forrest 's pika ( ochotona forresti ) is a species of mammal in the family ochotonidae .

it is found in bhutan , china , india , and myanmar .

in india , it has been recorded from arunachal pradesh in the north-east .

forrest ' s pika

forrest ' s pikas are found in bhutan , china , india and myanmar .
dark grey area behind each ear
and these two areas almost meet across the nape in the forrest ' s pika , unlike the pale buffy area which does not meet in the moupin pika
.
genome - wide comparative chromosome map between human and the forrest ' s pika ( ochotona forresti ) established by cross - species chromosome painting : further support for the glires hypothesis .
70 % of the forrest ' s pika studied had violin shaped
confluence of the incisive and palatal foramina due to the abrupt constriction of the sides , whereas this constriction is not found in the moupin pika
.
genome - wide comparative chromosome map between human and the forrest ' s pika ( ochotona forresti ) established by cross - species chromosome painting : f . . . - pubmed - ncbi
ochotona forresti can be distinguished from ochotona gaoligongensis - gaoligong pika by the gaoligong pika ' s brilliant rufous - brown head and neck , and a dull rufous - black back .
ochotona roylei - royle ' s pika
and with it may form a superspecies
. ( b605 . 3 . w3 )
there is little information on this species and so it is unclear if this species is a burrowing pika , a rock dwelling pika or an intermediate habitat type of pika .
ochotona gaoligongensis - gaoligong pika : from the original description , this form is likely to be proved a synonym or sister species of the forrest ' s pika which also occurs in the same area . they appear to be morphologically similar . ( b605 . 3 . w3 , b607 . w20 )
there is little information on this species and so it is unclear if this species is a burrowing pika , a rock - dwelling pika or an intermediate habitat type of pika .
note : there is little information on this species and so it is unclear if this species is a burrowing pika , a rock dwelling pika or an intermediate habitat type of pika .
genome - wide comparative chromosome map between human and the forrest’s pika ( ochotona forresti ) established by cross - species chromosome painting : further support for the glires hypothesis - fulltext - cytogenetic and genome research 2011 , vol . 132 , no . 1 - 2 - karger publishers
forrest ' s pika has fairly dark reddish fur , with a dark grey area behind each ear ( nearly meeting each other ) ; the dorsal fur becomes dark grey - brown in winter , while the ventral fur is slightly lighter than this . ( b605 . 3 . w3 )
r as , r fb r g ra nt s ( a . s . g . ) . f . y . is su pp or ted by th e we ll com e tr us t .
the name pika originated from the tungus of siberia who attempted to mimic the call
peeka
of the local pika species . ( b285 . w5g )
ochotona gloveri - glover ' s pika : ( the calloceps form ) , yunnan , china ; burma ; and sikkim , india . ( b605 . 3 . w3 ; b607 . w20 )
it is not known whether this is a rock - dwelling or burrowing pika .
embryo resorption may occur if the pika encounters adverse conditions . ( b285 . w5a )
the o . forresti fibroblast cell line was derived from skin biopsies of a male forrest’s pika collected from gaoligong mountain , yunnan , china . metaphase preparations were made following conventional methods as previously described [ yang et al . , 2003a ] . the o . forresti chromosomes were karyotyped based on inverted dapi - banding patterns that are similar to the g - banding patterns .
in main characters and size of the skull . however it can be distinguished from this pika
in this study , we have established the first genome - wide chromosomal homology map between human and forrest’s pika ( ochotona forresti , ofo , 2n = 54 ) by cross - species chromosome painting with human chromosome - specific painting probes . such a map sheds further insight into the ancestral lagomorph karyotype and chromosomal rearrangements underlying the karyotype divergence between leporidae and ochotonidae and provides further support for the glires hypothesis .
ochotona macrotis - large - eared pika ( the chinensis form ) ( b605 . 3 . w3 )
the upper incisors ' roots are found in the skull ' s premaxillary bones . however , the length of the lower incisors ' roots varies . (
note : there is very little data specific to this species so the details below are from general pika information .
this species appears to be a sister species of ochotona erythrotis - chinese red pika . ( b607 . w20 )
many species of pika inhabit very restricted ranges and so may be threatened by human environmental disruption . ( b147 )
ochotona thibetana - moupin pika based on a study of the holotype . ( b605 . 3 . w3 , b607 . w20 )
the fore claws are notably longer in this species than that of ochotona thibetana - moupin pika . ( b605 . 3 . w3 )
general pika information : pikas have small , rounded ears which are 12 - 36 mm in length . ( b147 ; b285 . w5g )
in general , the status of many species of pika is hard to assess because they inhabit such remote areas . ( b285 . w5g )
ochotona nigritia - black pika :
apparently close to o . forresti ; may be only a melanistic individual
. ( b607 . w20 )
general pika information : these are shorter in burrowing pikas , such as this species , compared to rock dwelling pikas . ( b605 . 3 . w3 )
pika species living in areas where winter snow is common may also make tunnels in the snow to reach and harvest any nearby vegetation . ( b285 . w5g )
this study is supported partly by a grant from the national natural science foundation of china ( no . 30770293 ) and mcb , sb ras , rfbr grants ( a . s . g . ) . f . y . is supported by the wellcome trust .
chantry - darmon c , bertaud m , urien c , chadi - taourit s , perrocheau m , et al : expanded comparative mapping between man and rabbit and detection of a new conserved segment between hsa22 and ocu4 . cytogenet genome res 111 : 134–139 ( 2005 ) .
pishchukha
is the russian common name for all species of pika and some gerbils ( rhombomys opimus , meriones tamareiscinus ( muridae - ( family ) ) . ( b605 . 3 . w3 )
70 % of the ochotona forresti studied had violin shaped
confluence of the incisive and palatal foramina due to the abrupt constriction of the sides , whereas this constriction is not found in the moupin pika
.
cross - species chromosome painting with human chromosome - specific painting probes has allowed us to construct the first genome - wide comparative chromosome map between human and o . forresti , a representative species of the pika family . an integrated analysis of our comparative chromosome map and the previously published maps [ korstanje et al . , 1999 ; robinson et al . , 2002 ] provides new insight into the comparative genome organization of the lagomorpha .
little or nothing is known regarding the behavior , ecology , and reproduction of ochotona forresti ( smith et al . 1990 ) . o . forresti inhabits mountain slopes at elevations of 2 , 600 to 4 , 400 m in the forest belt ( feng et al . 1986 ) . these high altitude habitats are typically coniferous or mixed broadleaf and conifer forests and shrubland ( smith and xie 2008 ) . it is speculated that this species of pika creates burrows ( smith and xie 2008 ) .
in spring , summer and / or autumn ( fall ) ( depending on species / location ) many pika species spend much time
haying
- harvesting mouthfuls of vegetation which are carried back to the den for storage . they build up these stores , resembling piles of hay , and use them for consumption during periods of sparse vegetation , often over - harvesting so that it is a rare occurrence for them to run out of food . ( b285 . w5g , b605 . 3 . w3 )
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lissovsky , a . a . 2014 . taxonomic revision of pikas ochotona ( lagomorpha , mammalia ) at the species level . mammalia 78 ( 2 ) : 199–216 .
justification : this species is listed as least concern because it is unlikely to be declining at nearly the rate required to qualify for listing in a threatened category .
ochotona forresti ranges from northwest yunnan to northern myanmar . the mapped range in myanmar is totally speculative ( duckworth and thaw pers . comm ) . ochotona forresti occupies elevations of 2 , 600 - 4 , 400 m .
the current population status of ochotona forresti is unknown ; there have been no ecological or natural history investigations of this species .
due to the general lack of data regarding the population status , behaviour , ecology , and reproduction research in these areas should be undertaken . this species has been regionally assessed in china as near threatened ( jiang et al . 2016 ) .
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this species is listed as least concern in view of its wide distribution , presumed large population , occurrence in protected areas , and because it is unlikely to be declining at nearly the rate required to qualify for listing in a threatened category .
this species occurs in nujiang and gaoligongshan ( yunnan ) nature reserves in china ( csis 2008 ) . due to the general lack of data regarding the population status , behavior , ecology , and reproduction research in these areas should be undertaken . this species has been regionally assessed in china as near threatened , nearly meeting the criteria for vulnerable under criteria a2c + 3c ( wang and xie 2004 ) .
. in india , it has been recorded from arunachal pradesh in the north - east .
smith , a . t . & johnston , c . h . ( 2008 ) . ochotona forresti . in : iucn 2008 . iucn red list of threatened species . retrieved 10 april 2009 . database entry includes a brief justification of why this species is of least concern .
choudhury , a . u . ( 2003 ) . the mammals of arunachal pradesh . regency publications , new delhi . 140pp .
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the generic name of ochotona is derived from the mongolian name for pikas :
ogdoi
. ( b285 . w5g )
pikas are small , egg - shaped , rodent - like lagomorphs which weigh under 500 g . they have rounded , relatively large ears , short legs , and a very short tail which is hardly visible . ( b285 . w5g )
newborn pikas are helpless and naked ( b147 , b287 ) or slightly furred . ( b287 )
rectangular shaped posterior end of the nasals are broader
. ( b605 . 3 . w3 )
male and female pikas are similar in size and can be difficult to tell apart from one another . ( b147 )
125 - 300 mm , with most species averaging around 200 mm or less . ( b147 )
range 4 . 1 - 12 . 7 g , depending on species . ( b287 )
in general , the head of pikas is blunt and short , and the skull is quite flattened rather than arched . there is also a constriction between the orbits . ( b147 )
there are 26 teeth in total - two less than other lagomorphs who have one more upper molar on each side . ( b285 . w5a , b605 . 1 . w1 )
the dental formula of pikas is 2 / 1 incisors , 0 / 0 canines , 3 / 2 premolars , and 2 / 3 molars . ( b147 , b605 . 1 . w1 )
lagomorphs , including pikas , differ from rodents by having two pairs of upper incisors rather than just the one pair . the additional set of incisors are called peg teeth and are found directly behind the long pair in the upper jaw . ( b147 , b285 . w5a , b605 . 1 . w1 )
at birth , lagomorphs actually have three pairs of upper incisors , but they quickly lose the outer incisor on each side . ( b147 )
[ note : lagomorphs have teeth which grow throughout their lives . for this reason the portion of the teeth which is not exposed ( not above the gum line ) is strictly speaking not a
root
; however , it is sometimes convenient to describe it as a root . ]
the first upper incisors have a cutting edge which is v - shaped . ( b147 )
pikas have high crowned cheek teeth with no roots [ the teeth grow continuously throughout life ] . ( b147 )
the lower tooth rows are closer together than the upper tooth rows . ( b147 )
adult : pikas have eyes positioned to give a broad field of vision ( b285 . w5a )
newborn : neonates are blind ; the eyes open at eight to ten days . ( b287 )
the hindlimbs are just slightly longer than the forelimbs . ( b147 , b430 . w2 , b605 . 2 . w2 )
in burrowing pikas , the claws are more straight and powerful than those of the rock dwelling pikas . ( b605 . 3 . w3 )
the tail of pikas is virtually absent at a length of 5 mm ( b285 . w5g ) ; it is not visible . ( b147 ; b430 . w2 )
fine , long , soft and dense coat with fur that covers the feet including the under surface . ( b147 , b285 . w5g )
most species have two moults per year with a brighter summer coat - often a yellowish red - and a greyer winter coat . ( b147 )
newborn pikas are hairless ( b147 , b287 ) or slightly furred . ( b287 )
the uterus is duplex . the placenta is discoid , deciduate and hemochorial , with a mesometrial , superficial implantation . ( b287 )
the testes are intra - abdominal outside the breeding season . ( b147 , b287 )
during the breeding season they are found in folds of skin at the base of the penis . ( b147 )
scent glands : pikas have scent glands , as do all lagomorphs . ( b285 . w5a )
in general , pikas breed twice a year in the spring and summer , and many species will have two or more litters per year . ( b147 )
pikas show induced ovulation and a post - partum oestrus . ( b285 . w5a , b287 )
in general , burrowing pikas have litters which are twice as large as those of rock dwelling pikas . (
rock dwelling pikas have few litters per year . they may have two litters annually but often only one is successfully weaned ( b285 . w5g , b605 . 3 . w3 )
burrowing pikas may have as many as five litters a year , depending on species . ( b285 . w5g )
burrowing pikas may mature and breed in their summer of birth . however , young rock dwelling pikas will first breed as yearlings . ( b605 . 3 . w3 )
the testes are intra - abdominal outside the breeding season ; during the breeding season they are found in folds of skin at the base of the penis ( in lagomorphs , the testes are in front of the penis ) . ( b147 )
high mortality as pikas are prey for many mammals and birds . ( b285 . w5a )
burrowers : up to three years of age , but usually only one year . ( b285 . w5g )
burrowing pikas have a high annual mortality , with few animals living more than two years . ( b605 . 3 . w3 )
rock dwellers : up to seven years old . ( b285 . w5g ) their average mortality is low compared to the burrowing pikas . ( b605 . 3 . w3 )
pikas are herbivorous - they eat grasses , flowering stalks , and leaves . pikas have a preference for those plants highest in protein or other chemicals important to them . ( b285 . w5g )
pikas eat a range of vegetable matter :
in the summer and early autumn the animals gather grasses , sedges , weeds , and many of the large flowering and woody plants , sometimes climbing a few meters up in trees and out on limbs to cut twigs . the material is sometimes place in exposed locations for curing by the sun
; many populations create haystacks to store food for winter . ( b147 )
pikas do not hibernate . ( b147 , b285 . w5g , b605 . 3 . w3 )
jaw motion : pikas have a vertical or transverse jaw motion . ( b147 ) ; pikas use a side - to - side jaw motion . ( b285 . w5g )
coprophagy : pikas produce two types of faeces , hard faeces like pepper seeds - small green spherical pellets - which are passed during the day ; and soft faeces , sticky and dark green / black , passed at night . faeces of the latter type have high a energy value and b vitamin levels , and are re - ingested . this behaviour , known as coprophagy , may have a similar function to the ruminant behaviour of chewing the cud . ( b147 , b285 . w5a )
pikas are known to be more vocal than other lagomorphs . ( b285 . w5a )
and a long call used by males during the breeding season which is
a series of squeaks lasting up to 30 seconds
. some rock dwellers rarely vocalise even weak calls . ( b285 . w5g )
muffle and transition calls used by young pikas , which are thought to promote cohesion among siblings . ( b285 . w5g )
pikas have scent glands , as do all lagomorphs . ( b285 . w5a )
pikas are unable to grasp plants with their forepaws ; they eat with a side - to - side jaw motion and carry vegetation in their mouths . ( b285 . w5g )
some species continue to forage throughout winter rather than haying , because snows are uncommon . ( b285 . w5g )
even at a fairly low population density of ten to twelve pikas per hectare , vegetation storage by pikas may be up to 30 kg per hectare . ( b605 . 3 . w3 )
in the burrowing pikas , the young may form a line behind an adult , usually their father , and follow . ( b285 . w5g )
it is rare that they interact and usually it is to repel an intruder if they do so . ( b285 . w5g )
even in a pair of pikas which are contributing to a shared hay pile , they spend a large part of the day apart . ( b285 . w5g )
the rock dwelling pikas have large territories defended by the individual ( in north american species ) or defended in pairs ( asian species ) . ( b285 . w5g , b605 . 3 . w3 )
the population density is therefore low , at 5 - 25 per acre , and reasonably stable over a period of time . (
population densities of pikas in rocky areas do not usually reach more than 20 per hectare . ( b147 )
these pikas are very friendly , sociable mammals that live within family groups where they may play - box , sit in contact , nose rub and spend time socially grooming . ( b285 . w5g )
communal dens house family groups which includes siblings of different ages . ( b285 . w5g )
the young may follow behind an adult , usually their father , in a line . ( b285 . w5g )
however , there may be aggression between members of different family groups , in particular , long chases of adult males occur . ( b285 . w5g )
population densities of burrowing steppe dwellers are often much higher than that of the rock dwellers but they are also prone to fluctuate more widely . ( b147 )
the population density may be greater than 750 per acre towards the end of the breeding season but this may fluctuate greatly both annually and seasonally . ( b285 . w5g , b605 . 3 . w3 )
the maximum density of some steppe pikas is reported to exceed 300 per hectare . ( b147 )
in general , it appears that pikas are monogamous . ( b285 . w5g , b605 . 3 . w3 )
mainly active by day . pikas are well - adapted to the cold and sensitive to even moderately warm conditions , therefore they tend to be active only during the cooler parts of the day . ( b285 . w5g )
pikas may be active at all hours , in particular , early morning and evenings . it seems that they are less active on sunny days compared with cloudy days . ( b147 )
pikas which live at high altitudes may be active all day , whereas pikas at warmer , lower altitudes emerge only in the morning and evening . ( b285 . w5g )
most pikas live in remote high mountains and wild country and are well adapted to the cold . pikas have become well adapted to living in rocky steppe and alpine habitats . ( b285 . w5g , b605 . 3 . w3 )
this species occupies mountain slopes that face the sun . ( b605 . 3 . w3 )
elevation : 2600 to 4400 metres in the forest belt . ( b605 . 3 . w3 )
burrowing pikas dig burrows in open alpine meadow , semi desert or steppe environments . ( b285 . w5g )
rock dwelling pikas nest among rocks or fallen logs . ( b285 . w5g )
certain pikas are intermediate in their habitat use in that they burrow but also sometimes live among rocks . however their life history is closer to that of the burrowing pikas . ( b285 . w5g )
it is common for burrows to be shared with birds or small mammals . ( b605 . 3 . w3 )
haypiles created by pikas may provide winter food for domestic cows and horses and also native species such as ungulates or smaller herbivores .
by recycling soil , burrowing pikas have a positive contribution to ecosystem - level dynamics . ( b605 . 3 . w3 )
currently no species or forms of ochotona are treated on any national list of endangered or threatened wildlife .
( b605 . 3 . w3 )
there are currently no ochotona species cites - listed . ( w354 . april08 . w1 )
iucn - lower risk / near threatened . ( w2 . apr08 . w44 )
the status of this species is unknown .
however , its restricted distribution suggests that it may be at some risk
. ( b605 . 3 . w3 )
due to their remote habitat , most pikas
rarely come into conflict with human economic activity .
( b147 )
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ye j 1 , nie w , wang j , su w , jing m , graphodatsky as , yang f .
state key laboratory of genetic resources and evolution , kunming institute of zoology , the chinese academy of sciences , kunming , pr china .
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the order lagomorpha consists of 2 families : leporidae ( hares and rabbits ) , and ochotonidae ( pikas ) , about 91 species [ hoffmann and smith , 2007 ] , of which the domestic rabbit ( oryctolagus cuniculus , 2n = 44 ) is the most well - studied species mainly due to its importance as a model for biomedical research [ korstanje et al . , 1999 ] .
comparative cytogenetic studies have made a great contribution to our understanding of karyotype evolution of species within the families leporidae and ochotonidae , respectively [ hsu and benirschke , 1971 ; stock , 1976 ; capanna et al . , 1991 ; ivanitskaya , 1991 ; robinson et al . , 2002 ; robinson , 2006 and literature cited therein ] . however , earlier cross - family comparison based on g - banding has only revealed a few conserved chromosomal segments between representative species from leporidae and ochotonidae , leading to the suggestion that tandem fusion could be responsible for the family - level karyotype divergence in lagomorpha [ stock , 1976 ] .
fluorescence in situ hybridization ( fish ) images were captured using the genus system ( applied imaging corp . ) as previously described [ yang et al . , 2004 ] . hybridization signals were assigned to specific chromosomes or chromosomal regions as defined by enhanced dapi - banding patterns .
examples of cross - species chromosome painting . a hsa chromosome9probe hybridized onto the short arm of o . forresti chromosome 3 . b dapi - banding of the same o . forresti metaphase . c hsa chromosome15probe hybridized onto the long arm of o . forresti chromosome 3 . d dapi - banding of the same o . forresti metaphase .
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1993 ; murphy et al . , 2001a ; reyes et al . , 2004 ; a
v , 2007 for review ; nie et al . , 2008 ] . nevertheless ,
rmed by a later study [ chantry - darmon et a l . , 2005 ]
. . . thus , o . huangensis is significantly diverging from the main group of the subgenus ochotona by main karyotypic characteristics that corresponds to the data of mtdna study ( yu et al . 2000 , niu et al . 2004 ) . the recent cytogenetic study of o . forresti ( 2n = 54 ) ( ye et al . 2011 ) also greatly expands karyotypic variability of the subgenus conothoa . as yet , the karyotypes of eight species of the subgenus conothoa and three species of the subgenus ochotona are not investigated . . . .
. . . the presence of the nox5 gene in rabbits makes the rabbit a potentially useful model to study nox5 gene functions . its presence in rabbits and its absence in rodents indicates it is a potential genetic marker to study the taxonomic position of lagomorpha in monophyletic clade glires [ 30 , 31 ] . . . .
. . . indicates that this specimen is currently accessioned as o . thibetana thibetana in the amnh collection . the forresti specimen used in ye et al . ( 2011 ) , was never accessioned into a collection and has been subsequently lost . to test potential morphological differences between l . melainus and sympatric l . mandshuricus , measurements of l . melainus and l . mandshuricus published by li and luo ( 1979 ) and luo ( 1988 ) were analyzed . . . .
. . . the 4 th group also from the same geographical area reported by hayata and shimba ( 1969 ) ; vorontsov and ivanitskaya ( 1973 ) which having the chromosome number 2n = 40 to 42 . the 3 rd group reported from qinghai tibet plateau having the chromosome number 2n = 46 to 54 reported by vorontsov and ivanitskaya ( 1973 ) ; ivanitskaya ( 1978 ) ; tan and bai ( 1987 ) ; ye et al . ( 2011 ) etc . the second group is having chromosome number 2n = 60 to 62 from the surrounding himalayan areas ( nadler et al . , 1969 ; vorontsov and ivanitskaya , 1973 ; capanna et al . , 1991 ; puget and berland , 2008 ) . . . .
reconstruction of karyotype evolution in core glires . i . the genome homology revealed by comparative chromosome painting
karyotypic evolution of the family sciuridae : inferences from the genome organizations of ground squ . . .
cross - species chromosome painting has made a great contribution to our understanding of the evolution of karyotypes and genome organizations of mammals . several recent papers of comparative painting between tree and flying squirrels have shed some light on the evolution of the family sciuridae and the order rodentia . in the present study we have extended the comparative painting to the . . . [ show full abstract ]
reconstruction of karyotype evolution in core glires . i . the genome homology revealed by comparative . . .
glires represent a eutherian clade consisting of rodents and lagomorphs ( hares , rabbits , and pikas ) . chromosome evolution of glires is known to have variable rates in different groups : from slowly evolving lagomorphs and squirrels to extremely rapidly evolving muroids . previous interordinal homology maps between slowly evolving glires were based on comparison with humans . here , we used sets of . . . [ show full abstract ]
karyotype evolution of giraffes ( giraffa camelopardalis ) revealed by cross - species chromosome painti . . .
considering the giraffe ( giraffa camelopardalis , gca , 2n = 30 ) as a primitive species , its comparative genomic data are critical for our understanding of the karyotype evolution of pecorans . here , we have established genome - wide chromosomal homologies between giraffe , chinese muntjac ( muntiacus reevesi , mre , 2n = 46 ) and human ( homo sapiens , hsa , 2n = 46 ) with whole sets of chromosome - specific . . . [ show full abstract ]
the number of rodent species examined by modern comparative genomic approaches , particularly chromosome painting , is limited . the use of human whole - chromosome painting probes to detect regions of homology in the karyotypes of the rodent index species , the mouse and rat , has been hindered by the highly rearranged nature of their genomes . in contrast , recent studies have demonstrated that . . . [ show full abstract ]


sigaus childi is an endangered protected species of grasshopper known only from the alexandra district of the south island of new zealand .

it is one of just two species of grasshopper listed for protection under the new zealand wildlife act 1953 ( the other is brachaspis robustus ) .

the genus sigaus is endemic to new zealand .

sigaus childi

dowle , morgan - richards & trewick . 2014 . bmc evol . bio . 14 ( 216 ) > > sigaus childi
the genus sigaus is endemic to new zealand and with all but one species endemic to the south island . there are nine species of sigaus .
sigaus childi was described as a new species in 1999 . it is closely related to s . minutus from the mackenzie basin . there are 16 species of grasshopper in nz .
jamieson , c . 1999 . new zealand j . zool . 26 ( 1 ) : 44 > > sigaus childi urn : lsid : orthoptera . speciesfile . org : taxonname : 58218
pca traditional species diagnostics . principle component analysis using morphological character states used in traditional species diagnostics of adult sigaus grasshoppers from south island new zealand . morphologically , sigaus childi is readily separated from s . australis , f = females ( triangles ) ; m = males ( squares ) .
discriminant analysis with cross - validation using the character states employed in traditional species diagnosis for adult grasshoppers of the sigaus australis complex in south island , new zealand . sigaus childi ( in bold ) individuals were correctly grouped together . the total squared difference between the two groups was 274 .
sample map . sample locations in south island , new zealand , of the sigaus australis complex grasshoppers used in this study . the two main species sigaus australis ( green ) and sigaus childi ( pink ) are morphologically very different ; s . childi tends to be smaller and more camouflaged to its local habitat than s . australis . the ‘central group’ and ‘area of sympatry’ defined here are used to analyse subsets of the specimens . hatched circles represent locations with both species present .
jamieson cd : distribution and abundance of sigaus childi jamieson ( orthoptera : acrididae ) , a central otago endemic grasshopper . science for conservation 110 . 1999 , department of conservation , wellington , new zealand
gene flow is traditionally considered a limitation to speciation because selection is required to counter the homogenising effect of allele exchange . here we report on two sympatric short - horned grasshoppers species in the south island of new zealand ; one ( sigaus australis ) widespread and the other ( sigaus childi ) a narrow endemic .
gene flow is traditionally considered a limitation to speciation because selection is required to counter the homogenising effect of allele exchange . here we report on two sympatric short - horned grasshoppers species in the south island of new zealand ; one ( sigaus australis ) widespread and the other ( sigaus childi ) a narrow endemic .
snp loci . ( a ) frequency distribution of locus specific f st values for each of the 74 snp loci sampled between the grasshoppers sigaus childi and sigaus australis in sympatry . ( b ) bayescan plot of 74 snp loci with a single marker ( log ( po ) > 1 ) showing slight departure from neutrality ; the vertical line is the 5 % po threshold of false discovery .
morris sj . 2003 . two new species of sigaus from fiordland , new zealand ( orthoptera : acrididae ) . the new zealand entomologist , 26 : 65 - 74
jamieson cd . 1999 . a new species of sigaus from alexandra , new zealand ( orthoptera : acrididae ) . new zealand journal of zoology , 26 : 43 - 48
minimum spanning networks . minimum spanning network of four of the six mtdna ( coi 519 bp ) haplogroups ( figure a ) within the sigaus australis grasshopper complex , show sharing ( grey ) of identical haplotypes between s . childi ( white ) and s . australis ( black ) . haplotype spot size is proportional to number of individuals with a particular haplotype and branch length estimates nucleotide differences .
extensive gene flow between the grasshoppers sigaus australis 1 and s . childi 2 in sympatry was revealed using migrate - n with 74 rad - seq snp makers . theta ? is an estimate of population size , ? = 4n e ? in the snps , where n e is population size and ? is mutation rate , population size was generally large as is expected for grasshoppers . mutation scaled migration rates ( m ) were converted into nm ( number of migrants per generation ) via ? 1 m 2 - > 1 = 4 nm 1 . the results show extensive gene flow in both directions .
as with the mtdna data we sought evidence of genetic structure concordant with taxonomy and morphology using analysis of the correlation of genotypes between species by grouping the samples according to the morpho - species s . australis and s . childi and estimating f ct . no significant genetic differentiation between the morpho - species was found within the central group : f ct = 0 . 01780 ( p ? = ? 0 . 18573 ) , or the area of sympatry f ct = ? 0 . 07424 ( p ? = ? 0 . 65494 ) , although these samples were not all taken from the same generation . this result was consistent with the inference from structure .
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dowle ej 1 , 2 , morgan - richards m 3 , trewick sa 4 .
ecology group , iae , massey university , private bag 11222 , palmerston north , 4442 , new zealand . eddy . dowle @ cawthron . org . nz .
coastal and freshwater group , cawthron institute , nelson , new zealand . eddy . dowle @ cawthron . org . nz .
ecology group , iae , massey university , private bag 11222 , palmerston north , 4442 , new zealand . m . morgan - richards @ massey . ac . nz .
ecology group , iae , massey university , private bag 11222 , palmerston north , 4442 , new zealand . s . trewick @ massey . ac . nz .
of the 79 putatively neutral markers ( mtdna , microsatellite loci , its sequences and rad - seq snps ) all but one marker we examined showed extensive allele sharing , and similar or identical allele frequencies in the two species where they co - occur . we found no genetic evidence of deviation from random mating in the region of sympatry . however , analysis of morphological and geometric traits revealed no evidence of morphological introgression .
based on phenotype the two species are clearly distinct , but their genotypes thus far reveal no divergence . the best explanation for this is that some loci associated with the distinguishing morphological characters are under strong selection , but exchange of neutral loci is occurring freely between the two species . although it is easier to define species as requiring a barrier between them , a dynamic model that accommodates gene flow is a biologically more reasonable explanation for these grasshoppers .
pmid : 25318347 pmcid : pmc4219001 doi : 10 . 1186 / s12862 - 014 - 0216 - x
alternative hypotheses . alternative hypotheses to explain the relationship between morphological differentiation and gene flow in this study of grasshoppers in new zealand . two morphologically defined species exist that have a common ancestor and may share identical alleles due to decent , but as distinct species derived alleles are also expected . ( a ) an abrupt speciation event is expected to result in accumulation of distinct traits and genotypes , including at neutral loci . ( b ) sharing of neutral genetic alleles might be maintained by ongoing gene flow whilst alleles at some loci are subject to selection . morphological difference is an observable expression of genetic loci under diverging selection . ( c ) hybridisation or reticulation is expected to result in individuals with intermediate forms .
references : batchelor cl . 196 ) . preliminary observations of alpine grasshoppers ina habitat modified by deer and chamois . proceedings of the new zealand ecological society , 14 : 15 - 26
bigelow rs . 1967 . the grasshoppers of new zealand , their taxonomy and distribution . university of canterbury publications , christchurch , new zealand .
dowle ej , morgan - richards m . & trewick sa . 2014 . morphological differentiation despite gene flow in an endangered grasshopper . bmc evolutionary biology , 14 : 216 - 231
eades dc , otte d , cigliano mm . & braun h . 2014 . family acrididae macleay , 1821 . orthoptera species file . version 5 . 0 / 5 . 0 . retrieved on : december 8th , 2014 : urltoken
hutton fw . 1897 . the grasshoppers and locusts of new zealand and the kermadec islands . proceedings and transactions of the new zealand institute , 30 : 135 - 150
hudson l . 1970 . identification of the immature stages of new zealand apine acridid grasshoppers ( orthoptera ) . transactions of the royal society of new zealand : biological sciences , 12 : 185 - 20
key khl . & colless dh . 1993 . a higher classification of the australian acridoidea ( orthoptera ) . ii . * subfamily cantanopinae . invertebrate systematics , 7 : 89 - 111
salmon jt . 1950 . a new species of acrididae ( insecta : orthoptera ) from new zealand . transactions of the royal society of new zealand , 78 : 69
trewick sa . 2001 . identity of an endangered grasshopper ( acrididae : brachaspis ) : taxonomy , molecules and conservation . conservation genetics , 2 : 233 - 243
trewick sa & morris s . 2008 . diversity and taxonomic status of some new zealand grasshoppers . doc research and development series no . 290 , department of conservation , wellington , new zealand , 40p .
trewick sa . 2008 . dna barcoding is not enough : mismatch of taxonomy and genealogy in new zealand grasshoppers ( orthoptera : acrididae ) . cladistics 24 : 240 - 254 .
von wattenwyl kb . & fea l . 1893 . révision du système des orthoptères et description des espèces rapportées par m . leonardo fea de birmanie . vol . 33 . tipografia dei r . istituto sordo - muti .
genetic introgression occurs when two genetically distinct populations come into contact enabling individuals from each to interbreed . when this occurs through secondary contact the process has usually been regarded as hybridisation [
] . the fertility of resulting offspring mediates gene flow between populations . this situation underpins the popular biological species concept [
] . here we report on flightless new zealand short - horned grasshoppers ( orthoptera : acrididae ) . most of the fifteen new zealand species , in four endemic genera , occupy subalpine native grasslands above the tree line [
are relatively large ( adult females ~ 26 mm ) and abundant in south island subalpine grasslands between 1000 and 1800 m asl . sympatric with this widespread species is the microendemic
is also present in this region , but the two species are readily distinguished by their appearance . an intriguing feature of
is that their colour patterns appear to be specific to the substrate on which individuals are found . colour patterns within
) that grows on rocks in some areas of central otago . although , inferences of camouflage are subjective they support the conjecture that these grasshoppers are under selection by visual predators .
are more boldly patterned , often with longitudinal stripes , and tend to be colour - pattern variable within locations .
] . or perhaps divergence has occurred and been maintained despite gene flow . genetic exchange between populations might be experienced at different rates across the genome ; selection could operate on some loci to limit local exchange of alleles even when net ( genome wide ) gene flow continues . these alternatives make different predictions about the pattern of morphological and genetic character sharing ( figure
in order to understand the evolution of this system we applied six types of data ; morphology , mtdna sequencing , microsatellite genotyping , multi - copy nuclear sequencing , single nucleotide polymorphisms ( snp ) and spatial position . we used these putatively independent data to contrast species integrity as characterised by morphology ( subject to natural selection ) and neutral characters that allowed us to test the stability of species delimitation , assess the extent and evenness of gene flow and thus gain an understanding of where these grasshopper populations are in the speciation continuum .
) . the first four components of the pca accounted for > 95 % of the variation . thus the morphometric data based on traditional taxonomic characters suggest there are just two morphological entities :
geometric analysis of the grasshopper pronotum gave a similar result to that of traditional morphology . pca analysis of the 14 landmark measurements showed two major groupings ( figure
was likely due to their extremely cryptic shape formed by the ‘broken’ edge of the pronotum resulting in little uniformity within species . the
grasshopper pronotum shape ( pca analysis from morphoj ) : ( i ) pca for both species from all areas , with pronotum shape changes indicated along the pc1 axis . the two major groupings comprise
) , with pronotum shape changes indicated along the pc1 axis . within the area of sympatry there was no clear evidence of morphological intermediates .
neighbour joining tree of mtdna haplotypes ( coi , 519 bp ) and a distribution map showing the spatial distribution of haplogroups . the circled clades ( dashed circles ) are used in the network analysis ( figure
? < ? 0 . 001 t - square 1291 . 4541 ) but there was no overlap between the two forms . thus we found no evidence of morphological intermediates in adults or juveniles .
b ) . the mtdna clusters did not correspond to current taxonomic groups or morphological types . mtdna sequences from
ancestry . the high genetic diversity detected at alexandra appears to be a result of the meeting of three distinct mtdna clades that otherwise have separate ranges .
( black ) . haplotype spot size is proportional to number of individuals with a particular haplotype and branch length estimates nucleotide differences .
? 0 . 1410 ) and the power of this test was limited by smaller within - clade sample sizes . there was no statistical support for population genetic differentiation between
the three microsatellite loci surveyed each had between 16 and 18 alleles . no evidence of linkage disequilibrium was detected and hardy - weinberg expectations were met in the majority of population samples . a positive relationship between geographic distance and genetic differentiation ( pairwise f
] showed evidence of extensive gene flow among populations . the optimum ? k was k = 2 ( figure
c ) . there was no support for k = 3 , which was unexpected given that the data encompassed two morphologically distinct species and spatial structure had already been indicated . it is important to note that although the microsatellite dataset covers a similar geographical range to that of the mtdna dataset there is little similarity in the genetic structure detected . to reduce the possible influence of uneven sample size of the two species we restricted the data to include sampling only from the area of sympatry ( figure
] analysis found no support for genetic partitioning within these data ( i . e . k = 1 ) , contrary to the expectation that the two morphologically defined species would represent discrete genetic units ( figure
a ) . a test for deviations from expected frequencies of neutral loci in bayescan indicated that one marker may have been subject to diversifying selection , log ( po ) > 0 alpha 0 . 878 ( figure
b ) . a blast search of the sequence containing this snp did not result in any matches to known sequences on genbank . mean population pairwise f
was low ( 0 . 025 ) , with a confidence interval that effectively included zero ( ci 2 . 5 % 0 . 001 , ci 97 . 5 % 0 . 053 ) , providing little evidence that these samples represent more than one population , with random mating . population differentiation estimated with structure suggested extensive sharing of genetic material among populations , with no species structure detected ( figure
has a comparatively wide geographic range that can be subdivided into a number of phylogeographically distinct mtdna haplogroups .
in terms of habitat , geographic range and genetic diversity ; in stark contrast to its clear morphological distinction . despite occuring in sympatry , no phenotypic intermediates were detected . we found no evidence of genetic partitioning in putatively - neutral mtdna sequence , microsatellite , its sequence loci or snp data . none of the mtdna diversity detected within
suggesting recent ( and on - going ) gene flow . the snp data show no population structure and extensive gene flow between the two species in sympatry , with one marker showing some sign of diversifying selection . the presence of more than one its1 - its2 sequence within the genomes of single grasshoppers is also consistent with recent gene flow near the township of alexandra . concerted evolution normally results in homogenisation of variation in the rrna cassette containing its [
generated by different local environmental conditions . extreme plasticity is known to occur in other grasshoppers such as the locust (
] , but despite altitudinal separated from subalpine areas , these habitats are climatically similar in terms of their extreme day / night and seasonal temperature cycles . more pertinent is the fact that
are sympatric in the lowland semiarid environment of central otago . they occur at the same places at the same times with , for example , specimens of both morphotypes used in our analysis collected as adults within metres of each other in little valley , alexandra on the same day . these circumstances are inconsistent with an interpretation of phenotypic differences being driven by environmental induced plasticity . although we cannot exclude this possibility , a novel type of micro - environmental control of grasshopper development would need to be invoked .
] , however the sharing of identical mtdna haplotypes and sharing of alleles across neutral nuclear loci in these two species suggests that very recent and / or ongoing reticulation is more likely . gene flow is expected to homogenise variation between species and it has traditionally been considered that speciation is unlikely to proceed in the presence of gene flow . many models of speciation have emphasized partitioning of populations by some extrinsic process ( e . g . allopatry ) as a prerequisite [
] . however , if selection on particular loci is sufficiently intense , the effects of gene flow could be mitigated . models that accommodate permeability of putative species boundaries and acknowledge that selection can be locus - specific rather than genome wide are not new [
, is a small , highly cryptic species of grasshopper suggesting it has been or is under selection from visual predators . in contrast ,
limited our sample sizes and precluded any observation of mating behaviour and juvenile colouration and survival where the species are sympatric . further examination might reveal morphological intermediates .
] . expansion of grass and herbs following reduction of forest that may previously have formed a habitat barrier between the alpine and lowland grasshoppers may have facilitated population mixing . the area now shared by
was further modified by european introduction of plants and grazing animals , and mining practices in the last 150 years . these changes could have facilitated increased gene flow , but were analogous to the effects of pleistocene climate cycling . disentangling their respective influence on the grasshoppers is not simple [
the taxonomic status of these species is problematic , as traditional methods cannot resolve the conflicting information from morphology and genetics resulting from the process of evolution . although
it is morphologically well differentiated , and in our relatively small snp dataset we were able to find one marker possibly under selection . models of speciation with gene - flow predict a continuum from partially isolated populations to reproductive isolation [
do not appear to be losing morphological distinction , but our neutral genetic data does show extensive gene flow . the absence of any morphological hybrids suggests selection is intense , removing relatively conspicuous intermediates and holding these two species apart . this may provide them the opportunity to diverge at other loci .
] . there is a grand irony that while for many , genetic methods are seen as tools for testing species status ( e . g . dna barcoding ) , genetic data are actually the key to revealing that speciation is not clear cut [
] . in our study we found that the only characters that reliably distinguished species were morphological , whilst 78 neutral genetic markers showed that distinct morphotypes do not correspond to genetically isolated units .
we collected grasshoppers by hand when they were active during the new zealand summer season ( december - march , between 1995 and 2009 ) . sampling included all recognised members of the
is widespread in subalpine habitat with a few populations extending down to low elevation ( ~ 300 m asl ) areas in some locations .
, sample sizes were limited and sampling spanned more than one overlapping generation ( c . f . usual assumptions of population genetic models ) . sampling from different generations , which are already overlapping is , however , not likely to increase the similarity of population allele frequencies , and therefore we do not consider this will have hindered any of the analyses .
specimens were confirmed by simon morris ( pers comm . to sat ) . individuals were preserved by freezing or in 95 % ethanol and identified following bigelow [
complex grasshoppers in two ways . the first used the traditional species diagnostic characteristics , although we note that much of the information used to distinguish some of these taxa has been geographic location and altitude [
] . male genitalia are taxonomically informative for some grasshopper species but consistent differences have not been reported among species in this complex . for instance , male genitalia in
adults were distinguished by the tegmina concealing the relictual hind wing , which is the case only in the last instar . juveniles were excluded from this morphometric analysis . the data were analysed using discriminant analysis and principle component analysis ( pca ) approaches implemented in minitab 15 [
. a pca was applied to all the morphological characters and the scores saved . pca required no
grouping , allowing us to determine whether the data could be partitioned into taxonomically meaningful groups based solely on the documented morphological character states .
as an alternative to the traditional taxonomic characters , we tested for shape differences of the pronotum among species using geometric analysis . this method is more powerful and avoids any circularity that could arise from using traditional species characteristics as the sole morphological traits analysed . much of the taxonomy in the
) that were obtained with the aid of a dissecting microscope we tested whether shape variation could be detected from metric data . using imagej [
] , 14 landmarks were identified around the perimeter of the dorsal surface of the pronotum on each image of each grasshopper and measured . the landmarks were selected to maximise variation among individuals . these measurements were analysed using morphoj [
] . a procrustes fit aligned by principal axes was performed to eliminate size differences before a procrustes anova was used to examine the error of image capture . this analysis revealed that the error arising from image capture variation was biologically irrelevant : mean squares for image capture was 32 times smaller than the variation found between individual grasshoppers . juveniles , adults and both sexes were included in the analyses and tested to confirm they did not partition in the results . principal component and discriminant analyses with cross validation were each preformed on the averaged value for each individual from all four species ( 34 ?
] . dna from specimens preserved for more than one year , was extracted using incubation at 55°c with proteinase k and a ctab buffer ( 2 % hexadecyltrimethylammonium bromide , 100 mmol / l tris - hcl ph8 . 0 , 1 . 4 mol / l nacl , 20 mmol / l edta ) , followed by a combined phenol / chloroform / isoamyl alcohol ( 25 : 24 : 1 ) cleanup . extractions were eluted in water and diluted as necessary for pcr reactions . primers c1 - j - 2195 and li - n - 3014 [
] were used to target the 3 ' portion of coi . polymerase chain reactions ( pcrs ) were performed in 10 μl volumes using abgene red hot taq ( thermo fisher scientific ) . thermocycling conditions were 94°c for three minutes ; 94°c for 45 seconds , 52°c for 45 seconds and 72°c for 75 seconds repeated 36 times ; followed by a 2 minute final extension . cycle sequencing used perkin elmer bigdye 3 . 1 chemistry following the manufacturer ' s protocols analysed on an abi prism 377 dna sequencer ( applied biosystems , inc . , foster city , california ) . sequences were checked using sequencher version 4 . 10 . 1 ( gene codes ) and aligned with existing data using seal version 2 . 0 and geneious pro version 5 . 3 . 4 [
geneious was used to estimate a neighbour - joining tree for all the lineages . network version 4 . 5 . 1 . 6 . [
] was used to estimate haplotype networks within clades . to test for a correlation between genetic and geographic distance ( expected under a model of isolation by distance ) , mantel tests [
] with 10 , 000 randomizations to assess the significance of distance correlations . distance by distance analysis was applied to all data and separately to data within haplo - groups .
screening of fifty microsatellite loci revealed three that were polymorphic and amplified consistently among a subset of dna samples from the target taxa . the loci were checked for large allele dropout , stuttering , and null alleles using 1000 randomisations in microchecker version 2 . 2 . 3 [
populations were treated as a single population for this purpose . although there was evidence of null alleles in some of the loci within some of the populations (
) . the analyses were run using an admixture model with correlated allele frequency , 100 , 000 generations of burn - in followed by 100 , 000 generations , and the number of groups ( k ) set from 1 to 20 ( 10 replicates each ) . the optimum value of k was found using the ? k method except for k = 1 , which was determined by examination of the bar - plots and structure harvester [
nuclear sequences representing the internal transcribed spacers ( its1 and its2 ) of the rrna cluster and the intervening rrna 5 . 8s gene were obtained using the primers its4 and its5 [
] . pcr conditions and sequencing followed standard protocols as above . sequences were aligned using geneious pro version 5 . 3 . 4 [
] and checked by eye . sequences were generated for all grasshoppers from the area of sympatry of
( the alexandra region ) . alignment and comparison of unambiguous with ambiguous sequences allowed us to identify the most likely combinations of sequences that gave the observed heterozygotes ( s3 ) . where sequence variants differed by single nucleotide substitutions we could identify and resolve the polymorphism . where sequence variation involved indels the resulting length polymorphism was evident by abrupt onset of sustained nucleotide ambiguity at the indel position , but sequencing in both directions allowed identification of the combination of sequences involved . to examine the number of families of its per grasshopper genome we interrogated a dna dataset generated by high throughput sequencing . genomic dna from a single
individual was sequenced on an illumina hi - seq 2000 ( beijing genomics institute ) resulting in > 1gb of sequence . the sequence was de - novo assembled via velvet [
] and the results viewed in tablet version 1 . 12 . 09 . 03 [
] . the resulting contigs were blasted to genbank and all matches to its were selected , aligned , mapped back and , checked for copy number .
single nucleotide polymorphic ( snp ) anonymous nuclear markers were generated using high throughput sequencing , with individual dna fragments coded so we could identify individual grasshopper genotypes . the double digest rad - seq protocol [
] was applied with minor modifications . we estimated genome size to help us optimise the selection of endonucleases and sequencing coverage . to do this we used flow - cytometry on a facscalibur system and cellquest software ( bd biosciences , san jose , ca , usa ) , following staining of cells with propidium iodide and reference to an internal control ( chicken or locust ) . our estimates of the
genome were approximately 11 . 9 pg ( consistent with estimates of other short - horned grasshopper species (
data were generated using an illumina hi - seq ( new zealand genomics limited ) , and sorted using the stacks version 0 . 99992 pipeline [
] . settings for coverage and sites per read were adjusted iteratively . read coverage settings vary in the literature [
] , so we initially ran trials ranging from 7 to 30 reads , but found no alteration in the results . we report results using an optimum coverage of 15 reads per individual ( excluding all stacks with a lower coverage ) , a maximum of two mismatches between reads for a single individual as well as allowing four mismatches between primary and secondary reads within ustacks . we allowed the program to remove any potentially spurious highly repetitive stacks . in cstacks we allowed 3 mismatches between samples when generating the snp set ( - m 15 - n2 - m4 - n3 - t ) . we restricted our analysis to a single snp per putative locus ( always the first ) , thus avoiding potential problems of non - independence between markers . data file conversion for programs was performed using pgdspider version 2 . 0 . 4 . 0 [
as calculated for each putative - locus across all loci in stacks , and an amova was run in genodive version 2 . 0b24 [
] was used to estimate population differentiation using an admixture model with correlated allele frequency . a burnin of 100 , 000 generations was followed by 10 replications of 100 , 000 generations with the number of groups ( k ) set from 1 to 3 . the optimum value of k was found from ? k method , via structure harvester , except for k = 1 , which was determined by examination of the bar - plots [
gene flow between the two populations in sympatry was estimated using migrate - n version 3 . 5 . 1 [
] , although algorithms that test for gene flow are often not ideal for situations where gene flow is very high , which is likely in this case . migrate - n was implemented with the bayesian inference strategy . initial runs involved only half the markers as we optimised settings . the starting values for ? and m were generated initially from f
with subsequent runs using the resulting ? and m values . the uniform prior distributions were used for both parameters with slice sampling ; one long chain was run recording every 5 steps after a burnin of 50 , 000 with a static heating scheme with five chains . four runs were conducted on half the data before three final runs on all loci were undertaken using the starting values for ? and m of the previous run . bayescan version 2 . 01 [
] was used to examine the individual markers for evidence of selection using the default settings . prior odds of a neutral model were 10 times more likely than the model with selection at a locus . this prior was tested further by changing it to one , without any identifiable change in results . the alpha value was used to determine the direction of selection with a positive value suggesting diversifying selection and a negative value suggesting balancing selection . results were viewed in r version 3 . 0 . 0 using an fdr of 0 . 05 [
assembly is available on genbank ( km576254 ) . its data indicating snp and indel variation is available in additional file
ejd , sat and mmr conceived and designed experiments and collected grasshoppers . ejd and sat carried out molecular analysis . ejd carried out morphometric analysis . ejd , sat and mmr drafted and edited paper . all authors read and approved the final manuscript .
additional file 1 : extended methods . further details on methodology . includes details on the microsatellite discovery methodology , along with primer sequences . as well as details on sample number / location at each analysis . as well as a table indicating snp and indel variations / positions in its sequencing . ( docx 146 kb )
additional file 2 : geometric morphometrics . procrustes fit values for geometric morphometrics . ( txt 94 kb )
additional file 3 : snp and microsatellites . spreadsheet of snps and microsatellites values across populations used in this study . ( xlsx 71 kb )
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