tenuibranchiurus is a genus of diminutive freshwater crayfish that live in the australian state of queensland .

only one species has been described , the swamp crayfish , t. glypticus .

t. glypticus is reportedly the smallest species of crayfish in the world .

it is distinguished from other crayfish by its small size , adults being only around 25 millimetres ( 1.0 in ) long , and its claws which open vertically rather than horizontally or obliquely .

tenuibranchiurus lives in coastal wallum swamps , and stays among the sedges rather than in more open water .

its habitat is highly fragmented , as land is used for the expansion of brisbane and the sunshine coast , and tenuibranchiurus glypticus is therefore listed as an endangered species on the iucn red list .

additional populations have been found at the periphery of its range , but these are thought to represent new , undescribed species .

tenuibranchiurus

existing and putative species identified within tenuibranchiurus and the newly proposed gen . nov . .
rt @ thepeerj : a novel genus and cryptic species harboured within the monotypic freshwater crayfish genus tenuibranchiurus riek https : / / t . co…
number of tenuibranchiurus specimens analysed for each gene fragment from each of the sampled localities , as well as outgroup sequences included ( see table s1 for sequence details ) .
based on our results , the genus tenuibranchiurus is represented only by specimens collected from queensland . as such , tenuibranchiurus glypticus remains a valid species and it is represented by populations grouped with samples from the type locality . five new putative species were identified within tenuibranchiurus ( table 8 ) . specimens collected from new south wales belong to a putative newly - proposed genus with two new putative species ( table 8 ) . until a formal description is completed , the new genus will be referred to as gen . nov .
although genetic diversity within tenuibranchiurus has previously been reported , no quantification of this diversity had been undertaken . the multi - gene approach taken by this study and use of several different analytical methods has identified not only several putative species within the formerly monotypic tenuibranchiurus , but a new genus with two putative species . although species identification of freshwater crayfish has traditionally been made through morphological methods , the use of molecular methods in this study allowed the potential pitfalls of plastic and / or cryptic morphological forms within crayfish to be avoided , and will contribute in the development of a standardised method for dealing with species identification within other freshwater crayfish .
there is no population information available for this species . tenuibranchiurus spp . are extremely rare and this species appears to have been extirpated at two of the sites it was known from ( one of which was developed for housing ) and one of the two remaining sites is at victoria point ( davie 2007 ) , a heavily urbanized area with some light industry on the outskirts of brisbane ( j . coughran , k . l . dawkins and j . m . furse pers . comm . 2008 ) .
a total of 127 tenuibranchiurus samples were sequenced for the coi gene fragment , 59 for 16s , 95 for gapdh , 58 for h3 , and 47 for ak ( table 2 ) . additional specimens from the genera gramastacus , geocharax , engaeus , engaewa , and cherax were also sequenced for inclusion as outgroups . where sequences from these outgroups could not be obtained ( i . e . , due to non - amplification ) , alternative sequences were retrieved from genbank ( details in table s1 ) . sequences obtained in this study were deposited in genbank under accession numbers kx669691 – kx670093 , kx753349 .
this species has a maximum length of 25 mm , and is the smallest known species of crayfish found in australia . it inhabits coastal wallum swamps and is found among sedges rather than in more open water ( harding and williamson 2003 ) . there is very little suitable habitat remaining within its range , as most has been cleared for the development of brisbane and the sunshine coast ( j . coughran , k . l . dawkins and j . m . furse pers . comm . 2008 ) . the habitat of other tenuibranchiurus species has recently been recorded ( dawkins et al . in prep ) and the biology of this genus is poorly understood .
two types of analyses were used to obtain a best - estimate of the species - level lineages present within tenuibranchiurus ; namely , groupings identified through use of a concatenated alignment phylogeny ( referred to henceforth as the ‘combined gene tree’ ) , and intra - versus inter - cluster variation through ϕ st analysis . a combined gene tree analysis was chosen over individual gene trees because , although preliminary phylogenetic analyses performed on the individual gene regions suggested that there were multiple genetic groups within t . glypticus , statistical support was not always strong for all genes . therefore , in order to increase the strength of the phylogenetic signal , and thus support for branching patterns , the five gene alignments were concatenated and analysed as a single data set for phylogenetic reconstructions .
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livingston , f . , livingston , f . , soulsby , a . - m . , batchelor , a . , dyer , e . , whitton , f . , milligan , h . t . , smith , j . , lutz , m . l . , de silva , r . , mcguinness , s . , kasthala , g . , jopling , b . , sullivan , k . & cryer , g .
has been assessed as endangered using criteria b1ab ( iii , iv ) . this species has an estimated extent of occurrence ( eoo ) of 1 , 700 km
and is only known from two locations . there is an ongoing decline in both the eoo of this species and the quality of habitat , with very little suitable habitat remaining . this species has strict habitat requirements which are being compromised by urbanisation , industry , pollution and salt water intrusion . the area in which this species is found is one of rapid human population growth ( davie 2007 ) . site protection for this species is urgently needed as this species faces an extremely high risk of extinction .
. the species was described from specimens collected in brisbane ( mt . gravatt ) and caloundra ( riek 1951 ) . however , the species has not been recorded in metropolitan
to the northeast ( harding and williamson 2003 ) . recent research ( dawkins 2008 , dawkins
to caloundra , a distance of approximately 100 km north - south . this species has an estimated extent of occurrence of 1 , 700 km
. however , this area has been very heavily developed ( i . e . includes
) and the actual area of occupancy is likely to be a small fraction of this . this species is now only known from two sites ( j . coughran , k . l . dawkins and j . m . furse pers . comm . 2008 ) .
there are no species specific conservation measures in place for t . glypticus . research should be urgently initiated to establish the species’ area of occupancy , and should also include population assessment and monitoring , biological and life history information , habitat requirements ( including salinity tolerance ) , investigations into thermal tolerance and resilience to exotic species . in addition , genetic and / or morphological analysis should be undertaken to elucidate any distinction between populations from the two extant localities , victoria point ( on the mainland ) and bribie island ( j . coughran , k . l . dawkins , j . m . furse pers . comm . 2008 ) .
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reportedly world ' s smallest crayfish , being fully grown at 25 mm . unlike other crayflish in south - east queensland , fingers of claws open and close vertically rather than horizontally or obliquely . body greyish - brown . difficult to find due to small size , cryptic colouration and well - developed burrowing habits .
paperbark swamps and shallow drainage channels . prefers to burrow into damp clay but is occasionally found in peaty sand . woodgate , qld , south to at least southern brisbane area .
originally recorded from bulimba creek , mt gravatt , it has not been recorded in metropolitan brisbane for more than 60 years , though it has been found at victoria point . habitat loss is probably a significant threat to the survival of this unique species .
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species are the fundamental unit of biodiversity , yet there has always been disagreement about criteria by which they should be recognised and the methods by which they should be delineated , with no general consensus reached thus far . the lack of one clearly accepted definition of a “species” creates obvious limitations , as what one person regards as a species may not be regarded as being so by another person , which is often further exacerbated by differences of opinion between fields of study . employing the general lineage concept ( glc ; de queiroz , 1998 ) , where a species is defined as a metapopulation lineage evolving separately from other lineages , somewhat unites the various species concepts by allowing any evidence of lineage separation ( and thus any property emphasised by the alternative concepts ) to be used as evidence for species delimitation ( de queiroz , 2007 ) . not only does this concept allow multiple lines of evidence to be used , but it also allows the evolutionary processes that have caused divergence between lineages to be examined .
once a species tree has been inferred , additional testing is often undertaken to provide support for the proposed species’ groups . a range of statistical analyses are available for testing species boundaries and , as there is currently no universally accepted way to define species , there are also a range of critiques on these methods ( e . g . , blaxter , 2004 ; brower , 1999 ; ebach & holdrege , 2005 ; lipscomb , platnick & wheeler , 2003 ; seberg et al . , 2003 ; sites & marshall , 2003 ; sneath & sokal , 1973 ; tautz et al . , 2002 ; tautz et al . , 2003 ; wiens & penkrot , 2002 ; wiens & servedio , 2000 ; will , mischler & wheeler , 2005 ; yang & rannala , 2010 ) . under the glc , any evidence of lineage separation can be evidence for the existence of different species ( de queiroz , 2007 ) ; as such , the identification of numerous corroborating lines of evidence ( through the use of multiple tests ) can be seen as lending support to any species boundaries that are defined . therefore , although no single test is currently universally accepted , the apparent need to choose a particular method is circumvented by using a selection of techniques and multiple gene regions as , under the glc , concordance between multiple lines of evidence is seen as increasing the rigour of species delimitation .
the triangle and bolded name denotes the type locality . grey lines denote drainage boundaries , and the black line denotes the border between queensland and new south wales . refer to table 2 for collection details .
a total of 133 specimens were collected across 16 field localities , including the type locality for t . glypticus . all specimens from this study were collected under permits witk08599510 , wisp08599610 , and twb / 01 / 2011 issued by the department of environment and resource management . dna was extracted from specimens preserved in 70 % ethanol using a variation of the cetyltrimethyl ammonium bromide / phenol - chloroform extraction protocol ( doyle & doyle , 1987 ) . two mitochondrial gene regions ( cytochrome oxidase subunit 1 ( coi ) and 16s rdna ( 16s ) ) and three nuclear gene regions ( glyceraldehyde - 3 - phosphate dehydrogenase ( gapdh ) , histone - 3 ( h3 ) , and arginine kinase ( ak ) ) were amplified ( see table 1 for primer list ) . sequences were edited using sequencher 4 . 9 ( genecodes , 2009 ) and aligned using the muscle addition in mega5 ( edgar , 2004 ) . alignments were then checked and edited by hand if necessary .
preliminary analyses of both individual and combined gene trees showed a prominent separation between qld and nsw populations . in light of this , genetic distances between qld and nsw populations , distances between these two groups and the outgroups , and distances between the outgroups were calculated using both coi and 16s data to compare the degree of separation . these distances were calculated in mega5 ( tamura et al . , 2011 ) using the net between group mean distances with 1 , 000 bootstrap replicates ( gamma distribution with shape parameter = 1 , maximum composite likelihood ( mcl ) model ; positions containing gaps and missing data were eliminated ) .
an analysis of molecular variance ( amova ) was used to calculate variation within and among clusters of sequences , as implemented in arlequin v . 3 . 1 ( excoffier , laval & schneider , 2005 ) . to determine what the most likely lineages were , the clades identified by the combined gene tree analysis , as well as additional splits evident within the tree that were deemed to plausibly represent lineages , were also tested , as well as groups based on the geographic division of populations ( i . e . , collection locality ; see table s2 ) . the amova calculates three statistics ; ϕ st , ϕ sc , and ϕ ct , all of which are based on both the haplotype frequency and genetic divergence . ϕ st measures variation among all populations , and ϕ sc measures variation among populations within groups , and ϕ ct estimates variation among groups . it has been suggested that an f ct value > 0 . 95 can represent evidence for accurate species groupings ( i . e . , > 95 % of the genetic variation can be attributed to differences among groups ) ( monaghan et al . , 2005 ) . using the ϕ ct estimate as a surrogate for f ct ( as this estimate includes genetic divergence as well as haplotype frequency ) , this can provide an approach to delineate taxa based on population genetic analyses by interpreting the amova results used to calculate intra - versus inter - cluster variation in a way analogous to f - statistics ( wright , 1978 ) . the criterion to determine the appropriate number of lineages using this method is where an increase in the number of suggested lineages does not appreciably increase the ϕ ct estimate for those lineages .
in order to validate the lineages that were identified using the species discovery approaches for species - status , two methods were used ; barcoding gap identification ( sensu hebert et al . , 2004 ) , and the k ∕θ method ( sensu birky , 2013 ) . these two methods were chosen as they both test species boundaries by comparison of intra - and inter - lineage differences , but do so in different ways ; thus allowing the results of each method to be tested and validated by the other . only the mitochondrial data were used to validate the species hypotheses , as the nuclear gene sample sizes were limited and individually were not very informative ; for instance , most of the nuclear gene trees contained numerous polytomies and thus could not be used to identify genetically divergent groups .
the genetic distances between the hypothesised lineages and between specimens for both coi and 16s were calculated and visualised to determine whether a barcoding gap existed . as the intent of this method was to provide support for , or refutation of , the lineage hypotheses formed through the species discovery approaches , lineages were pre - defined based on those results and genetic distances categorised as representing either intra - or inter - lineage distances . for the purposes of this study , a barcoding gap was defined as a clear separation ( or ‘gap’ ) between the highest intra - lineage and lowest inter - lineage genetic distances measured between the suggested lineages . although a standard threshold has been suggested by hebert et al . ( 2004 ) for recognising distinct species ( 10× average intraspecific difference ) , this approach was not followed as it has been shown that there are vastly different rates of divergence for both different taxa and different genetic markers ( avise , 2009 ) . rather , a recognisable distinction between the inter - and intra - lineage distances was considered potential evidence for distinct species . analyses were undertaken for qld and nsw specimens separately .
relative divergences between genetic groups were calculated in mega5 . to determine inter - lineage divergence , the number of base substitutions per site was estimated from the net average between groups of sequences and the diversity between specimens was determined by calculating the number of base substitutions per site between each pair of sequences , both using a mcl model with 1 , 000 bootstrap replicates . the rate variation among sites was modelled with a gamma distribution with a shape parameter of 1 , with positions containing gaps and missing data eliminated . this was performed separately for both coi and 16s , with all unique haplotypes included .
the species discovery hypotheses were also tested using the k ∕θ method ( birky et al . , 2010 ; birky & barraclough , 2009 ; birky et al . , 2005 ) . although this method was originally developed for asexually - reproducing organisms and termed the 4x rule , it has been further developed and shown to be effective for the mtdna region for sexually - reproducing organisms ( birky , 2013 ) . this method provides a simple way of defining species groups based on specimens / populations that form clusters ( i . e . , clades ) that are separated by genetic gaps too deep to be ascribed to random genetic drift within a species and , therefore , must be due to diversifying selection or long - term physical isolation ( apte , smith & wallis , 2007 ) .
using the groups from the species discovery hypotheses , sister clades were identified and statistical support for these was tested . sequence divergences were estimated within ( d ) and between each sister clade using uncorrected p - distances calculated in mega5 . nucleotide diversity ( π ) was then calculated using π = dn ∕ ( n − 1 ) , where n is the number of samples per clade . theta ( θ ) was then estimated as θ = 2 neμ ( where ne is the effective populations size and μ is mutation rate per base pair per generation ) by calculating π∕ ( 1 − 4π∕3 ) within each clade . if d = 0 ( as it did for one clade in this study ) , then π can alternatively be calculated as 2∕ ln ( n − 1 ) , where l is the length of the sequence . k was then calculated for each sister - clade comparison ( using mega5 ) as the uncorrected net between group mean distance , with this divided by the highest θ in the comparison ( as this is the more conservative approach ) to provide k ∕θ . where sister clades were poorly defined in the tree , k was estimated between all potential sister clades in the polytomy , with the clade of the lowest k considered to be the sister clade . finally , following the method of birky ( 2013 ) , if the k ∕θ value was greater than 4 , then the sister clades were accepted as different lineages .
the genetic distances calculated between the qld and nsw groups using coi and 16s were 16 . 0 % and 12 . 7 % , respectively ( table 3 ) . these distances were as large as , or in some cases larger than , the distances calculated between these two groups and other closely related genera . furthermore , some distances between pairs of the other genera were smaller than those between the qld and nsw groups for both coi and 16s ( e . g . , geocharax versus engaeus = 13 . 7 % and 6 . 7 % , gramastacus versus engaeus = 11 . 7 % and 8 . 1 % ; table 3 ) .
estimates of net evolutionary divergence between groups of coi ( below diagonal ) and 16s ( above diagonal ) sequences with a mcl model .
groups that are identified as potentially representing distinct species will be referred to herein as lineages , and will form the groups to be analysed through lineage validation methods .
although not all groupings were statistically supported , both the ml and bayesian combined gene trees suggested the presence of multiple groups within qld and nsw , and displayed the same topologies ( fig . 2 ) . six clades were evident within the qld populations , with the monophyly of all but two highly supported ( as these were represented by single specimens ) . the first clade included maryborough and some tuan state forest specimens ( lineage 1 ; bs 90 % , pp 1 ) , and the second contained the remaining tuan state forest specimens as well as bribie island , type locality , and some beerburrum specimens ( lineage 2 ; bs 96 % , pp 1 ) . the two groups for which monophyly could not be established were represented by the remaining beerburrum specimens ( lineage 3 ) and hervey bay ( lineage 4 ) . the final two clades consisted of tewantin and lake weyba specimens ( lineage 5 ; bs 100 % , pp 1 ) and gold coast specimens ( lineage 6 ; bs 100 % , pp 1 ) . there was also some geographic structuring evident within each of the clades .
maximum likelihood phylogeny showing the proposed lineages for queensland ( lineages 1 through 6 ) and new south wales ( lineages 7 and 8 ) .
bootstrap values from the maximum likelihood analysis are shown above the branches , and posterior probabilities from the bayesian analysis are shown below branches for the major nodes , as both analyses produced identical topologies for the major nodes .
the two monophyletic clades evident within the nsw populations were strongly supported , and form lineage 7 ( lennox head ) and lineage 8 ( lake hiawatha , broadwater national park 1 & 2 ) ( fig . 2 ) . although there was some structuring evident within lineage 8 , the branching patterns were very shallow and were therefore not explored as potential distinct lineages .
summary of possible lineages based on ϕ - statistics for qld specimens using coi and 16s data .
see table s2 for explanation of how potential lineages were determined . where specimens from the same collection locality are split into two or more groups , details are included below the table for clarification .
a total of eight lineage arrangements was deemed plausible based on apparent genetic groupings and collection localities , and were tested using amovas ( table 4 ) . the process of assigning the potential lineages is outlined in table s2 , where a hierarchical approach was taken to split the tree into major genetic groups , minor genetic groups , and geographic localities . as there was no logical reason for combining the nsw lineages for the amova analysis based on either the phylogenetic or geographic information , the nsw populations were considered to consist of the lh lineage and the lakeh / bnp lineage . further testing , however , was considered appropriate to determine the lineages present within qld . figure 3 shows an increase in the ϕ ct estimate , with a plateau reached at six lineages for both the coi and 16s estimates . these six qld lineages represent the most parsimonious arrangement of the specimens into lineages .
values for potential lineages for both coi ( open circles ) and 16s ( black circles ) for queensland specimens .
as the combined gene tree was inferred using only specimens that were successfully sequenced for at least four of the five genes , not all collection localities were represented on the tree ( i . e . , tsfn , knp , moo , eu ) . of these localities , only tsfn was represented in the amova analysis , as the remaining localities were represented by a single sequence and therefore could not be included in the amova . in order to assign these populations to a lineage for further testing , the individual gene trees and haplotype networks were examined and the localities were designated through the closest phylogenetic connection ( data not shown ) . both of the species discovery approaches suggested the presence of eight lineages ( six in qld and two in nsw ; table 5 ) , and formed the lineages to be validated .
lineages assigned through two species discovery approaches and the final lineage hypothesis , for queensland and new south wales localities .
the coi data showed some overlap of the intra - and inter - lineage estimates within qld , resulting in no usable barcoding gap for lineage separation ( fig . 4a ) . where the overlap occurred , the low inter - lineage estimates were attributable to the lineage 1 vs . lineage 2 comparison , and the high intra - lineage estimates were seen between specimens within lineage 1 . however , many estimates between these two lineages fell in the higher range of the inter - lineage estimates as well as the low range .
the 16s data for qld populations showed a clearer relationship between lineages ( fig . 4c ) . although there was a very small overlap between the intra - and inter - lineage distances ( occurring between two specimens from lineage 1 ) , this represented an overlap of less than 0 . 01 % . when the existence of this overlap was disregarded , there was a small gap at 2 . 8–3 . 0 % . however , despite there not being a distinguishable gap due to the overlap , identification of the majority of lineages through the comparison of intra - and inter - lineage distances was clear and distinguishable .
intra - and inter - lineage genetic distance estimates ( white and grey , respectively ) for queensland lineages showing ( a ) coi estimates for all lineages , ( b ) coi estimates without comparisons between lineage 1 and 2 , ( c ) 16s estimates for all lineages , and ( d ) 16s estimates without comparisons between lineage 1 and 2 .
when the estimates within and between lineage 1 and 2 specimens were removed from both the coi and 16s data ( with the comparison between these two lineages and all other lineages remaining ) , a clear barcoding gap was seen in both data sets ( fig . 4b , fig . 4d ) . for coi , the gap occurred between 1 . 7–4 . 7 % , and between 0 . 9–3 . 5 % for 16s . this shows that all other qld groups ( i . e . , lineage 3 through 6 ) represent clear lineages based on the barcoding approach using both coi and 16s data .
for nsw populations , there was a clear barcoding gap between the two lineages ( i . e . , lineage 7 and 8 ) , occurring between 1 . 5 and 6 . 6 % for the coi data and 0 . 7–3 . 0 % for the 16s data ( fig . 5 ) .
intra - and inter - lineage genetic distance estimates ( white and grey , respectively ) for new south wales lineages showing ( a ) coi and ( b ) 16s estimates for all lineages .
the sister clades within qld and nsw were tested using the k ∕θ method for a delimitation of eight lineages ( six from qld , two from nsw ) using both coi and 16s data ( table 6 ) . in some instances , sister clades that were defined by the lowest k - distance ( as they were ambiguous based on the combined gene tree ) differed between the coi and 16s datasets . in these cases , only the relevant k ∕θ comparison for the applicable gene was calculated .
k ∕θ values for both coi and 16s for comparisons between sister clades within queensland and new south wales .
where specimens from the same collection locality are split into two or more lineages , details are included below the table for clarification . dashes are used where sister clades differ between coi and 16s .
lineage 1 = mar & tsfn & tsfsa ( specimen 4 ) & tsfsc ( specimens 8 , 17 , 22 ) .
lineage 2 = tsfsa - h ( specimens 1 - 3 , 5 - 7 , 9 - 12 , 14 , 16 , 18 - 21 , 23 - 30 ) & brb & tl & ber ( specimens 1 , 2 , 5 ) .
lineage 3 = ber ( specimens 3 , 4 , 6 , 7 ) .
although there was some ambiguity in the barcoding analysis of the qld coi data regarding the separation of lineage 1 and 2 , the 16s data showed support for the species discovery lineage hypothesis . because of the deeper phylogenetic inferences provided by 16s in addition to the fact that there were many genetic distances within and between lineage 1 and 2 falling within the expected distributions , the lineage hypothesis for qld populations was considered supported by this analysis ( table 7 ) . the two nsw lineages were clearly separate based on both the coi and 16s data and were therefore also supported ( table 7 ) . in the k ∕θ analysis , all lineages were supported by both genes with the exception of the split between lineage 1 and 2 ( both genes ) , and lineage 1 and 3 ( 16s ) ( table 7 ) .
the species discovery lineage hypothesis and two lineage validation methods , with the final assignment of lineages for queensland and new south wales localities .
although it is difficult to define what degree of separation is necessary between genera at a molecular level ( rach et al . , 2008 ) , based on the genetic distances presented in this study there is strong support for division at genus level . for instance , the genetic distance between qld and nsw populations is larger than that seen between engaeus and both geocharax and gramastacus for the coi and 16s gene fragments , and between engaewa and both geocharax and engaeus for coi . other genera also show smaller genetic distances when compared to either qld or nsw than these two groups do with each other . regardless of which genera were genetically closer to each other , the distance between qld and nsw is at least as large as those between existing genera ( table 3 ) , thereby supporting their separation into two distinct genera .
all samples used in this study to infer the individual gene trees . * indicates samples obtained from genbank , qb indicates samples obtained from q . burnham unpublished data , and all others are from this study . where samples have more than one id name , the first is the identification used in this study , and the second is the identifier from genbank .
many thanks are given to the volunteers that helped with the field work ; dr seanan wild , amanda dawson , dr dianna virkki , and shane howard . we are also grateful for additional genetic material provided by dr andrew bentley and dr quinton burnham .
kathryn l . dawkins conceived and designed the experiments , performed the experiments , analyzed the data , wrote the paper , prepared figures and / or tables , reviewed drafts of the paper .
james m . furse conceived and designed the experiments , reviewed drafts of the paper .
jane m . hughes conceived and designed the experiments , contributed reagents / materials / analysis tools , reviewed drafts of the paper .
the following information was supplied relating to field study approvals ( i . e . , approving body and any reference numbers ) :
all specimens from this study were collected under permits witk08599510 , wisp08599610 , and twb / 01 / 2011 issue by the department of environment and resource management .
sequences obtained in this study were deposited in genbank under accession numbers kx669691 – kx670093 , kx753349 .
this project was completed as part of a phd project undertaken by the principal author , funded by an australian postgraduate award ( 2009 ) and the griffith school of environment , gold coast campus , queensland , australia . there was no additional external funding received for this study . the funders had no role in study design , data collection and analysis , decision to publish , or preparation of the manuscript .
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riek , e . f . ( 1951 ) . the freshwater crayfish ( family parastacidae ) of queensland . records of the australian museum . 22 ( 4 ) : 368 - 388 . , available online at urltoken [ details ]
crandall , k . a . & s . de grave . ( 2017 ) . an updated classification of the freshwater crayfishes ( decapoda : astacidea ) of the world , with a complete species list . journal of crustacean biology . page ( s ) : 381 [ details ]
kathryn l . dawkins a d , james m . furse b , clyde h . wild b and jane m . hughes c
a australian rivers institute , griffith university , gold coast , qld 4222 , australia .
b environmental futures centre , griffith university , gold coast , qld 4222 , australia .
c australian rivers institute , griffith university , nathan , qld 4111 , australia .
this study was conducted as a major part of a b . sc . ( hons ) by kathryn dawkins under the supervision of james furse , professor jane hughes and professor clyde wild . funding for this study was provided by both the australian rivers institute and the griffith school of environment , griffith university . additional funding for genetic analysis was also provided by rob mccormack and his company ‘aabio’ and was greatly appreciated . the authors would like to thank the two anonymous reviewers and the associate editor for their helpful comments , michael arthur for statistical guidance and rob mccormack , jason coughran and many other volunteers for field assistance . crayfish were collected under nsw scientific collection permit p05 / 0077 - 3 . 1 and qld general fisheries permit # 91210 .
austin , c . m . , and ryan , s . g . ( 2002 ) . allozyme evidence for a new species of freshwater crayfish of the genus cherax erichson ( decapoda : parastacidae ) from the south - west of western australia . invertebrate systematics 16 , 357–367 . | crossref |
baker , a . m . , williams , s . a . , and hughes , j . m . ( 2003
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) . mitochondrial dna reveals phylogenetic structuring and cryptic diversity in australian freshwater macroinvertebrate assemblages .
( decapoda : parastacidae ) on the mainland and islands of southeast queensland . bsc ( hons ) thesis , griffith university , nathan .
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) . extensive intraspecific genetic diversity of a freshwater crayfish in a biodiversity hotspot .
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) . phylogenetic relationships among the australian and new zealand genera of freshwater crayfishes ( decapoda : parastacidae ) .
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) . low genetic variability of the white - clawed crayfish in the iberian peninsula : its origin and management implications .
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) . arlequin ver . 3 . 0 : an integrated software package for population genetics data analysis .
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warning : the ncbi web site requires javascript to function . more . . .
kathryn l . dawkins , 1 james m . furse , 2 , 3 clyde h . wild , 2 and jane m . hughes 4
the parastacid crayfish genera are generally highly speciose , with novel species and genetically diverse groups commonly found ( e . g . ,
, which contains the species with the smallest body size in the parastacidae huxley , 1879 . although it has previously been highlighted as containing genetically diverse groups ( see
, on the basis of electrophoretic and geographical differences , that previously unrecognised genetic diversity existed within the genus . subsequently , two genetically divergent groups were identified within this region by
, both of which showed considerable internal genetic variability . the two identified groups aligned with populations from qld and nsw , respectively , and were suggested to represent species that diverged as a result of long - term historical geographic isolation (
, utilising molecular data across several gene regions and employing multiple species delimitation methods in order to determine the most likely species groups .
the triangle and bolded name denotes the type locality . grey lines denote drainage boundaries , and the black line denotes the border between queensland and new south wales . refer to
. all specimens from this study were collected under permits witk08599510 , wisp08599610 , and twb / 01 / 2011 issued by the department of environment and resource management . dna was extracted from specimens preserved in 70 % ethanol using a variation of the cetyltrimethyl ammonium bromide / phenol - chloroform extraction protocol (
were also sequenced for inclusion as outgroups . where sequences from these outgroups could not be obtained ( i . e . , due to non - amplification ) , alternative sequences were retrieved from genbank ( details in
the genetic distances calculated between the qld and nsw groups using coi and 16s were 16 . 0 % and 12 . 7 % , respectively (
) . these distances were as large as , or in some cases larger than , the distances calculated between these two groups and other closely related genera . furthermore , some distances between pairs of the other genera were smaller than those between the qld and nsw groups for both coi and 16s ( e . g . ,
) . six clades were evident within the qld populations , with the monophyly of all but two highly supported ( as these were represented by single specimens ) . the first clade included maryborough and some tuan state forest specimens ( lineage 1 ; bs 90 % , pp 1 ) , and the second contained the remaining tuan state forest specimens as well as bribie island , type locality , and some beerburrum specimens ( lineage 2 ; bs 96 % , pp 1 ) . the two groups for which monophyly could not be established were represented by the remaining beerburrum specimens ( lineage 3 ) and hervey bay ( lineage 4 ) . the final two clades consisted of tewantin and lake weyba specimens ( lineage 5 ; bs 100 % , pp 1 ) and gold coast specimens ( lineage 6 ; bs 100 % , pp 1 ) . there was also some geographic structuring evident within each of the clades .


spinomantis peraccae is a species of frog in the mantellid subfamily mantellinae , endemic to madagascar .

the specific epithet honours italian herpetologist mario giacinto peracca .

spinomantis peraccae

description of two tadpoles of malagasy treefrogs , spinomantis sp . aff . peraccae and spinomantis tavaratra
- - synonym : mantidactylus ( spinomantis ) peraccae — glaw and vences , 1994 .
mantidactylus ( spinomantis ) peraccae — glaw and vences , 1994 , fieldguide amph . rept . madagascar , ed . 2 : 403 .
no one has contributed data records for spinomantis yet . learn how to contribute .
rhacophorus ( rhacophorus ) peraccae — ahl , 1931 , das tierreich , 55 : 191 .
this recently described species is similar to spinomantis peraccae and s . elegans . there is confustion regarding the taxonomy of this group ( additional species might be involved ) , and a revision is needed .
- - . . . none , mantidactylus opiparis , unknown . none , mantidactylus peraccae , unknown . none ,
mantidactylus ( guibemantis ) peraccae — dubois , 1992 , bull . mens . soc . linn . lyon , 61 : 312 .
descriptions of two new spinomantis frogs from madagascar ( amphibia : mantellidae ) , and new morphological data for s . brunae and s . massorum
mantidactylus peraccae — blommers - schlösser , 1978 , genetica , 48 : 32 . blommers - schlösser , 1979 , beaufortia , 29 : 43 .
( pdf ) descriptions of two new spinomantis frogs from madagascar ( amphibia : mantellidae ) , and new morphological data for s . brunae and s . massorum
m 34 - 44 mm . the only arboreal spinomantis without distinct fringes or spines . usually light brown to greenish on the dorsum with distinct rounded brown patches .
spinomantis peraccae — vences and glaw , 2006 , in vences et al . ( eds . ) , calls frogs madagascar : 25 . glaw and vences , 2006 , organisms divers . evol . , 6 : 248 ; glaw and vences , 2006 , organisms divers . evol . , electron . suppl . , 11 ( 1 ) : 2 .
andreone , f . , glaw , f . , vences , m . and vallan , d . 1998 . a new mantidactylus from south - eastern madagascar , with a review of mantidactylus peraccae ( ranidae : mantellinae ) . herpetological journal 8 : 149 - 159 .
. . . for instance , 34 of 77 species of the treefrog genus boophis have been described since 2000 ( e . g . , glaw et al . 2010 ) . in contrast , only two species of spinomantis were described in the same period ( cramer et al . 2008 ) , although another 11 candidate species were identified by molecular data ( vieites et al . 2009 ; perl et al . 2014 ) . spinomantis bertini ( guibé , 1947 ) is a poorly known frog of 22−28 mm snout–vent length , inhabiting stream edges in primary rainforest of the south - eastern malagasy andohahela massif ( glaw & vences 2007 ) . . . .
rhacophorus peraccae boulenger , 1896 , ann . mag . nat . hist . , ser . 6 , 18 : 421 . holotype : bmnh 1947 . 2 . 9 . 70 , according to blommers - schlösser and blanc , 1991 , faune de madagascar , 75 : 153 . type locality :
ivohimanita
, northwestern madagascar .
spinomantis brunae — vences and glaw , 2006 , in vences et al . ( eds . ) , calls frogs madagascar : 26 . glaw and vences , 2006 , organisms divers . evol . , 6 : 248 ; glaw and vences , 2006 , organisms divers . evol . , electron . suppl . , 11 ( 1 ) : 2 .
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gland , switzerland , 5 july 2018 ( iucn ) – australia’s unique reptiles – including lizards and snakes – face severe threats from invasive species and climate change , with 7 % of th . . .
the value of medicinal and aromatic plant trade has increased three - fold in the past 20 years , but traditional harvesting practices are being replaced by less sustainable alternatives . . . .
a recently released iucn technical brief recommends increasing investments in sustainable land management practices , as well as better cooperation between agriculturalists and conservationists to conse . . .
frost , d . r . 2015 . amphibian species of the world : an online reference . version 6 . 0 . new york , usa . available at : urltoken .
the taxonomic status of this species is in doubt . some populations assigned to it might belong to other species ( andreone et al . 1998 ) .
justification : listed as least concern in view of its wide distribution and presumed large population .
this species ranges widely in northern , eastern and central madagascar from tsaratanana south to andohahela , at 500 - 2 , 000 m asl . recent records have confirmed it occurring in bemanevika ( rabearivony et al . 2010 ) .
it is a locally abundant species . due to ongoing declines in the extent and quality of habitat , the population is suspected to be decreasing .
it is an arboreal species living along streams in pristine rainforest and it is not found in altered habitats . the eggs are placed on leaves above water , and the larvae develop in slow - flowing streams .
its forest habitat is receding due to subsistence agriculture , timber extraction , charcoal manufacture , and invasive spread of eucalyptus , livestock grazing and expanding human settlements .
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variation : specimens from tsaratanana , have a more brownish colour and indistinct patterning and very melodious , “metallic” - sounding calls , and may represent a different species .
is larger , with a less granular skin and has only an inner metatarsal tubercle .
ambohitantely , andranomay forest , anjanaharibe , ankazobe , ankeniheny , chaines anosyennes , farihimazava , ranomafana ( maharira forest , ranomena , vohiparara ) . it occurs between 500m - 2 , 000m asl in pristine rainforest ( andreone et al . 2008 ) .
leaf axil . one calling male was captured in february , about 5 m above a brook in rain forest . several other specimens were heard along forest brooks .
call ( from ankeniheny ) : a single ' explosive ' short note that can be described as ' pom ' . note duration is about 75 ms . frequency is between 1 . 2 and 4 . 2 khz , with intensity maxima at 1 . 3 and 2 . 6 khz . call is repeated after intervals of at least about 30 seconds . similar calls were heard at andasibe .
breeding takes place in slow - moving streams ( andreone et al . 2008 ) .
andreone , f . , vallan , d . , and nussbaum , r . ( 2008 ) .
. in : iucn 2008 . 2008 iucn red list of threatened species . www . iucnredlist . org . downloaded on 29 april 2009 .
miguel vences and frank glaw ( m . vences at tu - bs . de ) , assistant professor and curator of vertebrates at the institute for biodiversity and ecosystem dynamics in the zoological museum at the university of amsterdam .
> university of california , berkeley , ca , usa . accessed jul 9 , 2018 .
> university of california , berkeley , ca , usa . accessed 9 jul 2018 .
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peracca ' s madagascar frog ( frank and ramus , 1995 , compl . guide scient . common names amph . rept . world : 101 ) .
holotype redescribed by andreone , glaw , vences , and vallan , 1998 , herpetol . j . , 8 : 152 . glaw and vences , 2007 , field guide amph . rept . madagascar , ed . 3 : 206 - 207 , provided an account .
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for access to available specimen data for this species , from over 350 scientific collections , go to vertnet .
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this species ranges widely in northern , eastern and central madagascar from . . .
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amphibian species of the world : an online reference v5 . 3 , database ( version 5 . 3 )
frost , darrel r . 2009 . amphibian species of the world : an online reference . version 5 . 3 ( 12 february , 2009 ) . electronic database accessible at urltoken american museum of natural history , new york , usa
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to cite this page : myers , p . , r . espinosa , c . s . parr , t . jones , g . s . hammond , and t . a . dewey . 2018 . the animal diversity web ( online ) . accessed at https : / / animaldiversity . org .
disclaimer : the animal diversity web is an educational resource written largely by and for college students . adw doesn ' t cover all species in the world , nor does it include all the latest scientific information about organisms we describe . though we edit our accounts for accuracy , we cannot guarantee all information in those accounts . while adw staff and contributors provide references to books and websites that we believe are reputable , we cannot necessarily endorse the contents of references beyond our control .
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living within fallen palm leaves : the discovery of an unknown blommersia ( mantellidae : anura ) reveal . . .
field observations on some dwarf chameleons ( brookesia spp . ) from rainforest areas of madagascar , wi . . .
six species of brookesia dwarf chameleons were recorded during two rainforest expeditions in madagascar . one of these is described as a new species brookesia valerieae based on the arrangement of the nine pairs of dorso - lateral spines , the lack of a pelvic shield , and dorsal chevron markings . observations were made on the defence and roosting behaviours shown by these six species . aspects . . . [ show full abstract ]
gephyromantis tricinctus guibé , 1947 ( anura : ranidae : mantellinae ) , known only from the type series and currently considered as a synonym of mantidactylus biporus , is resurrected as mantidactylus tricinctus , and included in the subgenus brygoomantis . a detailed redescription , based on specimens recently collected in central eastern madagascar , notes on natural history , and a description of . . . [ show full abstract ]
applications of ecological niche modeling for species delimitation : a review and empirical evaluatio . . .
although the systematic utility of ecological niche modeling is generally well known ( e . g . , concerning the recognition and discovery of areas of endemism for biogeographic analyses ) , there has been little discussion of applications concerning species delimitation , and to date , no empirical evaluation has been conducted . however , ecological niche modeling can provide compelling evidence for . . . [ show full abstract ]
a new phytotelmic species of platypelis ( microhylidae : cophylinae ) from the betampona reserve , easte . . .
we describe a new arboreal and diminutive species of the genus platypelis from the réserve naturelle intégrale n . 1 de betampona , one of the last low - altitude rainforest fragments of eastern madagascar . p . karenae sp . nov . is a phytotelmic species , living among leaves of pandanus spp . and those of a herbaceous plant of the genus crinum . amongst species of comparable size , the new species is . . . [ show full abstract ]
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c . michael hogan selected
madagascar subhumid forests habitat
to show in overview on
hapalemur aureus meier , albignac , peyriéras , rumpler and wright , 1987
.
jennifer hammock added an association between
madagascar subhumid forests habitat
and
prolemur simus
.
jennifer hammock added an association between
madagascar subhumid forests habitat
and
liopholidophis sexlineatus günther 1882
.
jennifer hammock added an association between
madagascar subhumid forests habitat
and
liopholidophis grandidieri mocquard 1904
.
jennifer hammock added an association between
madagascar subhumid forests habitat
and
pseudoxyrhopus ankafinaensis raxworthy & nussbaum 1994
.
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taxonomy - ontology - tool / amphibiaweb . txt at master · nescent / taxonomy - ontology - tool · github
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listed as endangered because its extent of occurrence ( eoo ) is 1 , 883 km 2 , all individuals are in fewer than five locations , and there is continuing decline in the extent and quality of its habitat in southeastern madagascar .
this species is known from andohahela national park and manantantely in extreme southeastern madagascar . it has been recorded from 300 - 600 m asl and its extent of occurrence ( eoo ) is 1 , 883 km 2 .
it is thought to be locally moderately common . however , due to ongoing declines in the extent and quality of habitat , the population is suspected to be decreasing .
it lives in crevices among boulders and rocky areas in pristine forest , usually close to flowing waters , and does not survive in secondary or degraded areas . its breeding biology is unknown , though it possibly takes place in water flowing among rocks .
the major threat to this species is habitat loss due to subsistence agriculture , timber extraction , charcoal manufacture , the invasive spread of eucalyptus , livestock grazing , and expanding human settlements .
conservation actions it occurs in andohahela national park . conservation needed improved protection and management of its habitat is required , including inside the boundaries of protected areas . research needed further work is required to better understand its population size , distribution , and trends , and its life history and ecology .
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andreone f , carpenter ai , cox n , du preez l , freeman k , furrer s et al ( 2008 ) the challenge of conserving amphibian megadiversity in madagascar . plos biol 6 : 943–946
andreone f , randriamahazo h ( 2008 ) sahonagasy action plan . conservation strategies for the amphibians of madagascar . mus reg sci nat torino , torino , italy
bielby j , cooper n , cunningham aa , garner twj , purvis a ( 2008 ) predicting susceptibility to future declines in the world’s frogs . conserv lett 1 : 82–90
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and amphibian chytridiomycosis in space , time and host . ann rev microbiol 63 : 291–310
gascon c , collins jp , moore rd , church dr , mckay je , mendelson jr iii ( 2007 ) amphibian conservation action plan . iucn , conservation international , gland , switzerland , cambridge , uk
gehring ps , köhler j , strauß a , randrianiaina rd , glos j , glaw f , vences m ( 2011 ) the kingdom of the frogs : amphibian radiations in madagascar . in : zachos fe , habel jc ( eds ) biodiversity hotspots . springer , heidelberg
glaw f , vences m ( 2007 ) a field guide to the amphibians and reptiles of madagascar , 3rd edn . vences and glaw verlag , cologne , germany
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lötters s . , rödder d . , kielgast j . , glaw f . ( 2011 ) hotspots , conservation , and diseases : madagascar’s megadiverse amphibians and the potential impact of chytridiomycosis . in : zachos f . , habel j . ( eds ) biodiversity hotspots . springer , berlin , heidelberg
humid forests in the anosy mountains from andohahela and manantantely , southeastern madagascar , 600 - 800 m elevation .
, but smaller and nearly without dermal flaps and fringes . females unknown . other characters see next section .
hand without web ; webbing of the foot 1 ( 0 . 5 ) , 2i ( 1 ) , 2e ( 0 . 25 ) , 3i ( 1 . 25 ) , 3e ( 0 . 25 ) , 4i / e ( 1 . 5 ) , 5 ( 1 ) . skin on the back granular . tubercles are present on the back , on the head , and on the eyes . on the posterior edge of foot and tarsus 8 - 9 small tubercles can be recognized . slightly distinct tubercles on the heels . only slight longitudinal furrows between the eyes .
colour in life light brown on the back , with rather irregular brown spots and larger markings , and some greenish colour . hindlegs with dark brown crossbands . some white patches are present on the flanks . underside uniformly whitish except the femurs which are marbled with brown . the bones are light green .
paratype : adult male , zfmk 57443 , from same locality as holotype . svl is 33 . 5 mm . morphological features very similar to the holotype . the general colour in life of the back and legs was mossy green ; in preservative this colour has disappeared . two distinct tubercles are present on each heel . the femoral glands measure 10x3 . 5 mm .
benavony , manongarivo , tsaratanana ( antsahamanara campsite ) . it occurs between 200 - 1 , 100m asl in streams near primary forest but not secondary habitats ( raxworthy and glaw 2008 ) .
habits : calling males were found in march along a brook in primary forest about 2 m above the ground . calling activity had a peak at dusk , starting before 17 h and was nearly finished at 19 . 30 h .
calls : a single pulsed note , consisting of a minimum of two pulses , but sometimes with more pulses , loud but not very conspicuous , and less “metallic” in appearance than in other species of the group .
it might occur in the réserve spéciale de manongarivo and réserve naturelle intégrale du tsaratanana ( raxworthy and glaw 2008 ) .
adults 50 - 60 mm . tibiotarsal articulation reaches the nostril . hand without webbing , foot webbing 1 ( 0 . 5 ) , 2i ( 1 ) , 2e ( 1 ) , 3i ( 2 ) , 3e ( 1 ) , 4i / e ( 2 ) , 5 ( 0 . 5 ) . dorsal skin smooth . colour dorsally light brown with distinct rounded dark brown patches that are delimited by a white or yellow line . males with distinct , prominent dark femoral glands ; shape and size of vocal sac unknown . similar species : mainly
andohahela , andohariana , chaines anosyennes , andringitra ( cuvette boby , imaitso forest ) , ivohibe , maharira summit ( ranomafana ) . it occurs between 1 , 350 - 2 , 500m asl along rocky outcrops in forested zones and above tree line ( cadle and raxworthy 2008 ) .
habits : a species restricted to high elevations , occurring among large boulders or in small caves above the tree - line or in rainforest above 1200 m elevation . at andringitra , subadults can be found hidden under rocks at high elevations above the tree line . males call from within caves and cavities during day and night . very large blackish tadpoles develop in streams .
calls : a very soft and inconspicuous sound for such a relatively large frog , reminding the dripping of water .
breeding takes place in streams . it takes at least a year for the tadpoles to begin metamorphosis ( cadle and raxworthy 2008 ) .
, its distribution is severely fragmented , and there is a continuing decline in the extent and quality of its forest habitat . it occurs in parc national de ranomafana , parc national d ' andringitra , parc national d ' andohahela , and probably parc national de midongy du sud ( cadle and raxworthy 2008 ) .
. in : iucn 2008 . 2008 iucn red list of threatened species . www . iucnredlist . org . downloaded on 05 may 2009 .
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- - elena ' s treefrog ( boophis elenae ) - images . everett ' s tree frog ( rhacophorus everetti ) -
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- - amphibian species of the world 3 . 0 an online reference . . . .
home | photos index | search | about | contact unless otherwise noted , all content and images are the property of rhett butler , content copyright 2004 - 2008 . all rights reserved .
1 . forest - > 1 . 6 . forest - subtropical / tropical moist lowland suitability : suitable season : resident major importance : yes 1 . forest - > 1 . 9 . forest - subtropical / tropical moist montane suitability : suitable season : resident major importance : yes 5 . wetlands ( inland ) - > 5 . 1 . wetlands ( inland ) - permanent rivers / streams / creeks ( includes waterfalls ) suitability : suitable season : resident major importance : yes
1 . residential & commercial development - > 1 . 1 . housing & urban areas
2 . agriculture & aquaculture - > 2 . 1 . annual & perennial non - timber crops - > 2 . 1 . 1 . shifting agriculture
2 . agriculture & aquaculture - > 2 . 1 . annual & perennial non - timber crops - > 2 . 1 . 2 . small - holder farming
2 . agriculture & aquaculture - > 2 . 3 . livestock farming & ranching - > 2 . 3 . 2 . small - holder grazing , ranching or farming
5 . biological resource use - > 5 . 3 . logging & wood harvesting - > 5 . 3 . 5 . motivation unknown / unrecorded
8 . invasive and other problematic species , genes & diseases - > 8 . 1 . invasive non - native / alien species / diseases - > 8 . 1 . 2 . named species
blommers - schlösser , r . m . a . 1979 . biosystematics of the malagasy frogs . i . mantellinae ( ranidae ) . beaufortia 29 ( 352 ) : 1 - 77 .
blommers - schlösser , r . m . a . and blanc , c . p . 1991 . amphibiens ( première partie ) . fauna de madagascar 75 : 1 - 379 .
glaw , f . and vences , m . 1994 . a fieldguide to the amphibians and reptiles of madagascar . second edition . zoologisches forschungsinstitut und museum alexander koenig , bonn .
glaw , f . and vences , m . 2007 . a fieldguide to the amphibians and reptiles of madagascar . third edition . vences & glaw verlag , cologne .
iucn . 2016 . the iucn red list of threatened species . version 2016 - 1 . available at : urltoken . ( accessed : 30 june 2016 ) .
rabearivony , j . , raselimanana , a . p . , andriamazava , m . a . , thorstrom , r . and rene de roland , l - a . 2010 . a new locality for the endangered microhylid frog scaphiophryne boribory from northern madagascar and a rapid survey of other amphibians of the bemanevika region . herpetology notes 3 : 105 - 109 .
raxworthy , c . j . and nussbaum , r . a . 1996 . amphibians and reptiles of andringitra massif : a study of elevational distribution and local endemicity . fieldiana zoology 85 : 158 - 170 .
the iucn red list of threatened species . version 2018 - 1 . < urltoken > . downloaded on 09 july 2018 .
a small conspicuously coloured frog ; males 22 - 23 mm , females 25 - 28 mm . colour in life unknown . in preservative , dorsum brownish with distinct dark spots , flanks darker with light spots . dorsal surface of legs white with distinct narrow black bands . venter white with dark spots , which are more or less circular on the throat and become more oblong towards the abdomen . skin smooth . nostrils nearer to tip of snout than to the eye . tympanum very distinct , about 1 / 2 of eye diameter . tibiotarsal articulation reaches the eye . lateral metatarsalia partly connected . webbing rudimentary . fingers with terminal disks . oblong femoral glands clearly visible in males .
similar species : colouration is unique . morphology is similar to the m . wittei - complex .