galaxiella munda is a species of fish in the galaxiidae family .

it is endemic to inland waters of southwest western australia , and known as the western mud minnow .

in 1999 the mud minnow was not included in the list of threatened species under the environment protection and biodiversity conservation act , but is listed as near threatened on the iucn red list .

galaxiella munda

mean genetic divergences between populations of galaxiella munda for cytochrome b calculated using p - distances .
information on the western mud minnow ( galaxiella munda ) is currently being researched and written and will appear here shortly .
martin f . gomon & dianne j . bray , galaxiella munda in fishes of australia , accessed 10 jul 2018 , urltoken
biology of galaxiella munda mcdowall ( teleostei : galaxudae ) , including a comparison of the reproductive strategies of this and three other local species .
western dwarf galaxias , galaxiella munda . source : stephen beatty / freshwater fish group & fish health unit , murdoch university . license : all rights reserved
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wager , r . 1996 . galaxiella munda . in : iucn 2011 . iucn red list of threatened species . version 2011 . 2 . < urltoken > . downloaded on 18 may 2011 .
berra , t . m . & g . r . allen . 1989 . clarification of the difference between galaxiella nigrostriata ( shipway , 1953 ) and galaxiella munda mcdowall , 1978 ( pisces : galaxiidae ) from western australia . rec . west . aust . mus . 14 : 293–297 .
berra , t . m . & g . r . allen . 1989 . clarification of the difference between galaxiella nigrostriata ( shipway , 1953 ) and galaxiella munda mcdowall , 1978 ( pisces : galaxiidae ) from western australia . rec . west . aust . mus . 14 : 293–297 .
mean genetic divergences between populations of galaxiella nigrostriata for cytochrome b calculated using p - distances .
mean genetic divergences between western lineage populations of galaxiella pusilla for cytochrome b calculated using p - distances .
mean genetic divergences between eastern lineage populations of galaxiella pusilla for cytochrome b calculated using p - distances .
molecular phylogeny and phylogeography of the australian freshwater fish genus galaxiella , with an emphasis on dwarf galaxias ( g . pusilla )
molecular phylogeny and phylogeography of the australian freshwater fish genus galaxiella , with an emphasis on dwarf galaxias ( g . pusilla )
male blackstriped dwarf galaxias , galaxiella nigrostriata . source : gerald r . allen / western australian museum . license : all rights reserved
galaxiella munda mcdowall 1978 , j . r . soc . n . z . 8 ( 1 ) : 119 , figs . 1 - 2 . type locality : pool in jarrah forest , about 8 km north of scott river , between nanup and augusta , 34°13 ' s , 115°29 ' e , western australia .
pen , l . j . , gill , h . s . , humphries , p . & potter , i . c . 1993 . biology of the black - stripe minnow galaxiella nigrostriata , including comparisons with the other two galaxiella species . journal of fish biology 43 : 847–863 . urltoken
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gill , h . s . & neira , f . j . 1994 . larval descriptions of three galaxiid fishes endemic to south - western australia : galaxias occidentalis , galaxiella mund a and galaxiella nigrostriata ( salmoniformes : galaxiidae ) . marine and freshwater research 45 : 1307–1317 . doi : 10 . 1071 / mf9941307
saddlier s , jackson j , hammer m . melbourne : state of victoria department of sustainability and environment ; 2010 . national recovery plan for the dwarf galaxias galaxiella pusilla .
wager , r . 1996 . galaxiella nigrostriat a . in : iucn 2012 . iucn red list of threatened species . version 2012 . 2 . . downloaded on 14 may 2012 .
pen , l . j . , gill , h . s . , potter , i . c . & humphries , p . ( 1993 ) . growth , age composition , reproductive biology and diet of the black - stripe minnow galaxiella nigrostriata ( shipway ) , including comparisons with the other two galaxiella species . journal of fish biology 43 : 847 - 864 .
galaxiella munda and g . nigrostriata show even more extreme examples of population differentiation . overall though , patterns were broadly similar to those in g . pusilla . some proximate populations had low divergences ( e . g . , sites 32–33 , 34 and 36 , 38–39 ) . populations 24–25 were unusual in that they are separated by considerable distance , yet they were genetically quite similar ( fig . 3 and table s5 ) . most other populations had high genetic divergences , typically from 0 . 7 % to 3 . 0 % ( tables s4 , s5 ) . interestingly , despite being more common in elevations closer to sea level , g . nigrostriata displayed a higher degree of genetic differentiation than g . munda ( table s1 ) .
gill , h . s . & morgan , d . l . 1996 . threatened fishes of the world : galaxiella nigrostriata ( shipway , 1953 ) ( galaxiidae ) . environmental biology of fishes 47 : 344 .
galeotti dm , mccullough cd , lund ma ( 2010 ) black - stripe minnow galaxiella nigrostriata ( shipway 1953 ) ( pisces : galaxiidae ) , a review and discussion . j r soc west aust 93 : 13–20 .
summary of pairwise comparisons of allele frequency between sites for galaxiella pusilla west . site 9 was excluded from the analysis due to small sample size and lack of a geographically - proximate neighbour . format as for table s8 .
colour varies in relation to the water inhabited . in tannin - stained waters g . munda is dark brown on the back and sides with a silvery - white belly . may also be greenish olive dorsally with a longitudinal silvery stripe , a brownish - orange stripe ( usually during the breeding season ) and a narrow dark stripe with a silvery belly .
smith , k . d . 1996 . an outlying population of galaxiella nigrostriata ( pisces : galaxiidae ) at melaleuca park , western australia : causes of habitat fragmentation . honours thesis , department of zoology , the university of western australia . 115 pp .
fish fact : the muddy , or western mud minnow ( galaxiella munda ) is the closest rellie to the black - strip minnow , but instead of being found in ephemeral waterbodies , it is found in spring - fed headwater streams that are the result of groundwater expression . take the yaragadee as an example , these species is most secure in yaragadee aquifer - fed streams . at least we aren ' t drawing that aquifer down - phew . the species has a one year life - cycle so if something negative happens in their streams then they are likely to go kaput . big worry considering our recent year of climate change .
galeotti , d . m . 2013 . metapopulation theory explains black - stripe minnow ( pisces : galaxiidae , galaxiella nigrostriata ) distribution in seasonal wetlands in south - west western australia . msc thesis , edith cowan university , perth , western australia , 165 pp . urltoken pdf
galeotti , d . m . , m . a . castalanelli , d . m . groth , c . mccullough & m . lund . 2014 . genotypic and morphological variation between galaxiella nigrostriata ( galaxiidae ) populations : implications for conservation . marine and freshwater research urltoken abstract
thompson gg , withers pc ( 1999 ) the metabolic response to hypoxia and emersion of aestivating fishes ( lepidogalaxias salamandroides and galaxiella nigrostriata ) and a non - aestivating fish ( bostockia porosa ) from south - western australia . australian journal of zoology 47 : 295 - 305 .
gill , h . s . & morgan , d . l . 2003 . ontogenetic changes in the diet of galaxiella nigrostriat a ( shipway , 1953 ) ( galaxiidae ) and lepidogalaxias salamandroides mees , 1961 ( lepidogalaxiidae ) . ecology of freshwater fish 12 : 151 - 158 .
the ml analyses of each gene recovered a nearly identical interspecific topology , with the only incongruence being among brachygalaxias relationships . cyt b placed b . gothei and b . bullocki as sister species , whereas s7 placed b . gothei in between the two b . bullocki lineages . bootstrap support was high ( > 98 % ) for all deep nodes in the cyt b dataset , with the exception of the node uniting g . munda and g . nigrostriata ( 52 % ; fig . 2 ) . our tcs networks provided a more detailed pattern of the relationships between cyt b haplotypes within each galaxiella species , highlighting the large number of nucleotide differences between many populations ( fig . 3 ) .
citation : unmack pj , bagley jc , adams m , hammer mp , johnson jb ( 2012 ) molecular phylogeny and phylogeography of the australian freshwater fish genus galaxiella , with an emphasis on dwarf galaxias ( g . pusilla ) . plos one 7 ( 6 ) : e38433 . urltoken
galeotti , d . m . , mccullough , c . d . & lund , m . a . 2010 . black - stripe minnow galaxiella nigrostriata ( shipway 1953 ) ( pisces : galaxiidae ) , a review and discussion . journal of the royal society of western australia 93 : 13–20 .
maximum likelihood trees for galaxiella based on analysis of cytochrome b ( a ) and s7 ( b ) sequences and a neighbour joining tree for allozymes ( c ) . panel a shows the complete tree with operational taxonomic units ( otus ) for each galaxiella species collapsed . expanded trees within each species are shown below , with the lower scale bar applying to each of the four subtrees . bootstrap values are based on 1000 pseudoreplicates . trees are rooted with brachygalaxias . each otu code is based on the sampling location number and name in table 1 and fig . 1 , while the coloured symbols match figs . 1 , 3 and 5 .
unmack pj , bagley jc , adams m , hammer mp , johnson jb ( 2012 ) molecular phylogeny and phylogeography of the australian freshwater fish genus galaxiella , with an emphasis on dwarf galaxias ( g . pusilla ) . plos one 7 ( 6 ) : e38433 . doi : 10 . 1371 / journal . pone . 0038433
smith , k . d . , pen , l . j . & knott , b . 2002 . genetic and morphological study of the black - stripe minnow , galaxiella nigrostriata ( salmoniformes : galaxiidae ) , including a disjunct population near perth , western australia . records of the western australian museum 21 : 285–290 . pdf open access
knott , b . , jasinska , e . j . & smith k . d . 2002 . limnology and aquatic fauna of epp 173 , melaleuca park , refuge for an outlier population of the black - stripe minnow galaxiella nigrostriata ( pisces : galaxiidae ) in southwestern australia . records of the western australian museum 21 : 291 - 298 .
unmack pj , bagley jc , adams m , hammer mp , johnson jb ( 2012 ) molecular phylogeny and phylogeography of the australian freshwater fish genus galaxiella , with an emphasis on dwarf galaxias ( g . pusilla ) . plos one 7 ( 6 ) : e38433 . pdf open access doi : 10 . 1371 / journal . pone . 0038433
smith , k . d . , knott , b . & jasinska , e . j . 2002 . biology of the black - stripe minnow , galaxiella nigrostriata ( shipway ) , in an acidic , black - water lake in melaleuca park near perth , western australia . records of the western australian museum 21 : 277 - 284 . pdf available open access
galeotti , d . m . , mccullough , c . d . & lund , m . a . 2008 . current state of knowledge of the black - stripe minnow galaxiella nigrostriata ( pisces : galaxiidae ) in western australia . edith cowan university , centre for ecosystem management report 2008 - 12 , unpublished report to kemerton silica sands pty . ltd . , perth , western australia . 36 pp
occur in slow - running , tea - colored streams usually in sandy areas . also found in swamps , small ponds and roadside ditches . also lives in the vegetated shallows of some freshwater lakes . water is typically acidic ( ph 4 . 5 - 6 . 5 ) and darkly tannin - stained . an inhabitant of temporary waters , capable of aestivating in damp bottom sediments over summer . spawning habits similar to g . munda . most individuals perish after their first spawning ( ref . 44894 ) . feed on small insects , larvae of aquatic insects and micro - crustaceans . breeding is associated with winter rains . males take on a bright color pattern at the start of the breeding season ( ref . 33842 ) .
refer to table 1 for corresponding locality details . each species is represented by a different symbol , with populations from both lineages of g . pusilla color coded to match figs . 2 , 3 and 5 . shaded areas refer to the general distribution of each galaxiella species . low sea level drainage patterns are shown to the minus 135 m bathymetric contour . bathymetry predicts a large depression that we refer to as lake bass . the small inset of australia shows the relevant biogeographic provinces , and the black central southern region represents the extent of the eucla basin .
summary of genetic diversity of cytochrome b sequences sampled within galaxiella pusilla populations and clades in this study . locality numbers , the number of individuals sampled , number of haplotypes ( h ) , haplotype diversity ( hd ) with its standard deviations , nucleotide diversity ( π ) with its standard deviations , and the results of coalescent simulations of three neutrality statistics , tajima’s d , fu’s f s , ramos - onsins and rozas’ r 2 , are shown . within - population values only represent genetically polymorphic sites ; unlisted populations were monomorphic ( h = 1 ; hd = π = 0 . 000 ) .
summary of pairwise comparisons of allele frequency between sites for galaxiella pusilla east . due to their small sample sizes , sites 16 and 20 were pooled with a geographically - proximate neighbour ( sites 15 and 21 respectively ) . lower triangle = number of loci displaying statistically - significant differences in allele frequency ( p < 0 . 05 after bonferroni correction ) . upper triangle = results of determining p - value for each population pair across all loci ( fisher’s method ) after bonferroni correction ; * * * = p < 0 . 001 , * * = p < 0 . 01 ; ns = not significant .
allozyme frequencies at all variable loci for the 22 sites surveyed for galaxiella pusilla . site codes follow table 1 . frequencies of all but the rarer / rarest alleles are expressed as percentages and shown as superscripts ( allowing the frequency of each rare allele to be calculated by subtraction from 100 % ) . maximum sample sizes per site are shown in brackets ( asterisks indicate sample sizes of n = 3 for the designated sites and loci ) . a dash indicates insufficient enzyme activity at this locus . invariant loci : ald1 * , ald2 * , enol * , gapd1 , glo * , gp , gpi2 , pgam * , and pk * .
a very small galaxiid found creeks and streams of south - west western australia .
adults are brownish - grey above with a white belly , and have several whitish blotches along the upper side , and a broad coppery - brown stripe edged in fine olive brown stripes just below the midline from behind the eye to the tail base .
video of western dwarf galaxias by the freshwater fish group of murdoch university , western australia .
endemic to temperate freshwaters of south western australia , from the albany district ( 118°10’e ) to north of perth ( 31°50’s ) .
the western dwarf galaxias prefers swift - flowing streams in karri forests , near submerged vegetation . the water in these streams is usually acidic ( ph 3 . 0 - 6 . 0 ) and darkly tannin - stained , and the water temperature fluctuates widely with the seasons . it also occasionally occurs in ponds , swamps and roadside drains .
dorsal fin 7 - 10 ( usually 8 - 9 ) ; anal fin 11 - 15 ( usually 12 - 14 ) ; pectoral fin 9 - 12 ( usually 10 - 11 ) ; pelvic fin 5 - 7 ( usually 6 ) ; gill rakers16 - 20 .
body elongate , slender , almost tubular ; body depth at anus 10 . 5 - 14 . 3 % sl ; head short , snout short and bluntly rounded ; mouth small , oblique , reaching to front of eyes ; jaws equal in size with no enlarged canine teeth ; eye large , upper margin at head profile ; gill rakers long and slender ; pyloric caeca absent ; compressed behind the vent ; caudal peduncle longer than deep ; scales absent .
fins small and membranous ; low membranous fold along caudal peduncle almost to dorsal and anal rays dorsal fin short - based ; dorsal fin origin behind anal fin origin ; caudal fin long , truncate to rounded ; pectoral fins narrow - based , small and rounded , inserted high laterally ; ventral fin very small , inserted at mid - body .
carnivore - feeds on small insects , aquatic insect larvae and microcrustaceans - usually near vegetation at the edges of streams .
the western dwarf galaxias has a year - long life cycle that is completed entirely in freshwater . the sexes are separate and fertilisation is external . spawning occurs between june and october , with a peak during august and september . females spawn several times , depositing several batches of eggs in flooded vegetation over several weeks . adults die soon after spawning .
allen , g . r . 1989 freshwater fishes of australia . t . f . h . publications , inc . , neptune city , new jersey .
allen , g . r . , midgley , s . h . & allen , m . 2002 . field guide to the freshwater fishes of australia . perth : western australian museum 394 pp .
gill , h . s . & f . j . neira 1998 galaxiidae ( galaxiinae ) : southern galaxias . p . 70 - 77 . in f . j . neira , a . g . miskiewicz & t . trnski ( eds . ) larvae of temperate australian fishes : laboratory guide for larval fish identification . university of western australia press . 474 p .
mcdowall , r . m . ( 1978 ) a new genus and species of galaxiid fish from australia ( salmoniformes : galaxiidae ) . j . r . soc . n . z . 8 ( 1 ) : 115 - 124 .
mcdowall rm , frankenburg rs ( 1981 ) the galaxiid fishes of australia . rec aust mus 33 : 443–605 .
mcdowall , r . m . & j . m . waters . 2004 . phylogenetic relationships in a small group of diminutive galaxiid fishes and the evolution of sexual dimorphism . journal of the royal society of new zealand 34 ( 1 ) : 23 - 57 .
merrick , j . r . & g . e . schmida 1984 australian freshwater fishes : biology and management . griffin press ltd . , south australia . 409 p .
morgan , d . , gill , h . & potter , i . ( 1996 ) . the distribution of freshwater fish in the southwestern corner of australia . report to water and rivers commission . waters and rivers commission , perth , western australia .
morgan , d . l . , gill , h . s . & potter , i . c . ( 1998 ) . distribution , identification and biology of freshwater fishes in south - western australia . records of the western australian museum . supplement no 56 , perth , western australia .
paxton , j . r . , d . f . hoese , g . r . allen & j . e . hanley 1989 zoological catalogue of australia . volume 7 . pisces . petromyzontidae to carangidae . australian government publishing service , canberra . zoological catalogue of australia . volume 7 . v . 7 : i - xii + 1 - 665 .
greek , galaxias , ou = a kind of fish , diminutive ( ref . 45335 )
oceania : endemic to australia . known only from the southwestern part in coastal streams between albany and ellen brook .
maturity : l m 4 . 7 range ? - ? cm max length : 6 . 0 cm sl male / unsexed ; ( ref . 44894 ) ; common length : 3 . 5 cm sl male / unsexed ; ( ref . 5259 )
dorsal spines ( total ) : 0 ; dorsal soft rays ( total ) : 6 - 8 ; anal spines : 0 ; anal soft rays : 9 - 12 ; vertebrae : 38 - 43 . adults are brown - grey dorsally and white ventrally with several whitish blotches near the dorsal midline , and a broad brown stripe just below midline from behind eye to caudal - fin base .
allen , g . r . , 1989 . freshwater fishes of australia . t . f . h . publications , inc . , neptune city , new jersey . ( ref . 5259 )
phylogenetic diversity index ( ref . 82805 ) : pd 50 = 0 . 5625 [ uniqueness , from 0 . 5 = low to 2 . 0 = high ] .
bayesian length - weight : a = 0 . 00389 ( 0 . 00180 - 0 . 00842 ) , b = 3 . 12 ( 2 . 94 - 3 . 30 ) , in cm total length , based on all lwr estimates for this body shape ( ref . 93245 ) .
trophic level ( ref . 69278 ) : 3 . 2 ±0 . 36 se ; based on food items .
resilience ( ref . 69278 ) : medium , minimum population doubling time 1 . 4 - 4 . 4 years ( k = 3 . 70 ; tm = 1 ; tmax = 1 ; mean fec = 65 ) .
vulnerability ( ref . 59153 ) : low vulnerability ( 14 of 100 ) .
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peter j . unmack , 1 , 2 , * justin c . bagley , 2 mark adams , 3 , 4 michael p . hammer , 3 , 5 and jerald b . johnson 2 , 6
conceived and designed the experiments : pju mph jbj . performed the experiments : pju ma . analyzed the data : pju jcb ma . contributed reagents / materials / analysis tools : pju mph ma jcb jbj . wrote the paper : pju jcb ma .
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southern australia provides an excellent geological setting for studying biogeographic patterns . long - term aridity since the oligocene created a vast desert region in southern australia , isolating two moist temperate regions in the southwestern and southeastern parts of the continent . the biota that occupy these two regions have been isolated since at least mid - miocene (
for corresponding locality details . each species is represented by a different symbol , with populations from both lineages of
color coded to match figs . 2 , 3 and 5 . shaded areas refer to the general distribution of each
species . low sea level drainage patterns are shown to the minus 135 m bathymetric contour . bathymetry predicts a large depression that we refer to as lake bass . the small inset of australia shows the relevant biogeographic provinces , and the black central southern region represents the extent of the eucla basin .
species is differences in the width of the continental shelf . regions with narrow continental shelf should present limited opportunities for drainage connections during low sea levels , since few rivers are likely to connect (
) , whereas a broader continental shelf allows a greater area for rivers to traverse and potentially interconnect . much of southern australia has a relatively narrow continental shelf , thus most drainages remain isolated during low sea levels (
are widespread across southern australia . as presently defined , the genus consists of three species of small , stout - bodied galaxiid fishes
is usually found in smaller creeks and can occur further inland . both species overlap to a limited extent in floodplain habitats that have seasonal connections to more permanent water bodies
, a more integrative approach is needed ; one that blends phylogeny and phylogeography . although obviously relevant , previous single - and multi - locus phylogenetic analyses of
used limited within - taxon sampling ( one sample / species ) . previous within - species comparisons in the two western
employed good sampling with 10 individuals from each population but included only three populations outside of victoria . here , we improve on previous studies of the biogeography and population genetics of
fishes . first , we use gondwanan fragmentation and the well known geology of southern australia to derive geologically - based calibration points for analyses and thereby estimate a time frame for diversification within the genus based on broader within - taxon sampling . this amounts to a ‘congeneric phylogeographical’ sampling approach . this has been shown to present a viable means of increasing systematic accuracy and improving historical inferences , by the recovery of polyphyly and ‘cryptic’ lineages , within vertebrate species
species based on more extensive geographic sampling . third , we investigate phylogeographic patterns within each
species in southern australia . specifically , if shelf width is important then populations of
in the western portion of their range in southeastern australia will have a high level of genetic divergence between drainages . in contrast , populations of
permission to undertake field work and collect specimens was obtained under the following permits : victorian fisheries research permit rp 581 , victorian flora and fauna permit 10002072 , victorian national parks permit 10002154 , tasmanian inland fisheries service permit 2003 / 12 , tasmanian department of primary industries , water and environment permit tfa 03106 , south australian primary industries and resources - section 59 exemption , western australian license to take fauna for scientific purposes sf006928 . specimens were obtained under arizona state university institutional animal care and use committee ( iacuc ) approval 09 - 1018r , brigham young university iacuc approval 070403 and university of adelaide animal ethics committee approval s - 32 - 2002 .
the rarity and conservation status of these species limited our sample sizes . in many cases , there were only one or a few populations known per river basin . our primary goal was to include one population per major drainage where each species is found . we examined samples of
were provided by david galeotti ( edith cowan university ) . fishes were collected with seine and dip nets and either frozen whole in liquid nitrogen or preserved in 95 % ethanol in the field . our sampling ranged from 1–11 individuals per locality , with most sites represented by 10 individuals within
, plus two sequences from genbank for a total of 219 ingroup samples . based on their close relationships to
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. the location column gives the general location of each sample . station code refers to the field number or sama ebu code that samples are catalogued as . the last three columns provide the number of individuals ( n ) examined for each marker . abbreviations : ck , creek ; hwy , highway ; r , river ; rd , road ; sa , southern australia ; tas , tasmania ; vic , victoria ; wa , western australia .
we employed a gis approach to quantify several environmental factors needed to evaluate our hypotheses linked to changes in sea level . datasets used to generate maps ( e . g . ,
and manipulated in arcinfo and arcmap version 10 ( environmental systems research institute , redlands , ca ) . bathymetric data were obtained from a 30 arc - second ( ca . 1 km ) dataset gebco 08 ( september 2010 release ,
we extracted genomic dna from muscle tissue for each specimen using dneasy tissue kits ( qiagen inc . , chatsworth , ca ) . we amplified the mtdna cytochrome
. when this failed to produce sufficient pcr product , the gene was amplified in two halves using glu21 - hd . gp ggrttgtttgagcctgtytcgt and gp . 505
we also used the forward primers glu18 taaccaggactaatgrcttgaa or glu31 tgrcttgaaaaaccaccgttgt with hd . gp as well as the internal forward primer sal . 484 caatgaatttgagggggrttctc or gn . 484
. we also included the first two introns and the second exon of the nuclear s7 gene . all nuclear sequences were obtained by nested pcr using the following primers : 1f
in the second reaction . in a few cases , we had to use the internal primers 1f . 2 and 2r . 67 tacctgggarattccagactc , and 2f . 2
. the first of these nested reactions were 25 µl . final concentrations for polymerase chain reaction ( pcr ) components per 25 µl reaction were as follows : 25 ng template dna , 0 . 25 µm of each primer , 0 . 625 units of taq dna polymerase , 0 . 1 mm of each dntp , 2 . 5 µl of 10x reaction buffer and 2 . 5 mm mgcl
. data files containing all individuals sequenced as well as various analysis files were deposited in dryad , doi : 10 . 5061 / dryad . c3g8h .
to reconstruct a species tree and to date clade divergences and coalescence times to the most recent common ancestor . we combined the data from cyt
and s7 , included all individuals sequenced , then ran analyses using a relaxed molecular clock in beast 1 . 6 . 1
. the full datasets for each gene were imported into beauti 1 . 7 ( prerelease ) , which we used to generate the input file for beast . we incorporated two calibrations based on geological events . the first was the root of the tree , representing the split between
, as potentially congruent with the separation of australia , antarctica and south america . the terrestrial separation of australia and antarctica occurred at approximately 52 ma
. three separate analyses were conducted using both calibration points in the same analysis , plus one analysis with each calibration used individually . analyses were also conducted excluding the sequence data to check that posterior distributions were not heavily driven soley by our priors rather than the sequence data . we used an uncorrelated lognormal relaxed molecular clock based upon a lognormal prior using the ‘speciation birth death’ process . both lognormal prior calibrations had a mean of 1 . 0 , with standard deviations of 1 . 5 for the calibration of 52 ma and 1 . 0 for the 14 ma calibration . the best - fitting model of molecular evolution for each gene ( cyt
: gtr + i + g ; s7 : gtr + g ) was estimated via aic in modeltest . analyses were run for 200 million generations , with parameters logged every 10 , 000 generations . multiple runs were conducted to check for stationarity and that independent runs were converging on a similar result . output from beast was examined in tracer 1 . 5 with 10 % burn - in and the tree results were summarized using treeannotator 1 . 7 ( prerelease ) .
we used multiple methods to examine within and between population genetic patterns in order to investigate the role of sea level changes in structuring genetic variation . we explored patterns of genetic variation in
population and clade , and for all populations combined . to test the relative contribution of genetic variation to within and between population structure in
. if genetic structuring is driven by isolation among drainages due to drainage basin boundaries , then we expect to see high levels of structuring among drainages . cases where high amounts of variation can be explained between groups suggest an important historic barrier to gene flow exists coincident with the structure of the model . to evaluate movement between drainage basins , we combined populations into 15 groups based on drainage basins or close geographic proximity ( i . e . , the following populations were combined : 1–4 , 11–12 , 13–14 , 15–16 , 17–18 ;
since frozen tissues were available for virtually all g . pusilla , we were able to obtain comparative allozyme profiles for this species . these were used to independently assess the finding by coleman et al . [ 35 ] of a major east - west dichotomy , and provide more appropriate molecular data for assessing whether any dichotomy reflected the presence of cryptic species . three specimens of an outgroup species , galaxias olidus , were also included in this analysis . we conducted allozyme electrophoresis of muscle homogenates on cellulose acetate gels ( cellogel™ , milan , italy ) following richardson et al . [ 66 ] . details of all enzyme and locus abbreviations , electrophoretic conditions , and stain recipes are contained in richardson et al . [ 66 ] or wallman and adams [ 67 ] , while the allozyme nomenclature follows hammer et al . [ 68 ] .
the genetic relationships among all sites were initially assessed by constructing a neighbour - joining ( nj ) tree from a pairwise matrix of nei’s [ 69 ] unbiased genetic distances ( nei d ) , following the methodology of hammer et al . [ 10 ] . we then used genepop 4 . 0 [ 70 ] to assess the raw allozyme genotypes for statistically - significant departures from hardy - weinberg expectations , for evidence of linkage disequilibrium at each individual site ( for n > 4 ) , and to detect statistically significant differences in allele frequency among sites within each of the two major lineages evident within g . pusilla . threshold probability values were adjusted for the use of multiple tests using sequential bonferroni correction [ 71 ] . finally , we employed the multivariate ordination technique principal coordinates analysis ( pco ) to visually assess the genetic affinities among all individuals within each lineage ( details in [ 10 ] ) .
our molecular datasets provided clear discrimination among all described species . in addition , they revealed major and concordant genetic discontinuities consistent with cryptic speciation in g . pusilla as per coleman et al . [ 35 ] .
) . removal of fish with identical haplotypes within populations reduced the dataset to 66 individuals . the number of individuals from a population with a given haplotype is provided after the population name in
. based on this reduced dataset , 680 characters were invariant , 28 characters were variable but parsimony uninformative and 433 characters were parsimony informative . both
codon , while the latter also had a premature stop codon in the third - to - last codon position ( codon 378 ) . neither change is unusual for cyt
among fishes ( p . j . unmack , pers . obs . ) or across various animal groups
) and was obtained via whole mtdna amplification , which minimizes the chance of amplifying nuclear pseudogene copies ( masaki miya , pers . comm . ) . maximum likelihood recovered one tree for cyt
( a ) and s7 ( b ) sequences and a neighbour joining tree for allozymes ( c ) . panel a shows the complete tree with operational taxonomic units ( otus ) for each
species collapsed . expanded trees within each species are shown below , with the lower scale bar applying to each of the four subtrees . bootstrap values are based on 1000 pseudoreplicates . trees are rooted with
) . inclusion of individual alleles from 14 heterozygous individuals increased the dataset to 66 sequences . the number of heterozygous positions within an individual varied from one to three ( all heterozygous individuals were phased into individual alleles prior to ml analyses ) . based on this dataset , 535 characters were invariant , 10 characters were variable but parsimony uninformative and 354 characters were parsimony informative . maximum likelihood recovered one tree for s7 with a likelihood score of −3378 . 210082 (
( a ) is color - coded relative to figs . 1 , 2 and 5 . circle size represents haplotype abundance ; the key to circle size is in the center of panel a . the ancestral haplotype in each network is indicated by a box . haplotype counts are given in parentheses when multiple populations share the same haplotype . haplotype labels consist of the population number and name from
) , with bayesian posterior probabilities equal to 1 . 0 for all internal nodes . divergence dating results obtained using * beast are presented in
. running the analysis with the same settings , but without data confirmed that our input settings actually produced the desired prior probability distributions on our calibrated nodes and that our data were responsible for our results rather than our priors . most statistics from all three analyses had ess scores > 400 , demonstrating the chain was well sampled . results were substantially different depending on the calibration used . when the single calibration of 14 ma was used , all dates were much younger (
the first line presents results based on both calibration points of 52 and 14 million years , while the second and third rows represent results under each individual calibration . the mean and 95 % highest posterior densities are given for each node ( in millions of years ) , and we report the per lineage mean rate of evolution per million years for each gene in the last two columns .
± one standard deviation ( s . d . ) = 0 . 212±0 . 090 ; mean π×100± s . d . = 0 . 059±0 . 014 ) , but moderate genetic diversity within each clade (
) . results from our amova analyses found high levels of among - group structure consistent with the effects of strong isolation between populations . when all groups ( n = 15 ) were compared , among - group comparisons accounted for 95 . 8 % of the genetic variation . within eastern
( n = 7 ) among - group comparisons accounted for 90 . 2 and 72 . 8 % of the variation respectively . all three among - group amova values were statistically significant ( p < 0 . 05 ) .
< 0 . 0001 ) , supporting demographic - spatial expansion . neutrality tests showed a similar pattern across
. comparing bsps to constant demographic models supported bsp patterns . bsp models provided a better fit to the data within each
mean posterior n e estimates for each species ( darker lines ) are bounded by upper and lower 95 % highest posterior densities ( g . pusilla west , pink shading ; g . pusilla east , blue shading ) . the x - axis represents units of time in thousands of years ago ( ka ) , scaled according to posterior mutation rates estimated in beast . the y - axis shows estimated population size in hundreds of thousands , calculated assuming a g . pusilla generation time equal to 1 . 0 . the fish shown is a male g . pusilla west ( by mph ) .
. no site displayed any statistical evidence of either linkage disequilibrium or of genotype frequencies being incompatible with hardy weinberg expectations . these data unequivocally recovered the same two primary clades (
) ; lineage e1 comprised victorian sites 11–18 , while lineage e2 consisted of the two most southerly victorian sites ( 15 and 16 ) plus all tasmanian sites ( sites 19–22 ) . these two lineages were diagnosable by near - fixed differences at two loci (
) , as well as by major differences in allele frequency ( δp > 40 % ) at three other loci . lineage e2 was further divisible into two sub - lineages e2a ( sites 15 , 16 , 19 , 22 ) and e2b ( sites 20 , 21 ) , diagnosable by a fully - fixed difference at a single locus (
) plus major differences in allele frequency at three other loci . importantly , this phylogeographic structure is concordant with the cyt
west ( b ) individuals . relative pco scores are plotted for the first and second dimensions , which individually explain 41 % and 10 % ( a ) and 25 % and 17 % ( b ) respectively of the total multivariate variation present . individuals are colour - coded relative to
west also revealed statistically significant differences in allele frequency , usually involving multiple loci , in all but four instances ( sites 6 vs . 7 , 1 vs . 2 , 1 vs . 3 , 2 vs . 4 ;
) . here lineage w2 consisted of the most easterly site ( site 10 ) , while lineage w1 comprised all other sites . additional phylogeographic structure was detectable within the widespread lineage w1 , although only one of the three sub - lineages ( w1b = site 5 ) was allozymically well - differentiated from the other two sublineages ( w1a = four western sites , w1c = four eastern sites ) by major differences in allele frequency at multiple loci (
based on a single individual per taxon , although their dataset included two additional genes ( rag1 and 16 s ) and morphological characters . our results , showing a deep divergence between
west using a suite of non - hypervariable nuclear genetic markers ( and therefore more suitable for assessing species boundaries ) , were also consistent with the conclusions of coleman et al .
based on mtdna coi and nuclear microsatellites . we therefore refer to these lineages hereafter as distinct candidate species . our results were strongly supported , with s7 and the combined analysis having high bootstrap values ( 100 and 1 . 0 respectively ) for all between species relationships . high support from the cyt
our divergence estimates varied greatly depending on whether we used the separation of australian and south american species ( min . age set to 52 ma ) or the east - west separation between
species ( min . age set to 14 ma ) . essentially , analyses using the older calibration point recovered mean ages slightly more than double those recovered using the younger calibration (
) . when both calibrations were included , estimates were up to ∼10 % lower , especially on the upper 95 % highest posterior distribution ( hpd ) . while we present results from all three calibration combinations , we favour the results obtained using a single calibration of 52 ma . the separation of southeastern and southwestern australia by the formation of the nullarbor plain (
may have separated long before that event . indeed , it has already been suggested that east - west separation of some groups predates this event by potentially tens of millions of years
. we are more confident that continental drift was responsible for their separation than some form of oceanic dispersal . neither
have any proclivity for marine environments today , nor do they possess any traits that imply a potentially diadromous life cycle . it is certainly not impossible that they were once diadromous , although this implies a substantial convergence in morphological and ecological characteristics , perhaps driven by adaptation to similar habitats . our age estimates based on continental drift are similar to burridge et al .
, they obtained a slightly older mean age than our analysis ( 58 vs . 55 . 8 ma ) . however , their credibility interval was broader ( 45–72 vs . 52–66 ma ) , probably due to their much larger dataset with multiple calibration points . the remaining divergences between each
were all younger than our results , with their mean ages usually being similar to our lowest 95 % credibility interval values . in contrast , the age for both
species was similar in both studies . it is also important to recognise that our estimate of 52 ma for the separation of australian and south american clades is only a minimum estimate . it is quite likely that the true divergence would have occurred earlier . this implies that all of our age estimates would be underestimated if the continental separation is an appropriate calibration point .
species across southern australia at a mean age of 34 . 3 ma ( 95 % hpd of 24 . 5–44 . 5 ma ,
are adapted to exploit more ephemeral environments , they might be expected to persist under more arid conditions . however , increasing aridity would also increase the likelihood of extirpation , given
species are more reliant on ephemeral habitats i . e . , seasonally wet and dry on an annual basis . as aridity intensified across southern australia
, these ephemeral habitats would be less reliably rewatered , thus eliminating populations through extended droughts .
was estimated to have a mean age of 22 . 5 ma ( 95 % hpd of 15 . 2–30 . 5 ma ,
adapted to survive in temporary habitats . alternatively , if aestivation was ancestral , then
evolved the ability to compete and survive in permanent habitats . it seems likely that temporary habitats like those used by
species had a mean age of 5 . 7 ma ( 95 % hpd of 2 . 2–9 . 1 ma ,
, with populations from mount emu creek / hopkins river representing the western species . this contrasts strongly with patterns in other co - distributed fishes , where species - level separations or western range limits occur at the eastern side of the lake bass drainage
, at least within freshwater fishes . no other fish species have divergences of this scale ( i . e . , species level differentiation ) at this location , although within - species genetic breaks are present in several groups
, consistent with this boundary representing a barrier that is not frequently crossed by aquatic organisms . one major exception to the separation on the western edge of lake bass is population 10 from the barwon river , which is thought to have become part of the lake bass drainage during low sea levels (
species at first seems perplexing , as it runs counter to predictions from current drainage patterns . based on the unique signature found at all molecular markers examined here (
, it seems unlikely that population 10 represents a recent translocation . instead , this outcome infers that the geomorphic history of the barwon river basin differs somewhat from that of adjacent basins . while several authors
, the lake expansion hypothesis needs to be re - examined geomorphically now that more accurate elevation and geological mapping is available . clearly though , evidence does support some faunal exchange via lake corangamite .
and allozyme analyses were broadly congruent , although they differed in details . the extent of within - species genetic divergence differed between species as well as markers ; for cyt
had an average within - group genetic divergence of 0 . 8 % , while the western
populations relative to the western species ( 90 . 2 % vs . 72 . 8 % respectively ) . five groups of populations had shared or closely related cyt
( p - distance of 0 . 5 % ) ; however allozymes grouped sites 6–9 together (
populations , which only differed from each other based on allele frequencies . coleman et al .
against expectations , we found that neither inferred phylogenetic patterns , nor patterns of genetic structuring , corresponded very closely to predictions based on estimated low sea level drainage patterns . we predicted that populations 10–16 , plus 19–20 would essentially show a low degree of genetic structure , since hydrological models predict that they would have been part of a larger continuous drainage system during historical periods of low sea level (
) . lastly , populations 1–9 were expected to show relatively higher levels of genetic divergence . although the latter pattern was indeed evident based on cyt
) . one extreme example is the relationship between sites 11–14 and 17–18 , which are quite geographically distant . based on low sea level drainage patterns , these populations should not have any lowland connections (