feeds mainly on insects , small mammals , snakes , lizards and vegetable matter .
distribution : madhya pradesh : throughout . elsewhere india : throughout . elsewhere outside : iran , russia , afghanistan , pakistan , nepal , myanmar , bangladesh and srilanka
body slender with a long tail ; snout rather acute , tympanum present , hind limbs well developed without 5 toe ; scales keeled , femoral pores absent ; tail long and slender . dorsum light / dark brown ; black edged , diamond shaped marks mouth lining dark blue ; belly cream ; males have bluish gular pouch during breeding season . svl : 100mm
diurnal and terrestrial and prefers rocky terrain , scrub jungles and sandy areas . very fast and agile .
distribution : madhya pradesh : balaghat , dindori , jabalpur , seoni , pachmari ( pbr ) , sagar ( ndws ) and indore ( rws ) . elsewhere in india : throughout except at heavy rainfall areas . elsewhere outside : sri lanka and nepal
digits free , moderately dilated and with almost straight transverse lamellae ; 10 - 11 lamellae beneath the first toe and 13 - 15 under the fourth toe .
not accessed . uncommon . no serious threat observedeast concern . uncommon . no serious threats observed .
distribution : madhya pradesh : chhattarpur . elsewhere in india : north - eastern maharashtra , chhattisgarh and odisha .
warning : the ncbi web site requires javascript to function . more . . .
j submicrosc cytol pathol . 2006 jun - sep ; 38 ( 2 - 3 ) : 93 - 102 .
department of studies in zoology , university of mysore , manasagangotri , mysore , india .
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nikaweratiya is located in north - western province about half - way from kandy to anuradhapura . we visited this location because the ' getapolonga ' at the university of peradeniya was reportedly collected there . we were faced by a sparce dry habitat that did not look very promising , but nikaweratiya turned out to be an excellent location for snakes .
scorpions were also in evidence , two species : a broad - clawed scorpionid ( heterometrus gravimanus ) and a narrow - clawed buthid ( buthoscorpio sarasinorum ) .
but it was snakes that were most abundant , we found two buff - striped keelbacks ( amphiesma stolata ) and i was called to a large and defensive long - nosed vinesnake ( ahaetulla nasuta ) in a hedge .
medically important snakes were also common . first capture was a sri lankan cobra ( naja naja ) caught on the dried our paddi - field .
next i was called to a small snake in a pile of palm - frond brush but in the process of searching it i lost my balance and fell into the dead fronds . fortunately i was not bitten because when i had moved the remained of the palm fronds i caught five humpnose pitvipers ( hypnale hypnale ) .
after a successful day at nikaweratiya we drove the 110 km to anuradhapura , slowly road - crusing for snakes and arriving well into the night . it was an interesting and productive drive .
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nausinoe pueritia is a moth in the crambidae family .

it was described by cramer in 1780 .

it is found in india , sri lanka , burma , malaysia , thailand and australia , where it has been recorded from the northern territory , queensland and new south wales .

the wingspan is about 30 mm .

adults are pale brown with complex white markings edged with dark brown on the wings .

the larvae feed on the leaves of abutilon species .

they live within a rolled leaf of their host plant .

the larvae are green and reach a length of about 20 mm .

nausinoe pueritia

shaffer ( pers . comm . ) treats nausinoe in the crambidae : spilomelinae .
{ author1 , author2 . . . } , ( n . d . ) . nausinoe pueritia cramer , 1780 . [ online ] india biodiversity portal , species page : { name of species field } available at : urltoken [ accessed date jul 10 , 2018 ] .
type - species : phalaena pueritia cramer , 1780 . uitl . kapellen 3 : 128 , pl . 264 , fig . f ( not fig . e as stated in text ) . [ bhl ]
( but included by hübner and cited by moore as neptis cramer ; see below ) . hübner , moore and several other authors have been misled by the incorrect figure references cited by cramer for two species that he described in ( 1780 ) ibidem 3 : 128 , and figured on plate 264 . phalaena pueritia ( pyralidae ) , described as ' phal . pyralis ' , is represented by figure f , not by figure e , as cited in cramer ' s text . phalaena neptis ( noctuidae ) , described as ' phal . noct . ' is represented by figure e , not by figure f , as stated in cramer ' s text . because of the transposition of the figure references and the consequent linking of the name neptis with the pyralid illustration phalaena neptis cramer , ( 1780 ) , has been incorrectly used in the pyralidae . the wrapper of part 22 of cramer ' s work , which included phalaena pueritia and phalaena neptis , is dated 1779 but the i . c . z . n . , 1958 , opin . decl . int . commn zool . nom . 19 ( opinion 516 ) : 42 , has shown that the part could not have been published before 1780 . the two names are there .
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urn : lsid : biodiversity . org . au : afd . taxon : 43ef1071 - e846 - 474d - b03e - 31864b241821
urn : lsid : biodiversity . org . au : afd . taxon : 49ccd0a8 - 2848 - 4b5e - ab06 - 46a804fe2cc2
urn : lsid : biodiversity . org . au : afd . taxon : 5571e767 - 0513 - 4d14 - b4ce - abaf0587cc05
urn : lsid : biodiversity . org . au : afd . taxon : 65360df2 - cfbf - 4b18 - 862b - 5f71105da9b3
urn : lsid : biodiversity . org . au : afd . taxon : 20f03500 - f30c - 4a1d - be7d - 9bb6976f9a88
urn : lsid : biodiversity . org . au : afd . name : 330224
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type - species designation : by subsequent designation by moore , 1886 . lepid . ceylon 3 : 309 . [ ( but included by hübner and cited by moore as neptis cramer ) ] [ bhl ]
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acyrthosiphon pisum , commonly known as the pea aphid ( and colloquially known as the green dolphin , pea louse , and clover louse ) , is a sap-sucking insect in the aphididae family .

it feeds on several species of legumes ( plant family fabaceae ) worldwide , including forage crops , such as pea , clover , alfalfa , and broad bean , and ranks among the aphid species of major agronomical importance .

the pea aphid is a model organism for biological study whose genome has been sequenced and annotated .

acyrthosiphon pisum

immunity and other defenses in pea aphids , acyrthosiphon pisum . - pubmed - ncbi
amino acid relations of the pea aphid acyrthosiphon pisum and its symbiotic bacterium buchnera aphidicola .
c . michael hogan changed the thumbnail image of
image of acyrthosiphon pisum
.
example 2 : the metabolic networks of buchnera aphidicola and acyrthosiphon pisum ( pea aphid ) .
wikipedia . acyrthosiphon pisum . available at : < urltoken > . access on june 10 , 2015 .
international aphid genomics consortium ( 2010 ) . genome sequence of the pea aphid acyrthosiphon pisum . plos biol 8 : e1000313 .
international aphid genomics consortium . ( 2010 ) . genome sequence of the pea aphid acyrthosiphon pisum . plos biol 8 : e1000313 .
host life stage - and temperature - dependent density of the symbiont buchnera aphidicola in a subtropical pea aphid ( acyrthosiphon pisum ) population . j
bommarco r , 1991 . action level threshold and economic injury level for the pea aphid , acyrthosiphon pisum ( homoptera : aphididae ) , on field peas , pisum sativum . vaxtskyddsnotiser , 55 ( 4 ) : 114 - 119
citation : the international aphid genomics consortium ( 2010 ) genome sequence of the pea aphid acyrthosiphon pisum . plos biol 8 ( 2 ) : e1000313 . urltoken
partitioning of symbiotic bacteria between generations of an insect : a quantitative study of a buchnera sp . in the pea aphid ( acyrthosiphon pisum reared at different temperatures
partitioning of symbiotic bacteria between generations of an insect : a quantitative study of a buchnera sp . in the pea aphid ( acyrthosiphon pisum ) reared at different temperatures
studies of the pea aphid , acyrthosiphon pisum harris ( hom . , aphididae ) , with special reference to the differences in the biology of the green and red forms
, a facultative symbiont of the pea aphid acyrthosiphon pisum : its phenotypic effect and interaction with coexisting symbiont . microbiol : appl . environ ; 2014 . doi :
anuj bhatnagar , 1996 . efficacy and economics of some insecticides and a neem formulation on incidence of pea aphid ( acyrthosiphon pisum ) on pea , pisum sativum . annals of plant protection sciences , 4 ( 2 ) : 131 - 133 ; 2 ref .
aphidbase is a reference information system providing genomic resources for the study of aphids . it hosts several reference aphid genomes , including the first aphid genome that was sequenced : acyrthosiphon pisum .
phylogenetic analysis of acyrthosiphon pisum aaap family transporters bayesian phylogenetic tree of aaap transporters ( t . c # 2 . a . 18 ) from a . pisum and other taxa , for details see legend of figure 1 . acyrthosiphon pisum , acypi007408 and drosophila , cg31668 solute : sodium symporter family ( t . c # 2 . a . 21 ) transporters were used to root the tree . acyrthosiphon pisum transporters with a similar genomic location are boxed in red and located as follows : ( 1 ) scaffold 345 , accession gl349965 ; ( 2 ) scaffold 385 , accession gl350005 ; and ( 3 ) scaffold 32 , accession gl349652 .
ranked expression levels of aaap transporters ( t . c # 2 . a . 18 ) in acyrthosiphon pisum bacteriocytes , shown relative to gapdh ( acypi009769 ) expression , gapdh = 1 .
cagán l ' ; barta m , 2001 . seasonal dynamics and entomophthoralean infection of the pea aphid , acyrthosiphon pisum harris . plant protection science , 37 ( 1 ) : 17 - 24 .
smith mah ; mackay pa , 1989 . genetic variation in male alary dimorphism in populations of pea aphid , acyrthosiphon pisum . entomologia experimentalis et applicata , 51 ( 2 ) : 125 - 132
c . michael hogan marked the classification from
species 2000 & itis catalogue of life : april 2013
as preferred for
acyrthosiphon pisum ( harris , m . , 1776 )
.
ali k ; louw svan den m ; swart wj , 2005 . components and mechanisms of resistance in selected field pea pisum sativum lines to the pea aphid acyrthosiphon pisum ( homoptera : aphididae ) . international journal of tropical insect science , 25 ( 2 ) : 114 - 121 .
olbrycht t , 1999 . the occurrence of pea aphid ( acyrthosiphon pisum harris ) and its parasitoids on lens ( lens esculata much . ) grown in the province of rzeszow . ( wystepowanie mszycy grochowej ( acyrthosiphon pisum harris ) i zwiazanych z nia parazytoidow na soczewicy ( lens esculata much . ) w okolicach rzeszowa . ) progress in plant protection , 39 ( 2 ) : 416 - 418 .
senoo n ; ochiai y ; nakashima y , 2002 . seasonal abundance of primary parasitoids and hyperparasitoids associated with acyrthosiphon pisum ( harris ) and acyrthosiphon kondoi shinji ( homoptera : aphididae ) on alfalfa . japanese journal of applied entomology and zoology , 46 ( 2 ) : 96 - 98 ; 19 ref .
douglas , a . e . 1996 . reproductive failure and the free amino acid pools in pea aphids ( acyrthosiphon pisum ) lacking symbiotic bacteria . j . insect physiol . 42 : 247 - 255 .
the international aphid genomics consortium , ( 2010 ) . genome sequence of the pea aphid acyrthosiphon pisum . plos biol ( 2 ) : e1000313 . doi : 10 . 1371 / journal . pbio . 1000313
douglas ae , françois clmj , minto lb . facultative ‘secondary’ bacterial symbionts and the nutrition of the pea aphid , acyrthosiphon pisum . physiol entomol . 2006 ; 31 ( 3 ) : 262–9 . doi :
wale m , 2015 . acyrthosiphon pisum ( harris ) pushes cool - season food legumes to extinction in ethiopia . in : pisum sativum : cultivation , functional properties and health benefits [ ed . by becket , s . ] . new york , usa : nova science publishers , 91 - 113 .
mandrioli m , bizzaro d , giusti m , manicardi g . c , bianchi u ( 1999 ) the role of rdna genes in x chromosome association in the aphid acyrthosiphon pisum . genome 42 : 381–386 .
global expression analysis of a . pisum amino acid transporters within a phylogenetic framework facilitates initiation of functional assignment
ekbom b , 1994 . arthropod predators of the pea aphid , acyrthosiphon pisum harr . ( hom . , aphididae ) in peas ( pisum sativum l . ) , clover ( trifolium pratense l . ) and alfalfa ( medicago sativa l . ) . journal of applied entomology , 117 ( 5 ) : 469 - 476
hinz b , 1991 . assessment of damage by the pea aphid , acyrthosiphon pisum , in peas of different consumption type ( homoptera : aphidinea : aphididae ) . entomologia generalis , 16 ( 2 ) : 139 - 145
last time ago we presented a model organism , the red flour beetle , and this week i’m introducing one more . its name is acyrthosiphon pisum , commonly known as pea aphid or sometimes pea louse or green dolphin .
seidenglanz m ; hunady i ; poslusná j ; løes ak , 2011 . influence of intercropping with spring cereals on the occurrence of pea aphids ( acyrthosiphon pisum harris , 1776 ) and their natural enemies in field pea ( pisum sativum l . ) . plant protection science , 47 ( 1 ) : 25 - 36 . urltoken
digilio mc , 1995 . rapid identification of the nymphal stages of acyrthosiphon pisum ( harris ) ( homoptera aphidoidea ) . bollettino di zoologia agraria e di bachicoltura , 27 ( 1 ) : 111 - 116 ; 11 ref .
charles h . , h . ishikawa . 1999 . physical and genetic map of the genome of buchnera , the primary endosymbiont of the pea aphid acyrthosiphon pisum . j . mol . evol . 48 : 142 - 150 .
wilkinson , t . l . and a . e . douglas . 1996 . the impact of aposymbiosis on amino acid metabolism of pea aphids ( acyrthosiphon pisum ) . entomol . exp . appl . 80 : 279 - 282
mondor eb ; baird ds ; slessor kn ; roitberg bd , 2000 . ontogeny of alarm pheromone secretion in pea aphid , acyrthosiphon pisum . journal of chemical ecology , 26 ( 12 ) : 2875 - 2882 ; 27 ref .
wilkinson , t . l . and a . e . douglas . 1995 . why aphids ( acyrthosiphon pisum ) lacking symbiotic bacteria have elevated levels of the amino acid glutamine . j . insect physio . 41 : 921 - 927
of those aphid species , baker ( 2015 ) lists 17 as occurring in britain : acyrthosiphon loti , acyrthosiphon pisum , aphis craccivora , aphis fabae , aphis gossypii , aphis medicaginis , aulacorthum solani , brachycaudus helichrysi , macrosiphum euphorbiae , megoura viciae , myzus ascalonicus , myzus ornatus , myzus persicae , nearctaphis bakeri , smynthurodes betae , therioaphis riehmi and therioaphis trifolii .
sasaki t . & ishikawa h . 1995 : production of essential amino acids from glutamate by mycetocyte symbiont of the pea aphid acyrthosiphon pisum . - j . insect physiol . 41 : 41 - 46 go to original source . . .
number of raw sequences , selected ests , sizes , contigs formed , and redundancy in a . pisum est database
ximenez - embuna , m . g . et al . ( 2014 ) . seasonal , spatial and diel partitioning of acyrthosiphon pisum ( hemiptera : aphididae ) predators and predation in alfalfa fields . biological control 69 , 1 - 7 . abstract
charles h ; ishikawa h , 1999 . physical and genetic map of the genome of buchnera , the primary endosymbiont of the pea aphid acyrthosiphon pisum . journal of molecular evolution , 48 ( 2 ) : 142 - 150 ; 40 ref .
derron jo ; goy g , 1993 . le puceron du pois ( acyrthosiphon pisum harris ) : biologie , nuisibilite et moyens de lutte sur le pois proteagineux . revue suisse d ' agriculture , 25 ( 1 ) : 43 - 47 .
brisson j . a , davis g . k , stern d . l ( 2007 ) common genome - wide patterns of transcript accumulation underlying the wing polyphenism and polymorphism in the pea aphid ( acyrthosiphon pisum ) . evol dev 9 : 338–346 .
boyar dm , 2005 . development of monitoring of the dynamics of numbers and structure of populations of pea aphid ( acyrthosiphon pisum harris ) . vestsi¯ natsyyanal ' nai akademi¯i¯ navuk belarusi¯ . seryya bi¯yalagi¯chnykh navuk , no . 2 : 55 - 60 .
chakraborty a ; dutta sk , 1998 . development and reproduction of pea aphid , acyrthosiphon pisum ( harris ) on pea . journal of the agricultural science society of north east india , 11 ( 2 ) : 162 - 166 ; 5 ref .
lykouressis dp ; polatsidis cp , 1990 . seasonal abundance of acyrthosiphon pisum ( harris ) ( homoptera : aphididae ) and therioaphis trifolii ( monell ) ( homoptera : callaphididae ) on lucerne in central greece . entomologia hellenica , 8 : 41 - 46
montlor c . b . , maxmen a . & purcell a . h . 2002 : facultative bacterial endosymbionts benefit pea aphids acyrthosiphon pisum under heat stress . - ecol . entomol . 27 : 189 - 195 go to original source . . .
mclean ahc ; ferrari j ; godfray hcj , 2009 . effects of the maternal and pre - adult host plant on adult performance and preference in the pea aphid , acyrthosiphon pisum . ecological entomology , 34 ( 3 ) : 330 - 338 . urltoken
morgan d ; walters kfa ; aegerter jn , 2001 . effect of temperature and cultivar on pea aphid , acyrthosiphon pisum ( hemiptera : aphididae ) life history . bulletin of entomological research , 91 ( 1 ) : 47 - 52 ; 27 ref .
fukatsu , t . , n . nikoh , r . kawai and r . koga . the secondary endosymbiotic bacterium of the pea aphid acyrthosiphon pisum ( insecta : homoptera ) . appl . environ . microbiol . 2000 . 66 : 2748 - 2758 .
wilkinson , t . l . and a . e . douglas . 1998 . host cell allometry and regulation of the symbiosis between pea aphids , acyrthosiphon pisum , and bacteria , buchnera . j . insect . physiol . 44 : 629 - 635 .
andarge a ; westhuizen mcvan der , 2004 . mechanisms of resistance of lentil lens culinaris medikus , genotypes to the pea aphid acyrthosiphon pisum harris ( hemiptera : aphididae ) . international journal of tropical insect science , 24 ( 3 ) : 249 - 254 .
nunes mv ; hardie j , 1996 . differential photoperiodic responses in genetically identical winged and wingless pea aphids , acyrthosiphon pisum , and the effect of day length on wing development . physiological entomology , 21 ( 4 ) : 339 - 343 ; 12 ref .
wale m ; jembere b ; seyoum e , 2000 . biology of the pea aphid , acyrthosiphon pisum ( harris ) ( homoptera : aphididae ) on cool - season legumes . insect science and its application , 20 ( 3 ) : 171 - 180 .
montllor cb , maxmen a , purcell ah : facultative bacterial endosymbionts benefit pea aphids acyrthosiphon pisum under heat stress . ecological entomology . 2000 , 27 : 189 - 195 . 10 . 1046 / j . 1365 - 2311 . 2002 . 00393 . x .
charles h . & ishikawa h . 1999 : physical and genetic map of the genome of buchnera , the primary endosymbiont of the pea aphid acyrthosiphon pisum . - j . mol . evol . 48 : 142 - 150 go to original source . . .
douglas a . e . & prosser w . a . 1992 : synthesis of an essential amino acid tryptophan in the pea aphid ( acyrthosiphon pisum ) symbiosis . - j . insect physiol . 38 : 565 - 568 go to original source . . .
the bcm - hgsc sequenced the pea aphid ( acyrthosiphon pisum ) . the pea aphid displays both sexual and asexual reproduction , and is a model for bacterial endosymbiosis and developmental plasticity . it is also an important agricultural pest quickly gaining resistance to multiple insecticides .
honda k , 1989 . seasonal changes in population density of the pea aphid acyrthosiphon pisum harris in pea fields and environmental conditions for the emergence of alate virginoparp . annual report of the society of plant protection of north japan , no . 40 : 110 - 111
mcvean rik ; dixon afg , 2002 . the host plant range of the pea aphid subspecies acyrthosiphon pisum ssp . destructor ( johnson ) ( hom . , aphididae ) . journal of applied entomology , 126 ( 6 ) : 281 - 286 ; 36 ref .
stary p , 1968 . impact of an indigenous parasite , aphidius ervi hal . , on pea aphid , acyrthosiphon pisum ( harris ) populations on alfalfa in czechoslovakia . boll . lab . ent . agr . ` filippo silvestri ' , 26 : 293 - 313 .
fukatsu t . , tsuchida t . , nikoh n . & koga r . 2001 : spiroplasma symbiont of the pea aphid acyrthosiphon pisum ( insecta : homoptera ) . - appl . environ . microbiol . 67 : 1284 - 1291 go to original source . . .
bizzaro d ; mandrioli m ; zanotti m ; giusti m ; manicardi gc , 2000 . chromosome analysis and molecular characterization of highly repeated dnas in the aphid acyrthosiphon pisum ( aphididae , hemiptera ) . genetica , 108 ( 2 ) : 197 - 202 ; 28 ref .
wale m , 2004 . relative efficacy of some botanicals , detergent and kerosene in controlling the pea aphid acyrthosiphon pisum ( harris ) ( hemiptera : aphididae ) on grass pea lathyrus sativus . international journal of tropical insect science , 24 ( 2 ) : 143 - 149 .
4 . wilkinson t . l . , and douglas a . e . , why pea aphids ( acyrthosiphon pisum ) lacking symbiotic bacteria have elevated levels of the amino acid glutamine . j insect physiol , 1995 . 41 ( 11 ) : p . 921 - 7 .
giles kl ; obrycki jj ; degooyer ta , 1994 . prevalence of predators associated with acyrthosiphon pisum ( homoptera : aphididae ) and hypera postica gyllenhal ( coleoptera : curculionidae ) during growth of the first crop of alfalfa . biological control , 4 ( 2 ) : 170 - 177
wale m ; jembere b ; seyoum e , 2003 . effect of aphid density on reproduction and survival of the pea aphid acyrthosiphon pisum ( harris ) ( homoptera : aphididae ) on cool - season food legumes . pest management journal of ethiopia , 5 : 12 - 19 .
wale m ; jembere b ; seyoum e , 2003 . occurrence of the pea aphid acyrthosiphon pisum ( harris ) ( homoptera : aphididae ) on wild leguminous plants in west gojam , ethiopia . sinet : ethiopian journal of science , 26 ( 1 ) : 83 - 87 .
chen d . q . , campbell b . c . & purcell a . h . 1996 : a new rickettsia from a herbivorous insect , the pea aphid acyrthosiphon pisum ( harris ) . - curr . microbiol . 33 : 123 - 128 go to original source . . .
fukatsu t . , nikoh n . , kawai r . & koga r . 2000 : the secondary endosymbiotic bacterium of the pea aphid , acyrthosiphon pisum ( insecta : homoptera ) . - appl . environ . microbiol . 66 : 2748 - 2758 go to original source . . .
wilkinson t . l . , fukatsu t . & ishikawa h . 2003 : transmission of symbiotic bacteria buchnera to parthenogenetic embryos in the aphid acyrthosiphon pisum ( hemiptera : aphidoidea ) . - arthr . struct . dev . 32 : 241 - 245 go to original source . . .
tsuchida t . , koga r . , meng x . - y . , matsumoto t . & fukatsu t . 2005 : characterization of a facultative endosymbiotic bacterium of the pea aphid acyrthosiphon pisum . - microbiol . ecol . 49 : 126 - 133 go to original source . . .
chen , d . - q . ; montlor , c . b . 2000 . fitness effects of two facultative endosymbiotic bacteria on the pea aphid , acyrthosiphon pisum , and the blue alfafa aphid , a . kondoi . entomologia experimentalis et applicata , 95 ( 3 ) : 315 - 323 .
humphreys , n . j . and a . e . douglas . 1997 . the partitioning of symbiotic bacteria between generations of an insect : a quantitative study of buchnera in the pea aphid ( acyrthosiphon pisum ) reared at different temperatures . appl . environ . microbiol . 63 : 3294 - 3296
the pea aphid is a model organism whose genome has been sequenced and annotated . acyrthosiphon pisum is a rather large aphid whose body can reach 4mm in adults . pea aphids also show hereditary body color variations of green or red / pink . the green morphs are generally more frequent in natural populations .
tsuchida t . , koga r . , shibao h . , matsumoto t . & fukatsu t . 2002 : diversity and geographic distribution of secondary endosymbiotic bacteria in natural populations of the pea aphid acyrthosiphon pisum . - mol . ecol . 11 : 2123 - 2135 go to original source . . .
eastop vf , 1971 . keys for the identification of acyrthosiphon . bulletin of the british museum of natural history ( entomology ) , 26 : 1 - 115 .
additional data file 2 : a list of the 921 perfect microsatellite motifs found in the 12 , 082 a . pisum contigs ( xls 130 kb )
miura t , braendle c , shingleton a , sisk g , kambhampati s , stern dl : a comparison of parthenogenetic and sexual embryogenesis of the pea aphid acyrthosiphon pisum ( hemiptera : aphidoidea ) . journal of experimental zoology . 2003 , 295b : 59 - 81 . 10 . 1002 / jez . b . 3 .
szwejda j , 1992 . atraycynosc odmian i zwalzanie szkodnikow wystepujacych na grochu ( pisum sativum l . ) . biuletyn warzywniczy , 39 : 129 - 137 .
additional data file 1 : a list of the 25 most abundant a . pisum unique transcripts from the 12 , 082 collection . ( xls 21 kb )
stary p ; gonzalez d , 1978 . parasitoid spectrum of acyrthosiphon - aphids in central asia ( hymenoptera : aphidiidae ) . entomologica scandinavica , 9 ( 2 ) : 140 - 145
additional data file 3 : a table giving the description by cdna libraries of the 135 contigs specific to a given a . pisum tissue ( xls 175 kb )
model outline for branch analyses of acyrthosiphon pisum apc slimfast expansion molecular evolution . strategy used to test for heterogeneous rates of evolution along branches in a . pisum - specific apc family ( t . c # 2 . a . 3 ) slimfast expansion . ( a ) bayesian phylogeny for a . pisum - specific apc slimfast expansion is shown , including insect orthologs and two outgroup genes ( see fig . 1 ) . each branch is assigned a ω category as indicated . ( b ) description of the models of molecular evolution tested , where model names indicate the number of ω categories assumed ( e . g . , one ratio assumes one ω for all branches ) .
the set of a . pisum contigs orthologous to d . melanogaster sequences have been compared to the whole set of d . melanogaster genes using flybase gene ontology terms . the last column indicates the p value of the hypergeometric test . en , enhanced and d , depleted in a . pisum transcripts . / , no bias .
aphids are the leading pests in agricultural crops . a large - scale sequencing of 40 , 904 ests from the pea aphid acyrthosiphon pisum was carried out to define a catalog of 12 , 082 unique transcripts . a strong at bias was found , indicating a compositional shift between drosophila melanogaster and a . pisum . an in silico profiling analysis characterized 135 transcripts specific to pea - aphid tissues ( relating to bacteriocytes and parthenogenetic embryos ) . this project is the first to address the genetics of the hemiptera and of a hemimetabolous insect .
miura t . , braendle c . , shingleton a . , sisk g . , kambhampati s . & stern d . l . 2003 : a comparison of parthenogenetic and sexual embryogenesis of the pea aphid acyrthosiphon pisum ( hemiptera : aphidoidea ) . - j . exp . zool . ( b ) 295 : 59 - 81 go to original source . . .
the bcm - hgsc would like to thank everyone in the international aphid genome consortium , and others involved in the analysis and annotation of this genome that led to such an amazing set of publications . the genome and its analysis was published in the february issue of plos biology
genome sequence of the pea aphid
acyrthosiphon pisum by the international aphid genome consortium .
“the question is whether the production of nutrients changes with supply and demand and if so , how it happens , ” says wilson . investigators examining the relationship between pea aphids ( acyrthosiphon pisum ) and the bacteria buchnera aphidicola have now learned how aphids’ cells tightly control the metabolic activity of their bacterial endosymbionts . another team of investigators has discovered genetic arrangements that support this symbiosis .
strikingly , compared with other insects with fully sequenced genomes , the a . pisum genome contains extensive gene family duplications ( international aphid genomics consortium 2010 ) . phylogenomic analysis of a . pisum identified 2 , 459 gene families that possess a . pisum lineage - specific duplications ( international aphid genomics consortium 2010 ) , most of which are small to moderate in size ( < 10 paralogs ) , but 9 % of which represent large a . pisum - specific expansions containing > 10 closely related duplicates ( huerta - cepas et al . 2010 ) . to date , detailed analyses have been made of large gene family expansions in a . pisum odorant and gustatory receptors ( smadja et al . 2009 ) , sugar transporters ( price et al . 2010 ) , and proteases ( rispe et al . 2008 ) . we now report in detail on the large expansion of aats belonging to the apc transporter superfamily .
genomic organization of acyrthosiphon pisum apc and aaap transporters . genomic organization of a . pisum duplicated aats belonging to ( a ) apc transporter family ( t . c # 2 . a . 3 ) slimfast expansion and ( b ) aaap transporter family ( t . c # 2 . a . 18 ) from expansion ( 1 ) , ( 2 ) , and ( 3 ) . full - length coding sequences for each of the transporter genes ( as indicated by acypi identifier ) are shown mapped to a . pisum genomic scaffolds ( release acyr 2 . 0 ) . regions of each genomic scaffold containing the aats are drawn to scale ( red line ) , and the location of each transporter is depicted as a shaded block with an arrow indicating gene orientation .
background . many aphid species , including the pea aphid acyrthosiphon pisum , exhibit a behaviour where they drop or fall from their host plant , a commonly used strategy to avoid predation , parasitism or physical disturbance . we hypothesised that there was a physiological non - consumptive cost due to such dropping behaviour because aphids would expend energy re - establishing themselves on a host plant and also lose feeding time .
ranked expression levels of apc transporters ( t . c # 2 . a . 3 ) in a . pisum bacteriocytes , shown relative to gapdh ( acypi009769 ) expression , gapdh = 1 .
acyrthosiphon pisum strain lsr1 was used in this study . parthenogenetic aphid clones were reared on broad bean , vicia faba , in growth chambers with a long - day photoperiod ( 16 l : 8 d cycle ) at 19°c . l3–l4 nymphs and young adults of parthenogenetic females were used to prepare bacteriocyte and whole body samples . more details on dissections and rna extraction are provided in the electronic supplementary material , materials and methods .
in this example , we present a host - endocytobiont pair . the seed set of the metabolic network of b . aphidicola was previously published in borenstein et al . ( 2008 ) ( supplement ) . the metabolic network and seed set of a . pisum was generated using the same approach . the bss of b . aphidicola on a . pisum is 0 . 41667 , which reflects the role of b . aphidicola as an obligate endocytobiont of a . pisum in this mutualistic interaction ( borenstein & feldman 2009 ) . only the giant components are considered here . download the example as a pair of text files
over the years , a variety of aphid species have been used to study the symbiosis with the bacteria budinera ( e . g . ehrhardt , 1969 ; mittler , 1971 ; hinde , 1971 ; campbell & nes , 1983 ) . recent research , however , has focused almost exclusively on one species , the pea aphid acyrthosiphon pisum . between 1992 and june , 1995 , 12 of the 14 publications on symbiotic interactions in aphids concerned a . pisum ( data obtained from bids , the isi data service at bath , uk ) . this situation reflects the amenability of a . pisum for biochemical and nutritional studies ; it is a big aphid , and large numbers can readily be cultured in the laboratory on plants and on chemically - defined diets . it is not known whether the data obtained for the pea aphid symbiosis are relevant to other aphids or peculiar to this species .
acyrthosiphon pisum were used as the model organism for this experiment , were reared in culture on broad bean ( vicia faba var . ‘exhibition long pod’ ) seedlings in ventilated plastic cages ( 75 × 40 × 40 cm ) , and cultured in a controlled temperature growth room at 20 °c , 50–70 % rh with a photoperiod of l16 : d8 h . lighting in the growth room was provided by a bank of horticultural grow - lamps ( grolite e40 ) .
inra and cnrs researchers studied two different aphid species , the pea aphid ( acyrthosiphon pisum ) and the peach - potato aphid ( myzus percicae ) . they discovered that both have several types of macrophage migration inhibitory factors ( mifs ) . the pea aphid has five and the peach - potato aphid has three . these molecules are proteins that play a significant role in mediating the immune responses of vertebrates . this is the first time they have been found to be used by aphids .
pickering j ; gutierrez ap , 1991 . differential impact of the pathogen pandora neoaphidis ( r . & h . ; ) humber ( zygomycetes : entomophthorales ) on the species composition of acyrthosiphon aphids in alfalfa . canadian entomologist , 123 ( 2 ) : 315 - 320
sabater - muñoz b , legeai f , rispe c , bonhomme j , dearden p , dossat c , duclert a , gauthier jp , giblot ducray d , hunter w , dang p , kambhampati s , martinez - torres d , cortes t , moya a , nakabachi a , philippe c , prunier - leterme n , rahbé y , simon jc , stern d , wincker p , tagu d : large - scale gene discovery in the pea aphid acyrthosiphon pisum ( hemiptera ) . genome biology .
phylogenetic reconstruction of a . pisum aaap transporters with related sequences from other taxa reveals a conspicuous arthropod expanded clade ( labeled “arthropod expanded clade” in figures ) that shows evidence of several ancient gene duplications in the common ancestor of arthropods ( fig . 2 ) . the arthropod expanded clade possesses a many - to - many orthologous relationship with human genes slc36a1 - 4 . in contrast , a . pisum and other arthropod aaap transporters found outside the arthropod expanded clade more frequently cluster with single gene representatives from each taxon and tend to have one - to - one human orthologs . a . pisum , drosophila , and tribolium transporters in the arthropod expanded clade have undergone independent species - specific expansions with each small expansion typically containing three to four closely related transporters . we designate the three a . pisum expansions ( 1 ) , ( 2 ) , and ( 3 ) ( fig . 2 ) . in contrast , arthropod gene duplications outside of the arthropod expanded clade are only observed for a . pisum transporters ( acypi26212 and acypi44297 ) and i . scapularis transporters ( iscw018960 and iscw018959 ) with all other aaap transporters ( with the exception of some human transporters ) present as singletons ( fig . 2 ) .
the international aphid genomics consortium has selected the pea aphid acyrthosiphon pisum as the model aphid species ( it has a genome of four holocentric chromosomes and approximately 525 mb ) , and its genome sequencing project has recently been funded . we present here a collection of 40 , 904 high - quality annotated expressed sequence tags ( ests ) generated from different organs of the pea aphid . these ests form 12 , 082 different contigs and singletons , and represent a first significant step towards the comprehensive description of cellular functions involved in aphid biology .
the origin of pea aphid is likely europe or asia , although it is now found throughout the world in regions with temperate climates . in recent years , a closely related species with similar biology , acyrthosiphon kondoi shinji , has also spread nearly worldwide and attacks some of the same crops .
the duplication events defined by the above mentioned species overlap algorithm that only comprised paralogs from a . pisum were considered lineage - specific duplications . whenever more than one round of duplication followed an a . pisum speciation event ( family expansion ) , all resulting paralogs were grouped into a single group of “in - paralogs” . results from all the trees in the phylome were merged into a non - redundant list of in - paralogs groups , by merging groups sharing a significant fraction of their members ( 50 % ) .
members of large a . pisum gene family expansions , those that contain > 10 in - family paralogs , are frequently arranged as tandem duplicates in the genome ( rispe et al . 2008 ; smadja et al . 2009 ; price et al . 2010 ) . clusters of apc and aaap gene duplicates follow this pattern . a total of seven members of the a . pisum apc family expansion share a similar genomic location and map to a 235 . 8 kbp region of genomic scaffold 434 ( acyr 2 . 0 , accession gl350054 ) ( fig . 3 ) . genomic clustering of transporters is also seen for duplicated aaap transporters ; all four a . pisum aaap transporters from expansion ( 2 ) are grouped on a 103 . 5 kbp region of scaffold 385 ( acyr 2 . 0 , accession gl350005 ) , and similar genomic clustering is observed for some members of a . pisum aaap expansions ( 1 ) and ( 3 ) ( fig . 3 ) . we anticipate that the remaining five apc and aaap family expansion members that currently do not map to these four genomic scaffolds are likely located on neighboring scaffolds , an hypothesis that will be resolved in the future by international aphid genomic consortium efforts to physically map the a . pisum genome ( international aphid genomics consortium 2010 ) .
transcripts showing differential accumulation between winged and unwinged morphs of both males and females ( fourth instars or adults ) of the pea aphid ( a . pisum ) , or between winged and unwinged adults females of the peach potato aphid ( m . persicae ) . a
we investigated cross - hybridization of the most upregulated heat shock genes on our array by spiking control rna with labeled pcr products corresponding to hs genes . four buchnera genes ( dnak , ibpa , mopa and yjea ) and one aphid gene ( hsp70 ) were amplified . pcr reactions were carried out for each gene in two 25 μl volumes containing 50 mm kcl , 10 mm tris - hcl , 1 . 5 mm mg 2 + , 2 . 5 mm of each datp , dctp and dgtp , 1 mm dttp , 1 . 5 mm amino - allyl linked dutp ( sigma a0410 - 1mg ) , 0 . 25 units of eppendorf taq dna polymerase and 10 ng of dna ( buchnera genes ) or cdna ( est6 – hsp70 ) . primers used to amplify genes selected for spiking were as follows : buchnera of a . pisum dnak : f 5 ' caccagagagaactcctccc 3 ' and r 5 ' atggatggcaataaaccacg 3 ' , buchnera of a . pisum ibpa : f 5 ' tcaattaacactaagtattcctgg 3 ' and r 5 ' taggtttttcctcttccgg 3 ' , buchnera of a . pisum mopa : f 5 ' accacaacagcaacattattagc 3 ' and r 5 ' acctccagcaactacaccttc 3 ' , buchnera of a . pisum yjea : f 5 ' gaagcgtttattagcatcag 3 ' and r 5 ' tgaacaaggaggtaagcc 3 ' , a . pisum hsp70 : f 5 ' gtgttgatatttgacctgggc 3 ' and r 5 ' aatggtagagttgcgttcgac 3 ' .
recent genomic analyses of arthropod defense mechanisms suggest conservation of key elements underlying responses to pathogens , parasites and stresses . at the center of pathogen - induced immune responses are signaling pathways triggered by the recognition of fungal , bacterial and viral signatures . these pathways result in the production of response molecules , such as antimicrobial peptides and lysozymes , which degrade or destroy invaders . using the recently sequenced genome of the pea aphid ( acyrthosiphon pisum ) , we conducted the first extensive annotation of the immune and stress gene repertoire of a hemipterous insect , which is phylogenetically distantly related to previously characterized insects models .
recent genomic analyses of arthropod defense mechanisms suggest conservation of key elements underlying responses to pathogens , parasites and stresses . at the center of pathogen - induced immune responses are signaling pathways triggered by the recognition of fungal , bacterial and viral signatures . these pathways result in the production of response molecules , such as antimicrobial peptides and lysozymes , which degrade or destroy invaders . using the recently sequenced genome of the pea aphid ( acyrthosiphon pisum ) , we conducted the first extensive annotation of the immune and stress gene repertoire of a hemipterous insect , which is phylogenetically distantly related to previously characterized insects models .
given that “ ca . serratia symbiotica” cluster a has been demonstrated to confer heat tolerance in a . pisum ( 19 , 29 ) , it is likely that other symbiont lineages in this clade have similar effects on their aphid hosts , although this has not been tested experimentally .
rates of molecular evolution were estimated for a subset of a . pisum apc transporters that form an a . pisum - specific slimfast expansion . the analysis was conducted using the ten a . pisum paralogs that form the apc family expansion ( acypi003523 , acypi005156 , acypi003240 , acypi000537 , acypi000584 , acypi05472 , acypi005118 , acypi008904 , acypi08323 , and acypi002633 ) , insect slimfast orthologs from a . aegypti ( aaslif , aat73699 ) , d . melanogsater ( slimfast , cg11128 ) , t . castaneum ( xp _ 967023 ) , a . mellifera ( xp _ 393144 ) , and p . humanus ( phum258130 ) , and outgroup sequences from the sister clade of the expansion ( a . pisum , acypi003923 and d . melanogaster , cg5535 ) . the amino acid alignment of these 17 sequences was generated in mafft ( katoh et al . 2005 ) and used to reconstruct the phylogeny of the genes in mrbayes v . 3 . 1 . 2 . bayesian trace plots were analyzed , and the burn - in value determined as described above and the resulting majority rule consensus tree used as input for the molecular evolution analyses .
vertical dotted lines show the estimated average ds between orthologs from different aphid species . 1 : a . pisum and myzus persicae ( two species of the tribe macrosiphini ) , mean ds = 0 . 25 ; 2 : a . pisum and aphis gossypii ( tribe aphidini ) , mean ds = 0 . 35 ( estimates from ) . paralogs resulting from ancient duplications ( ds > 1 . 5 ) are also abundant in all three genomes ( 1 , 449 pairs in aphid , 1 , 726 in drosophila , 1 , 010 in bee ; not shown ) .
vertical dotted lines show the estimated average ds between orthologs from different aphid species . 1 : a . pisum and myzus persicae ( two species of the tribe macrosiphini ) , mean ds = 0 . 25 ; 2 : a . pisum and aphis gossypii ( tribe aphidini ) , mean ds = 0 . 35 ( estimates from [ 128 ] ) . paralogs resulting from ancient duplications ( ds > 1 . 5 ) are also abundant in all three genomes ( 1 , 449 pairs in aphid , 1 , 726 in drosophila , 1 , 010 in bee ; not shown ) .
hamity vc ; amg de hamity mg , 2000 . monitoreo de la poblacion de aphidae y sus predadores en cultivos de vicia faba y pisum sativum con manejo agrecologico . presented at the xxi congreso nacional de entomologia , november 1999 , arica , chile . idesia , 18 : 41 - 48 .
the evolution of ecological specialization generates biological diversity and may lead to speciation . genetic architecture can either speed or retard this process . if resource use and mate choice have a common genetic basis through pleiotropy or close linkage , the resulting genetic correlations can promote the joint evolution of specialization and reproductive isolation , facilitating speciation . here we present a model of the role of genetic correlations in specialization and speciation , and test it by analysing the genetic architecture of key traits in two highly specialized host races of the pea aphid ( acyrthosiphon pisum pisum ; hemiptera : aphididae ) . we found several complexes of pleiotropic or closely linked quantitative trait loci ( qtl ) that affect key traits in ways that would promote speciation : qtl with antagonistic effects on performance on the two hosts are linked to qtl that produce asortative mating ( through habitat choice ) . this type of genetic architecture may be common in taxa that have speciated under divergent natural selection .
the genome sequences used in this study were retrieved from genbank database . the four b . aphidicola strains are b . aphidicola acyrthosiphon pisum str . aps ( bap , accession no . ba000003 [ 20 ] ) , b . aphidicola schizaphis graminum ( bsg , accession no . ae013218 [ 21 ] ) , b . aphidicola baizongia pistaciae ( bbp , accession no . ae016826 [ 22 ] ) , and b . aphidicola cinara cedri ( bcc , accession no . cp000263 [ 15 ] ) . escherichia coli was used as out - group in all comparisons : e . coli str . k12 substr . mg1655 ( eco , accession no . u00096 ) .
genome distribution of genes with strong selective constraints in endosymbiotic bacteria . we identified genes with strong constraints in each of the five endosymbiotic bacterial lineages : ( a ) buchera aphidicola ( from acyrthosiphon pisum ) ; ( b ) candidatus blochmannia ( blochmannia floridanus ) ; ( c ) blattabacterium from periplaneta americana ; ( d ) wigglesworthia glossinidia ; and ( e ) ca . baumannia cicadellinicola . genes indicated in the circle - representation of the endosymbiotic genomes are those that were specifically identified in that endosymbiotic genome and not the others from the same group . these genes are mostly related to the metabolism of the bacterium that interacts with the metabolism of the insect host .
dls provided cdna libraries . wbh sequenced and provided clones of a . pisum ests . dls and nam designed the array . gkd pcr - amplified aphid ests . hed designed primers for and pcr - amplified buchnera of a . pisum genes . accw and hed executed heat shock experiments and prepared rna for array hybridization . accw annotated the array and analyzed the microarray data . hed designed , executed and analyzed rt - qpcr reactions . accw verified spot identity by resequencing . hed , accw and nam designed and executed the spike experiment . accw and nam drafted the manuscript . all authors read and approved the final manuscript .
huybrechts h , bonhomme j , minoli s , prunier - leterme n , dombrovsky a , et al . ( 2010 ) neuropeptide and neurohormone precursors in the pea aphid acyrthosipon pisum . insect mol biol . ( doi : 10 . 1111 / j . 1365 - 2583 . 2009 . 00951 . x ) .
approximately 200 ma , aphids evolved novel cells , called bacteriocytes , that house the gamma - proteobacterial endosymbiont , buchnera aphidicola . physiological studies and whole - genome sequence analysis of acyrthosiphon pisum and buchnera have revealed an intimate metabolic collaboration between partners . buchnera provides the host with nutrition such as amino acids that aphids cannot synthesize and that are deficient in plant phloem sap , aphids ' sole dietary component ( reviewed in [ 1 , 2 ] ) . the aphid– buchnera association is obligate and mutualistic with neither partner being able to reproduce in the absence of the other [ 1 , 2 ] . the symbionts are passed directly from mother to offspring by transovarial transfer [ 3 , 4 ] .
the best studied insect - symbiont system is that of aphids and their primary bacterial endosymbiont buchnera aphidicola . buchnera inhabits specialized host cells called bacteriocytes , provides nutrients to the aphid and has co - speciated with its aphid hosts for the past 150 million years . we have used a single microarray to examine gene expression in the pea aphid , acyrthosiphon pisum , and its resident buchnera . very little is known of gene expression in aphids , few studies have examined gene expression in buchnera , and no study has examined simultaneously the expression profiles of a host and its symbiont . expression profiling of aphids , in studies such as this , will be critical for assigning newly discovered a . pisum genes to functional roles . in particular , because aphids possess many genes that are absent from drosophila and other holometabolous insect taxa , aphid genome annotation efforts cannot rely entirely on homology to the best - studied insect systems . development of this dual - genome array represents a first attempt to characterize gene expression in this emerging model system .
acyrthosiphon pisum , with 34 604 predicted genes in its 464 - mb draft genome assembly , has the highest gene count of any insect sequenced to date , although a precise gene count awaits better genome assembly and further analyses [ 5 ] . this high gene count reflects both extensive gene duplications and the presence of aphid - specific orphan genes . orphan genes , which display no significant similarity with genes identified in other sequenced organisms , comprise 20 per cent of the total number of genes in the pea aphid genome [ 5 ] . the functions of these orphan genes are unknown , but their evolution may be related to the evolution of lineage - specific traits in aphids , such as symbiosis with buchnera .
to understand the host ' s role in this symbiotic system , mrna of the host bacteriocyte of the pea aphid acyrthosiphon pisum has been assessed by expressed sequence tag ( est ) - based transcriptome analyses [ 6 , 7 ] and , more recently , by deep - sequencing transcriptome analysis [ 8 ] . many of the over - represented transcripts are associated with the utilization of essential amino acids , biosynthesis of non - essential amino acids and transport , which support the metabolic collaboration between the host and symbiont . in addition , laterally acquired genes of alpha - proteobacterial origin are also preferentially transcribed in bacteriocytes [ 6 , 9 , 10 ] . orphan genes have not been the focus of previous studies .
finally , when unmapped reads were re - mapped to the partially assembled genome of spiroplasma isolated from a . pisum , individuals which had been found to be positive for spiroplasma by a pcr - based diagnostic assay but which had a negligible coverage when their reads were mapped to s . melliferum registered a high coverage on contigs from the a . pisum - derived spiroplasma ( up to 1185 × for some individuals , supplementary table s2 , supplementary material ) . in one case , the sensitivity of the sequence analysis may have exceeded that of the pcr test as individual lc3 was pcr - negative for spiroplasma but recorded on re - mapping , suggesting a possible infection under the threshold of pcr detection .
we examined gene expression under heat shock of an insect and its bacterial symbiont in a single assay using a dual - genome microarray . further , our results indicate that microarrays are a useful tool for inferring functional roles of genes in a . pisum and other insects and suggest that the pea aphid genome may contain many gene paralogs that are differentially regulated .
unmapped reads are often discarded from the analysis of whole - genome re - sequencing , but new biological information and insights can be uncovered through their analysis . in this paper , we investigate unmapped reads from the re - sequencing data of 33 pea aphid genomes from individuals specialized on different host plants . the unmapped reads for each individual were retrieved following mapping to the acyrthosiphon pisum reference genome and its mitochondrial and symbiont genomes . these sets of unmapped reads were then cross - compared , revealing that a significant number of these unmapped sequences were conserved across individuals . interestingly , sequences were most commonly shared between individuals adapted to the same host plant , suggesting that these sequences may contribute to the divergence between host plant specialized biotypes . analysis of the contigs obtained from assembling the unmapped reads pooled by biotype allowed us to recover some divergent genomic regions previously excluded from analysis and to discover putative novel sequences of a . pisum and its symbionts . in conclusion , this study emphasizes the interest of the unmapped component of re - sequencing data sets and the potential loss of important information . we here propose strategies to aid the capture and interpretation of this information .
real - time quantitative polymerase chain reaction ( qpcr ) was performed on a . pisum cdna from bacteriocytes , gut and head ( including salivary glands ) tissues , and whole parthenogenetic adult females of three a . piusm lineages that included lsr1 ( caillaud et al . 2002 , the parental line of the inbred line used in the a . pisum genome project ; international aphid genomics consortium 2010 ) , 5a ( sandstrom and moran 2001 ) , and 9 - 2 - 1 ( russell and moran 2006 ) . tissues were dissected from 100 parthenogenetic females in nuclease - free 0 . 9 % saline and immediately lysed in tri reagent ( ambion ) . total rna was isolated and further purified by rna cleanup with an rneasy mini kit ( qiagen ) , which included a dnasei digest . whole insect total rna was isolated from ten wingless adult females using an rneasy mini kit and on - column rnase - free dnasei treatment . first - strand cdnas were synthesized from 500 ng total rna from whole a . pisum , and head and gut tissues and 100 ng of total rna from bacteriocyte tissues , using qscript cdna supermix ( quanta biosciences ) in standard 20 μl reactions .
the pea aphid acyrthosiphon pisum clone ll01 was reared on preflowering vicia faba cv . sutton at 20°c with 18 l : 6 d . it was confirmed to bear no detectable internal bacteria other than buchnera by microscopical examination , pcr assays with general 16s rrna gene primers and microbiological cultivation experiments . aposymbiotic aphids , i . e . lacking the symbiotic bacterium buchnera , were generated by oral administration of the antibiotic rifampicin at 50 µg ml −1 diet over 2 days after birth ; and aphids were used when 7 days old , at which time buchnera were undetectable microscopically and by buchnera - specific pcr assay [ 20 ] . the drosophila s2 cells were cultured aseptically in schneider ' s drosophila medium ( sdm ) + 10 % foetal calf serum ( fcs ) ( invitrogen ) at 20°c and maintained in exponential phase by passaging every fourth day .
cpg ratios were calculated using refseq data for each insect species . for each sequence the observed ( obs ) cpg frequency and the expected ( exp ) cpg frequency were calculated . the expected cpg frequency was calculated based on the gc content of each sequence and the cpg ratio was calculated as obs / exp . the frequency of each cpg ratio was plotted against the observed / expected ratio . a bimodal distribution was observed for a . pisum and a . mellifera , both of which show dna methylation within the coding sequence of genes , . d . melanogaster and t . castaneum both show a unimodal distribution , and there is only limited evidence of methylation in both of these species . in addition a . pisum and a . mellifera have all the dna methyltransferases while d . melanogaster only has dnmt2 and t . castaneum has dnmt1 and dnmt2 .
quantitative gene expression analysis of acyrthosiphon pisum apc and aaap transporters . gene expression profiles of a . pisum ( lineage lsr1 ) aats from : ( a ) apc transporter family ( t . c # 2 . a . 3 ) slimfast expansion ; and ( b ) aaap transporter family ( t . c # 2 . a . 18 ) expansions ( 1 ) , ( 2 ) , and ( 3 ) together with representative a . pisum aaaps with one - to - one orthology to aaaps in other taxa . transcript levels for each transporter gene were quantified using qpcr on cdna from adult female whole insects ( wi ) and cdna from head ( h ) , gut ( g ) , and bacteriocyte ( b ) tissues . the relative abundance of each transporter gene is normalized to gapdh ( acypi009769 ) , and expression data are compiled into a heat map , as described in the materials and methods . transporter genes that are enriched in head ( h ) , gut ( g ) , and bacteriocyte ( b ) tissues > 4 - fold relative to wi expression levels are indicated to the right of the heat map and mean fold - enrichment is shown for indicated tissue , with confidence limits ( livak and schmittgen 2001 ) indicated in parentheses ( n = 3 ) . transporters with < 4 - fold enrichment relative to whole insect in analyzed tissues are indicated . expression of aaap transporter acypi53198 was not detectable ( n . d . ) in wi or any of the tissues analyzed . finally , because of high sequence similarity , it was not possible to design gene - specific primers to determine the expression profiles of closely related apc family transporters acypi000537 , acypi000584 , and acypi005472 and aaap family transporters acypi008957 and acypi007681 ( shown in gray ) .
cpg ratios were calculated using refseq data for each insect species . for each sequence the observed ( obs ) cpg frequency and the expected ( exp ) cpg frequency were calculated . the expected cpg frequency was calculated based on the gc content of each sequence and the cpg ratio was calculated as obs / exp . the frequency of each cpg ratio was plotted against the observed / expected ratio . a bimodal distribution was observed for a . pisum and a . mellifera , both of which show dna methylation within the coding sequence of genes [ 37 ] , [ 129 ] . d . melanogaster and t . castaneum both show a unimodal distribution , and there is only limited evidence of methylation in both of these species . in addition a . pisum and a . mellifera have all the dna methyltransferases while d . melanogaster only has dnmt2 and t . castaneum has dnmt1 and dnmt2 .
in the same way , primers able to amplify the targeted genes in all the sequenced wolbachia strains were designed to confirm the absence in wpn of genes that were found in the bpn genome . for each test , an acyrthosiphon pisum was used as a positive control for buchnera . for each primer pair , pcr amplifications were performed in 50 - μl reactions containing 2 μl of dna , 5 μl of 10× reaction buffer ( roche ) , 5 μl of dntp ( 2 . 5 mm each ) , 1 . 5 μl of mgcl 2 ( 50 mm ) , 1 μl of each primer ( 25 μm ) , 0 . 2 μl of taq polymerase ( roche , 1 u / μl ) , and 34 . 3 μl of water . the temperature cycle corresponded to an initial denaturation step at 95 °c followed by 40 cycles of 30 s at 95 °c , 1 min at 52 °c , and 2 min at 72 °c .