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

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

the genus sigaus is endemic to new zealand .

sigaus childi

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

it is also distributed in alaska and mexico .

pachymerium ferrugineum

pachymerium ferrugineum ( c . l . koch , 1835 ) - natural history museum
worms - world register of marine species - pachymerium ferrugineum ( c . l . koch , 1835 )
jennifer hammock split the classifications by nmnh entomology resource from pachymerium ferrugineum koch ( 1835 ) to their own page .
hox gene sequences from the geophilomorph centipede pachymerium ferrugineum ( c . l . koch , 1835 ) ( chilopoda : geophilomorpha : geophilidae ) : implicatio . . . - pubmed - ncbi
hox gene sequences from the geophilomorph centipede pachymerium ferrugineum ( c . l . koch , 1835 ) ( chilopoda : geophilomorpha : geophilidae ) : implications for the evolution of the hox class genes of arthropods
hox gene sequences from the geophilomorph centipede pachymerium ferrugineum ( c . l . koch , 1835 ) ( chilopoda : geophilomorpha : geophilidae ) : implications for the evolution of the hox class genes of arthropods .
hox gene sequences from the geophilomorph centipede pachymerium ferrugineum ( c . l . koch , 1835 ) ( chilopoda : geophilomorpha : geophilidae ) : implications for the evolution of the hox class genes of arthropods - sciencedirect
in the majority of habitats one species was overwhelmingly dominant ( from 41 . 5 % to 95 . 2 % ) . in rushes at oxbows and in the wet meadows it was lamyctes emarginatus , in riparian forests lamyctes curtipes , in thermophilous thickets lamyctes mutabilis , in sandy grasslands lithobius dudichi , and in the mesic meadows pachymerium ferrugineum . it was only in xerothermic grasslands that two species co - dominated ( lithobius dudichi and pachymerium ferrugineum ) ( table 1 ) .
chilopoda were collected throughout all the months of the study , although one can observe certain tendencies in particular species – especially with regard to four , most frequent species . lithobius curtipes was active throughout all the months , though predominantly in october and november . lamyctes emarginatus occurred from june to november , while it was most numerous in september . pachymerium ferrugineum was reported from april until november , especially in may and june , but also in august and september . finally , lithobius dudichi was most active in may and november .
( of geophilus ferrugineum c . l . koch , 1835 ) matic , z . ( 1972 ) . clasa chilopoda , subclasa epimorpha . fauna republicii socialiste românia , 6 ( 2 ) : 1 - 224 . bucuresti page ( s ) : 160 [ details ]
( of geophilus ferrugineum c . l . koch , 1835 ) zapparoli , m . ( 2002 ) . a catalogue of the centipedes from greece ( chilopoda ) . fragmenta entomologica , 34 ( 1 ) : 1 - 146 . roma page ( s ) : 94 [ details ]
distribution probably one of the most widespread chilopod species in the world . there are very few records , all from coastal shingle , . . .
distribution probably one of the most widespread chilopod species in the world . there are very few records , all from coastal shingle , from great britain . rather eastern distibution in scandinavia . [ details ]
barber , a . d . ( 2018 ) . world database of littoral myriapoda .
barber , a . d . ( 2009a ) centipedes . linnean society synopses of the british fauna ( new series ) 58 . field studies council , shrewsbury . [ details ]
iorio , e . ( 2005 ) contribution à la connaissance des chilopodes de bretagne ( myriapoda , chilopoda ) . bull . soc . linn . bordeaux 140 ( 33 ) : 149 - 156 [ details ]
johns , p . m . ( 2010 ) . phylum arthropoda subphylum myriapoda : centipedes , millipedes , pauropods symphylans , in : gordon , d . p . ( ed . ) ( 2010 ) . new zealand inventory of biodiversity : 2 . kingdom animalia : chaetognatha , ecdysozoa , ichnofossils . pp . 90 - 97 . [ details ]
palmen , e . ( 1949 ) the chilopoda of eastern fennoscandia . ann . zool . soc . zool . bot . fenn . vanamo 13 ( 4 ) : 1 - 45 [ details ]
przhiboro , a . ( 1994 ) fauna of terrestrial arthropods of the intertidal zone of kandalaksha bay , the white sea . unpublished phd thesis , st . petersburg state university . [ details ]
ecology a terrestrial species also occurring on seashores . apparently quite tolerant of seawater and so likely to be dispersed by rafting . [ details ]
a large ( to 50 mm ) reddish - orange centipede with 43 - 45 leg pairs and numerous small pores distributed over both the dorsal and ventral surfaces of the coxae of the last legs .
it is known from a handful of coastal sites in south - east england , invariably in shingle above the storm drift line .
a terrestrial species also occurring on seashores . apparently quite tolerant of seawater and so likely to be dispersed by rafting .
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as far as i know , no soil centipedes have official common names . i made this one up , to refer to the elongate fangs and the fact that this species often occurs in the intertidal zone .
; also japan , alaska , mexico [ info from wikipedia , not checked . = v = ]
in soil , beneath rocks and debris , often in moist areas . this species is well - known for often inhabiting the intertidal zone where it can tolerate long periods of immersion in salt water .
small invertebrates . in the intertidal zone they eat barnacles , amphipods , polychaete worms , etc .
de jong y . s . d . m . ( ed . ) ( 2011 ) fauna europaea version 2 . 4
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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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barber , a . d . ( 2009 ) . littoral myriapods . a review . < em > soil organisms . < / em > 81 ( 3 ) : 735 - 760 .
barber , a . d . ( 2009a ) centipedes . linnean society synopses of the british fauna ( new series ) 58 . field studies council , shrewsbury .
iorio , e . ( 2005 ) contribution à la connaissance des chilopodes de bretagne ( myriapoda , chilopoda ) . bull . soc . linn . bordeaux 140 ( 33 ) : 149 - 156
johns , p . m . ( 2010 ) . phylum arthropoda subphylum myriapoda : centipedes , millipedes , pauropods symphylans , in : gordon , d . p . ( ed . ) ( 2010 ) . new zealand inventory of biodiversity : 2 . kingdom animalia : chaetognatha , ecdysozoa , ichnofossils . pp . 90 - 97 .
koch c . l . ( 1835 ) deutschlands crustaceen myriapoden und arachniden . in : ( panzer ) herrich - schaeffer ' s deutschlands insecten : pustet , regensburg . heft 136
matic z . ( 1972 ) fauna republicii socialiste romania . clasa chilopoda subclasa epimorpha : academiei republicii socialiste romania bucuresti . 6 : 1 - 220
palmen , e . ( 1949 ) the chilopoda of eastern fennoscandia . ann . zool . soc . zool . bot . fenn . vanamo 13 ( 4 ) : 1 - 45
przhiboro , a . ( 1994 ) fauna of terrestrial arthropods of the intertidal zone of kandalaksha bay , the white sea . unpublished phd thesis , st . petersburg state university .
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department of general zoology , adam mickiewicz university ul . umultowska 89 , 61 - 614 poznań , poland
“natura” ecology research laboratory marek wierzba , ul . kubusia puchatka 78 , żabokliki 08 - 110 siedlce , poland
siedlce university of natural science and humanities , faculty of natural sciences , department of zoology , ul . b . prusa 12 , 08 - 110 siedlce , poland
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leśniewska m , jastrzębski p , stańska m , hajdamowicz i ( 2015 ) centipede ( chilopoda ) richness and diversity in the bug river valley ( eastern poland ) . in : tuf ih , tajovský k ( eds ) proceedings of the 16th international congress of myriapodology , olomouc , czech republic . zookeys 510 : 125–139 . doi : 10 . 3897 / zookeys . 510 . 8763
habitats in valleys of european rivers are relatively poorly known in terms of species diversity , habitat selection and the dynamics of chilopoda communities . the very few studies in this field include , for instance , zerm ( 1997a , b , 1999 ) . in poland , such studies have not been conducted so far .
centipedes from river valleys have been studied mainly in the context of changes in the communities as a result of seasonal flooding ( zulka 1991 , pižl and tajovský 1998 , tajovský 1999 , tuf 2000 , 2003 , tufová and tuf 2005 , marx et al . 2009 ) , and in the context of life strategies enabling survival in periodically flooded habitats ( adis et al . 1996 , adis and junk 2002 ) . xerothermic environments ( often naturally occurring in the river valleys ) have rarely been the subject of research on chilopoda ( voigtländer 2003 ) .
river valleys , especially the natural ones , only slightly changed – unregulated , are extremely valuable areas with habitats found more and more rarely , which already start to disappear across the continent . studies on these habitats provides an opportunity not only to learn about the biodiversity but also to develop appropriate management and protection schemes . the bug river is one of the few rivers of such a size in europe , which still remain almost unregulated ( dombrowski et al . 2002 ) . its length in poland amounts to 587 km .
research in the bug river valley was conducted since march to november in 2007 and 2008 .
in the lower course of the river , where the valley is much wider ( it stretches up to several kilometers wide ) with an overgrown flood terrace at its bottom . in this section , the bug river slowly meanders and sometimes changes its course ( near localities : morzyczyn , płatkownica , in the protected area ‘bug landscape park’ ) .
location of the study area in poland . study area . morzyczyn , płatkownica – lower section of the bug valley : mogielnica , zabuże , gnojno – central section of the bug valley .
in the bug valley we can come across habitats that vary in terms of moisture content and structure – two important parameters from the point of view of chilopoda biology . on the side of the river – from flooded and very humid habitats through medium moist ones to grasslands . riparian forests within the floodplain are closest to the river , then there are meadows of lower flooded terrace – submerged during river floods . mesic meadows are found in higher terraces , while slopes of terraces feature xerothermic grasslands , and thermophilous thickets . sandy grasslands and rushes at oxbows are located in the mosaic of meadows , sometimes closer to the river bed , and sometimes closer to the edge of the valley ( głowacki et al . 2002 ) . in terms of the structure , according to the classification by voigtländer ( 2005 ) one can distinguish high ( riparian forests ) , middle ( thermophilous thickets , rushes at oxbows ) and low ( meadows , grasslands ) vegetation cover .
below , the data about the habitat ( along with an abbreviation used throughout the study ) , coordinates , phytocoenosis , location , substrate , at every site are given .
3 ) płatkownica , sandy grassland ; 52°69 ' 02
n , 21°84 ' 53
e ; diantho - armerietum ; the phytocenosis occupies the raised , flat flooded terrace at the foot of the southern slopes of the flood embankment , within the base of the bug valley cut off from the inundation area ; desiccated river alluvial soils .
4 ) gnojno , xerothermic grassland ; 52°27 ' 59
n , 23°13 ' 88
e ; impoverished form of the adonido - brachypodietum arrhenatheretosum ; slope of the upper terrace , southern exposure and inclination of approx . 30° ; proper parendzina ;
5 ) mogielnica , xerothermic grassland ; 52°40 ' 08
n , 22°57 ' 04
e ; adonido - brachypodietum ; slope of the upper terrace , eastern exposure and inclination of approx . 30° ; proper pararendzina ;
6 ) morzyczyn , xerothermic grassland ; 52°68 ' 43
n , 21°91 ' 50
e ; tunico - poetum compresse ; the analyzed patch of the phytocoenosis evolved in an anthropogenic habitat , with a slightly alkaline ph . it is the southern slope of the flood embankment with an inclination of about 30° ; anthropogenic pararendzinas ;
7 ) płatkownica , xerothermic grassland ; 52°69 ' 00
n , 21°84 ' 43
e ; grassland with carex praecox of the agropyretea intermedio - repentis class ; southern slope of the flood embankment with an inclination of about 35 degrees ; anthropogenic pararendzinas .
11 ) płatkownica , rushes at oxbows ; 52°69 ' 21
n , 21°84 ' 54
e ; glycerietum maximae ; located on the edge of the shallow old riverbed of the bug river ; alluvial soils .
12 ) gnojno , thermophilous thickets ; 52°28 ' 19
n , 23°13 ' 47
e ; rhamno - cornetum sanguinei ; on the edge of the bug valley . the phytocoenosis habitat is a moraine slope with eastern exposure and inclination of approx . 30° ; leached brown soils ;
13 ) mogielnica , thermophilous thickets ; 52°40 ' 02
n , 22°57 ' 08
e ; rhamno - cornetum sanguinei ; on the edge of the bug valley . the phytocoenosis habitat is a moraine slope with eastern exposure and inclination of approx . 30° ; brown soils .
17 ) płatkownica , mesic meadow ; 52°69 ' 06
n , 21°84 ' 53
e ; poo - festucetum ; raised upper flooded terraces of the bug valley ; river alluvial soils .
18 ) zabuże , riparian forest ; 52°33 ' 33
n , 23°00 ' 86
e ; salicetum albae - fragilis ; n slope of the low flooded terrace in the patch of the riparian willow ; alluvial processes accumulate coarse - grained material of sand and river sediments . during periods of low water the initial alluvial soil may undergo significant desiccation ;
21 ) płatkownica , riparian forest ; 52°69 ' 19
n , 21°84 ' 06
e ; salicetum albae - fragilis ; flooded terrace ; proper alluvial soils .
24 ) płatkownica , wet meadow ; 52°69 ' 16
n , 21°84 ' 42
e ; violo - cnidietum ; upper flooded terrace of the bug valley , submerged regularly during the floods of the river ; river alluvial soils .
pitfall traps were used as a sampling method . an aqueous solution of propylene glycol ( about 50 % ) , containing a few drops of a detergent per 1 liter to reduce the surface tension of the fluid , was used as a preservation liquid . in each of the sites ten pitfall traps were placed in one straight line , at a distance of two meters one from another . the beginning of the trapping period was in the middle of march and the end was in the middle of november . the traps were replaced every two weeks .
the material analyzed in the current work was obtained during studies related to different groups of arthropods – including primarily spiders , carabids , diplopods and butterflies – under the project titled “the diversity of habitats and the biological diversity of selected groups of arthropoda in the bug valley” ( oleszczuk et al . 2011 , jastrzębski 2012 , hajdamowicz et al . 2014 ) . since chilopoda were not taken into account during the planning of the study , the specificity of this group of animals was not accounted for in the applied methodology .
in this work , standard methods and analysis indicators were applied : the shannon - weaver diversity index ( h ) , pielou’s measure of species evenness ( j ) , morisita index values as modified by horn , the cluster analysis – distance / similarity measure bray and curtis ; cluster method : nearest neighbor .
444 specimens belonging to 12 centipede species of two orders – geophilomorpha ( four species ) and lithobiomorpha ( eight species ) were caught ( table 1 ) .
lithobius ( monotarsobius ) dudichi – present in five habitats , prevalent in sandy grasslands ( 35 % of specimens ) and in xerothermic grasslands ( 32 % ) .
these species constitute 88 % of all centipedes caught during the study , thus establishing themselves as the most typical ones of almost all habitats of the study area . it is only in thermophilous thickets that a species from outside this group of four dominates – lithobius ( lithobius ) mutabilis ( table 1 ) .
interspecies occurrence similarity ( figure 2 ) [ the analysis does not include lithobius ( lithobius ) tenebrosus , as regrettably the information about the site on the label describing the specimen was completely obscured ] .
similarity of species ( distance / similarity measure bray and curtis ; cluster method : nearest neighbor ) . species nomenclature in table 1 .
in terms of ecology and zoogeography european eurytopic species prevail ( table 1 ) .
two to ten chilopoda species were found in each habitat under investigation . the greatest species richness was found in thermophilous thickets ( ten species ) , sandy grasslands ( eight species ) , xerothermic grasslands ( eight species ) and mesic meadows ( six species ) . the fewest number of species was found in rushes of reed mannagrass ( glyceria maxima ) and in wet meadows ( two species at each location ) ( table 1 ) .
the greatest number of specimens was found in the following habitats : riparian forests , mesic meadows and xerothermic grasslands ( table 1 ) .
the shannon - weaver diversity index ( h ) and pielou’s measure of species evenness ( j ) reached their highest values in thermophilous thickets , xerothermic grasslands , mesic meadows , while the lowest values – in rushes at oxbows and riparian forests ( table 1 ) .
the cluster analysis conducted on the basis of the species composition and dominance structure demonstrated the greatest similarity between communities of warm and dry habitats on the one hand , and wet and flooded – on the other ( figure 3 , table 2 ) .
dendrogram of the similarities of centipede composition in different habitats ( distance / similarity measure – bray and curtis , cluster method –nearest neighbor ) for designations see table 1 .
similarity of dominance structures – morisita index values as modified by horn . for designations see table 1 .
we note differences in the species composition and the number of chilopoda between the lower ( 102 specimens from 6 species ) and the middle section of the river valley ( 324 specimens from 11 species ) . this result reflects the differences in the structure and the vegetation of the two regions of the bug valley – especially the presence of thermophilous habitats in the middle section of the valley .
although several studies from european river valley areas have been conducted , this habitat is still poorly explored in terms of chilopoda . meanwhile natural and seminatural habitats associated with valleys of big rivers are already disappearing throughout the continent ( głowacki et al . 2002 ) .
this study is based on the materials obtained in studies on groups of arthropods other than chilopoda . this should explain the applied methodology , which does not take into account the specificity of chilopoda . this undoubtedly affected the results . pitfall traps limited the set of species to active epigeic forms . the expansion of the method to include quantitative soil samples and direct qualitative capture would contribute to a more complete picture of this group of chilopoda .
as noted by zulka ( 1999 ) and voigtländer ( 2011 ) , the one - year life cycle of lamyctes emarginatus is a strategy that allows this species to populate the same habitats as lamyctes curtipes – a species of a similar body size and probably very similar ecological requirements . adult specimens of the annual species lamyctes emarginatus appear in the environment in the summer and fall , when there is enough food to suffice for perennial species , such as lamyctes curtipes .
the riparian forest habitat is dominated by lamyctes curtipes – a species that prefers wet and humid habitats with high vegetation cover . this species , was found alive after 34 days of inundation ( adis and junk 2002 ) . in central europe it inhabits primarily riparian forests , alder swamp forests , river and brook sides , wet meadows with flooding and more rarely humid deciduous forests ( voigtländer 2005 ) . in the investigated area a small number of the specimens of this species was also collected from dry and xerothermic habitats ( table 1 ) . it can be assumed that these specimens have only immigrated from surrounding habitats , and they do not form stable populations in these habitats .
most species in the bug river valley inhabit thermophilous thickets – an environment that is already similar to forest habitats . this is clearly manifested by the composition of chilopoda community , in which we can find typically forest - dwellers – such as lithobius mutabilis or strigamia acuminata .
our results confirm the need to protect xerothermic habitats , unique almost throughout entire central europe , which due to their dispersion and their small area covered are fairly easily subject to the process of destruction . these environments are refuges for rare species of animals – including centipedes , as our research shows . lithobius dudichi presumably belongs to the relict xerothermic species of steppe provenance and is presumably in danger of extinction .
the results from our research in the bug valley also show that centipedes are a valuable indicator group for the assessment of habitat conditions . the information about the species composition of chilopoda communities , the dominance structure and their dynamics may thus be useful in characterizing specific location types .
we wish to extend our gratitude to karin voigtländer and ivan tuf for the extremely valuable comments to the first version of the manuscript .
m . stańska and i . hajdamowicz were supported by grant of siedlce university of sciences and humanities 222 / 05 / s .
terrestrial invertebrates inhabiting lowland river floodplains of central amazonia and central europe : a review .
[ tausend - und hundertfüßer ] ) an trockenheit und überflutung . – mitt .
impact of inundations on terrestrial arthropod assemblages in southern moravian floodplain forests , the czech republic .
even through the snowey winter i was able to find these guys under leaf litter .
based on around 43 pairs of legs ( not many northeastern soil centipedes have so few ! ) and the fact that the fangs extend forward past the edge of the head . this is an amazing species that often lives in the intertidal zone , eating barnacles , amphipods , worms etc . it can even survive several hours in salt water !
they often live outside of the intertidal zone , they just prefer moist areas . it ' s just that they ' re one of the few centipedes that can live on the shore .
moved from id request . one pair of legs per body segment = centipede .
that ' s good basic info . thank you ! i ' m learning so much more than i thought i could about ecology since getting into macro photography ! everyone here at te guide are the best ! : )
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