drosophila subobscura is a species of fruit fly from the family drosophilidae .

originally found around the mediterranean , it has spread to most of europe and the near east .

it has been introduced into the west coasts of canada , the united states , and chile .

its closest relative is drosophila madeirensis , found in the madeira islands , followed by d. guanche , found in the canary islands .

these three species form the paraphyletic drosophila subobscura species group .

drosophila subobscura

drosophila subobscura <EOT> solved : in drosophila subobscura , the presence of a recessive g . . . | urltoken

introduction history of drosophila subobscura in the new world : a microsatellite - based survey using abc methods .

introduction history of drosophila subobscura in the new world : a microsatellite - based survey using abc methods . - pubmed - ncbi

asymmetry of overall wing shape for females raised at 23°c same as in table 5 for drosophila subobscura flies reared at 23°c .

asymmetry of overall wing shape for males raised at 23°c same as in table 5 for drosophila subobscura flies reared at 23°c .

krimbas cb : drosophila subobscura : biology , genetics and inversion polymorphism . 1993 , hamburg , germany : verlag dr . kovac

llopart a , aguadé m ( 2000 ) nucleotide polymorphism at the rpii215 gene in drosophila subobscura : weak selection on synonymous mutations . genetics 155 : 1245–1252

additional file 3 : physical map of differentially expressed genes . localization by in situ hybridization on the salivary gland chromosomes of drosophila subobscura of 88 differentially expressed genes . ( pdf 77 kb )

gilchrist gw , huey rb , serra l : rapid evolution of wing size clines in drosophila subobscura . genetica . 2001 , 112–113 : 273 - 286 . 10 . 1023 / a : 1013358931816 .

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in drosophila subobscura , the presence of a recessive gene called grandchildless ( gs ) causes the offspring of homozygous females , but not those of homozygous males , to be sterile . can you offer an explanation as to why females and not males are affected by the mutant gene ?

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. drosophila inversion polymorphism . edited by : krimbas cb , powell jr . 1992 , boca raton : crc press , 127 - 220 .

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comparisons of recent with historical samples of chromosome inversion frequencies provide opportunities to determine whether genetic change is tracking climate change in natural populations . we determined the magnitude and direction of shifts over time ( 24 years between samples on average ) in chromosome inversion frequencies and in ambient temperature for populations of the fly drosophila subobscura on three continents . in 22 of 26 populations , climates warmed over the intervals , and genotypes characteristic of low latitudes ( warm climates ) increased in frequency in 21 of those 22 populations . thus , genetic change in this fly is tracking climate warming and is doing so globally .

the results strongly support the hypothesis that environmental canalization and developmental stability share underlying regulatory mechanisms , but environmental and genetic canalization are not functionally the same . a likely explanation for this lack of association is that natural wing shape variation in drosophila populations is loosely related to individual fitness .

seasonal changes in field temperature and chilling tolerance of d . subobscura living in close association with a heap of discarded apples . ( a ) air temperature ( light grey ) and the temperature at the bottom ( black ) and top ( i . e . surface ; dark grey ) of the heap of apples were recorded during the transition from spring to summer . adult d . subobscura were collected approximately every four weeks ( circles with vertical lines ) . ( b ) ct min and ( c ) ccrt measured in flies collected during the experimental period ( mean ± s . e . ) regressed against mean temperature at the surface of the heap in the 24 h preceding collection . shaded areas represent 95 % confidence intervals of regressions . ( online version in colour . )

andres aj , thummel cs : methods for quantitative analysis of transcription in larvae and prepupae . drosophila melanogaster : practical uses in cell and molecular biology . methods in cell biology . edited by : goldstein l , fyrberg e . 1994 , new york : academic press , 44 : 565 - 573 .

we thank david salguero for his excellent technical assistance . we also thank servei de genòmica , serveis cientifico - tècnics , universitat de barcelona , for automated sequencing facilities . this paper was prepared with full knowledge and support of the barcelona subobscura initiative ( bsi ) . this work was supported by grants bfu2012 - 35168 and bfu2014 - 63732 from ministerio de economía y competitividad , spain , and 2014sgr - 1055 from comissió interdepartamental de recerca i innovació tecnològica , generalitat de catalunya , spain to ma .

our data suggest that a sizeable number of genes appear to be involved in thermal adaptation in drosophila , with a substantial fraction implicated in metabolism . this apparently illustrates the formidable challenge to understanding the adaptive evolution of complex trait variation . furthermore , some clustering of genes within inverted chromosomal sections was detected . disentangling the effects of inversions will be obviously required in any future approach if we want to identify the relevant candidate genes .

hl sampled the thermal populations , made the rna extractions , participated in the design of the experiment , carried out statistical analysis , gene ontology searches , and drafted the manuscript . fg - f , bec - s and vt designed primers , carried out in situ hybridizations on the polytene chromosomes of d . subobscura , and read all salivary gland squashes from the in situ hybridizations . sb and mc coordinated the microarray experiments for generating the original data . ms set up and maintained the thermal populations , designed the study , carried out statistical analyses and drafted the final version of the manuscript . all authors read and approved the final manuscript .

water and ion balance in d . subobscura before ( i . e . control ) and immediately after 24 h at −5°c on five dates during the spring–summer transition . ( a ) haemolymph volume , ( b ) haemolymph [ na + ] and ( c ) haemolymph [ k + ] measured in flies before and after cold exposure . the net change in haemolymph [ k + ] with cold exposure ( δk + ) was significantly associated with the net change in haemolymph volume ( d ) , as well as both indices of chilling tolerance : ( e ) the ct min and ( f ) ccrt . shaded areas represent 95 % confidence intervals of regressions . all values are means ( ±s . e . ) . haemolymph volume was not measured ( nm ) on july 21st . ( online version in colour . )

subsequently , samples from all nine populations were obtained in may - june 2004 ( 25 generations at 13°c , 35 at 18°c , and 46 at 22°c ) by placing eggs into twelve 130 - ml bottles ( ~ 200–250 eggs per bottle ) . these bottles were cultured at 18°c and emerging adults were dumped into plexiglas cages for egg collections . eggs for the experiment were collected over a six days period by placing petri dishes containing non - nutritive agar with a generous smear of live yeast in the cages . as before , ~ 100 eggs ( ± 4 h ) were placed at 18°c in 120 - ml plastic chambers with stained drosophila medium to sample the treatment third instar larvae for further mrna extraction .

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pascual m 1 , chapuis mp , mestres f , balanyà j , huey rb , gilchrist gw , serra l , estoup a .

departament de genètica , facultat de biologia , universitat de barcelona , diagonal 645 , 08028 barcelona , spain . martapascual @ urltoken

research support , u . s . gov ' t , non - p . h . s .

many properties of organisms show great robustness against genetic and environmental perturbations . the terms canalization and developmental stability were originally proposed to describe the ability of an organism to resist perturbations and to produce a predictable target phenotype regardless of random developmental noise . however , the extent to which canalization and developmental stability are controlled by the same set of genes and share underlying regulatory mechanisms is largely unresolved .

phenotypic robustness refers to the invariance of the specified target phenotype given the genetic makeup and environmental conditions . whereas the presence of naturally occurring phenotypic variation is at the core of evolutionary biology , developmental geneticists have traditionally considered it as a nuisance . instead , they have relied on the study of single or multiple mutant combinations to reveal the generation of phenotypic patterns ( e . g . [

] , where development appears to be buffered so that slight abnormalities of genotype or slight perturbations in the environment do not lead to the production of abnormal phenotypes . however , evolutionary geneticists define canalization as the tendency of traits to evolve a reduction in variability [

] . therefore , canalization and ds are subcategories of developmental buffering : the first can be appraised by estimating interindividual variance whereas the most commonly used estimate of ds in bilaterally symmetrical organisms is fluctuating asymmetry ( fa ) ; i . e . the intraindividual variation due to random differences between left and right sides . the question of whether or not canalization and ds are different buffering mechanisms has been a constant source of debate . two recent reviews implicitly [

] assume that ds is a special case of canalization , a viewpoint also embraced by several authors ( e . g . [

] found that the vectors describing inter - and intraindividual variation of landmark position for fly vein traits were highly concordant . on the other hand , debat et al . [

] came to the opposite conclusion applying the same methods to cranial landmarks in the house mouse – although klingenberg ' s et al . [

] work with mouse mandibles found patterns of intra - and interindividual variation that were only partly consistent – . at first glance , the different results may suggest that the mechanisms that affect canalization and ds are related in some developmental contexts but not in others . the problem is , however , that according to the causes of phenotypic variation a distinction between genetic and environmental canalization is necessary [

) : the first refer to influential factors ( e . g . temperature ) that are shared by groups of individuals , whereas the latter are residual deviations from the phenotype that would be specified on the basis of genotype and general environmental effects . such deviations are unique to individuals and are largely unpredictable . the variance associated with special environmental effects can be estimated when experiments are performed on completely inbred lines ( i . e . , there is no genetic variance ) . in bilaterally symmetrical organisms it is also feasible to estimate the two sources that contribute to those special environmental effects : among - individual (

the third process involved in the control of phenotypic variability is plasticity , which can be defined as the ability of an individual to express one phenotype under one set of environmental circumstances and another phenotype under another set . the expressed phenotypes can be discontinuous thus eliciting discrete morphs ( i . e . , polyphenism ) , or there can be a continuous range of potential phenotypes ( i . e . , reaction norm ) . the reaction norm is thus a property of the genome : genetic canalization and phenotypic plasticity are not mutually exclusive and can combine to form canalized reaction norms [

] , and the vein pattern is highly conserved across species ( e . g . [

] . however , there is circumstantial evidence suggesting that developmental and evolutionary temperature - related cell size divergence have contrasting effects on wing shape . thus , birdsall et al . [

] ) for which a balancer stock is available . in colonizing populations of the new world only six gene arrangements are segregating for that chromosome : o

] . therefore , six independent isochromosomal lines for each of these three chromosome arrangements ( i . e . ,

= st , 3 + 4 , 3 + 4 + 7 ) were used in the present experiments .

the experimental flies were obtained from 54 crosses , which will be referred to as inbred ( isogenic ; i . e . ,

) with 18 crosses in total , or outbred ( including both structural homo - and heterokaryotypes ) with 36 ( 18 + 18 ) crosses in total . the six lines with a given gene arrangement were crossed to produce the three different outbred homokaryotypes ( i . e . ,

) . the three kinds of heterokaryotypic flies were similarly obtained but using lines with different gene arrangements ( i . e . ,

) . since all isochromosomal lines were homogeneous for the same genetic background ( except for the male sex chromosome ) , maternal effects were not considered to be critically important . anyhow , experimental flies were randomly derived form reciprocal crosses for all outbred combinations . two developmental temperatures were used in the experiment : optimal ( 18°c ) and warm ( 23°c ) .

ranging from 0 . 032 for inbred females at 18°c to 0 . 073 for inbred males at 23°c ;

= 0 . 030 ( - 0 . 005 , 0 . 064 ) for inbred females at 23°c ) , thus suggesting independence between size and size fa .

) . size variation ( cs : centroid size ) among individuals comprised the largest part ( > 90 % ) of the variation . the fraction of the total phenotypic variance in wing size associated to genetic differences among karyotypes and / or lines ( i . e . ,

) ranged from 0 . 235 ( inbred males at 18°c ) to 0 . 602 ( inbred females at 23°c ) . ( bear in mind that there is nothing in the anova method of estimation that will prevent a negative variance estimate [

flies raised from inbred ( isogenic ) and outbred crosses reared at 18°c . centroid size ( cs , estimated in a normalized form [ 22 ] ) is the dependent variable ( values in pixels

) provide here unbiased estimates of the among - fly ( i . e .

asymmetry of overall wing size for males raised at 18°c same as in table 1 .

inbreeding and temperature effects on size homokaryotipic averages for centroid size and centroid size fa ( index fa1 in [ 39 ] ) in inbred ( black symbols ) and outbred ( open symbols ) crosses . small symbols give the average values for each of the three different homokaryotypes to appreciate the dispersion from the corresponding grand average ( large symbols connected by lines ) . squares give the values at 23°c and circles at 18°c .

] a quite remarkable finding here was that left wings were consistently bigger than the right ones , thus causing a generally highly significant da ( i . e . , " sides " effect in tables

] . when wings were mounted no attempt was made to standardize the surface position : in females 394 ( 60 . 8 % ) left and 387 ( 59 . 7 % ) right wings were mounted on the slides with the dorsal side up (

= 0 . 691 ) ; in males the corresponding figures were 383 ( 59 . 1 % ) and 401 ( 61 . 9 % ) , respectively (

= 0 . 306 ) . potential biasing effects when measuring wings ; namely , dorsal or ventral bitmap images or possible differences between left and right wings when bitmap images are captured from the top or bottom of the microscope slide , were checked from a subset of 75 females and 75 males . an additional set of two images for each wing were taken in the same session from the top and bottom of the slide and digitized once . the centroid size differences between the averages of both measurements was apparently random with respect to digitizing procedure and always lower than 0 . 07 % , whereas left wings were 0 . 26 % bigger than the right ones in females and 0 . 34 % in males . we are , therefore , quite confident that the fairly subtle da for wing cs is not an experimental artifact but a real phenomenon .

in addition to da , there was subtle but significant fa in all crosses ( i . e . , " individuals × sides " interaction effect in tables

) together with a small amount of genetic variation for da in some of them . this last finding could hardly be attributable to a type i error because similar results had been previously obtained [ [

] ) failed to show any statistically significant effects whatsoever ( variance components ranging from - 0 . 0047 to 0 . 0071 for karyotypes , and from - 0 . 0343 to 0 . 0406 for crosses within karyotypes ; values in pixels

( pixels ) , but results were qualitatively identical without a log - transformation ) as the dependent variable , with karyotype , temperature and inbreeding as fixed effects , and crosses nested within karyotypes , clearly indicated inbreeding depression together with temperature by inbreeding interaction ( i . e . , inbreeding was most noticeable at the sub - optimal temperature of 23°c ) , but no karyotype by temperature interaction was detected . these results confirm that wing size is not a purely additive trait in

both inbreeding and ( sub - optimal ) temperature effects were also apparent in females when overall size fa ( index fa1 ) was used as the dependent variable in three - way factorial anovas , with no differences among karyotypes . on the other hand , no statistically significant effects were detected for males , basically because inbred crosses performed approximately equal at both temperatures ( fig .

it is worth mentioning here that in outbred crosses overall size fa was about the same for homokaryotypes and heterokaryotypes : the only significant effect was again an increase in fa at the sub - optimal temperature ( more than two - fold ; c . f .

in conclusion , overall size ds was positively correlated with levels of heterozygosity ( i . e . , inbred vs . outbred homokaryotypes ) and development at the optimal temperature of 18°c . however , no positive association was found between ds and chromosomal heterozygosity in outbred crosses .

. for the present study of 13 landmarks , with 2 coordinates each , the shape dimension is 22 . sums of squares and cross - products ( sscp ) matrices are therefore not full - ranked , and we retained 22 pc ( principal components [

] ) scores for outbred crosses and only 15 pc scores – which accounted for more than 98 % of the total shape variance – for inbred crosses to be capable of testing for genetic components . the degrees of freedom in tables

) times the number of pc scores retained in each sample . likewise , the overall covariation in wing shape ( " individuals " effect ) was decomposed into causal components ( karyotypes , crosses in karyotypes , and among flies ) ; and the overall covariation in wing shape fa ( " individuals × sides " interaction effect ) was decomposed into causal components attributable to wing shape da ( karyotypes , crosses in karyotypes , and within flies ) .

reared at 18°c . for the inbred crosses 15 pc scores were retained for analyses ( proportion of total shape variance accounted is given in parenthesis ) . for the outbred crosses 22 pc scores were retained . (

asymmetry of overall wing shape for males raised at 18°c same as in table 5 .

similarly to what had been found for cs , differences between left and right wings were also highly significant ( " sides " effect ) , thus indicating that da was present for overall wing shape . this finding is contrary to our previous claim from a subset of o

isochromosomal lines , where da for some landmarks ( e . g . those defining the position of the anterior crossvein ) but not for overall wing shape was detected [

] . after plotting the procrustes grand mean shapes of both wings it also became apparent here that the location of the anterior crossvein was indeed slightly more distal in the right wings . furthermore , the individuals × sides interaction effects were highly significant in all cases and , hence , wing shape fa greatly exceeded measurement error .

] shape is an inherently multidimensional concept and cannot be easily reduced to a scalar index without severe loss of information . therefore , for a quantitative genetic analysis of shape data a multivariate approach is required [

are , respectively , the covariance matrices for karyotypes , crosses within karyotypes , and the residuals .

shows the amount of variation associated with the different dimensions in shape space . much of the variation was concentrated in the first few pcs , but the

matrices showed the clearest trend to quickly decrease after the first pc . permutation tests indicated that matrix correlations ( mcs ) between

= 0 . 2665 ) , but none of the mcs was statistically significant . on the other hand , vcv matrices were correlated across rearing temperatures ( females : mc

) covariance matrix onto the total variance of the phenotypic covariance matrix was lower at 18°c in both sexes ( females : 0 . 1312

eigenvalues of causal covariance matrices for wing shape first 15 eigenvalues of the phenotypic ( black bars ) , karyotype ( hatched ) and crosses ( open ) covariance matrices from outbred crosses .

also point to the presence of genetic variation for overall shape da , mainly at 18°c ( i . e . , the " crosses in karyotypes " component from the decomposition of the i × s interaction effect ) . as far as we are aware , these are the first experiments that found detectable genetic variation in da for wing traits . the uncovering of da ( i . e . , " side " effect ) for fly wings is quite general when quantitative analyses of form are carried out using the powerful methods of geometric morphometrics to reveal even small morphological variation that otherwise would remain hidden with less effective techniques [

] because it provides compelling evidence that da in fly wings may signal the presence of genetic variation in a phylogenetic conserved left - right developmental axis ( i . e . , an imaginary plane between the two lateral sides of the body ) , as discussed by klingenberg et al . [

] . actually , modern treatises in developmental biology ( e . g . [

] ) distinguish the left - right axis besides the customary anterior - posterior and dorsal - ventral axes , and several asymmetrically expressed genes ( e . g .

] were the first ( and to our knowledge the only ones ) who showed a developmental mechanism for the developmental asymmetry . it seems , therefore , that the detection of genetic variation for da in this genus appears to be basically a methodological problem , including statistical power and the environmental conditions where the experiments are performed . the mechanisms that constitute the genetic basis of morphological asymmetry in

] a multivariate equivalent of fa1 ( i . e . , the " unsigned " left - right differences ) was defined by changing the signs of all coordinate differences ( from left - right to right - left ) whenever the inner product ( also referred to as the dot product ) of a left - right difference vector with the vector of mean left - right difference was negative . for the univariate case ( cs ) this procedure would render here the absolute (

) . however , the approach used to define the multivariate equivalent of fa1 might be influenced by the arbitrary choice of the plane ( i . e . , the mean left - right differences ) to subdivide the shape space into " positive " and " negative " halves ( christian p klingenberg , pers . comm . 2004 ) . a modified procrustes shape distance for non - isotropic variation ( i . e . , landmarks usually differ in their amounts of variation ) has been recently developed by klingenberg and monteiro [

] , and can be used here as a scalar measure of the amount of shape asymmetry because fa is random in origin ( i . e . , only the magnitude and not the direction may usually be the interesting component of fa shape variation ) . when this scalar was used in our data set the same conclusion was obtained ; namely , there was no detectable genetic variation for wing shape fa in any case ( results not shown ) .

manovas for female wing shape fluctuating asymmetry a multivariate equivalent of fa1 ( i . e . , the " unsigned " left - right differences ) was defined as explained in the text . flies raised from outbred crosses of

to investigate allometric and nonallometric temperature effects on overall wing shape we performed a multivariate analyses of covariance ( mancova ) of the procrustes coordinates ( after averaging both sides and the two replicated measurements per side ) considering temperature and inbreeding ( i . e . , isogenic

( pixels ) ) as the covariate . temperature effects were only significant in males , but inbreeding and temperature × inbreeding interaction effects were highly significant in both sexes ( results not shown ) , which suggests a strong effect of the categorical predictors on the nonallometric component of shape . size effects were also found to be significant ( females : wilks '

< 0 . 001 ) . the association between size and temperature ( fig .

] ) , was found to be lower than the suggested guideline for serious collinearity ( i . e .

≥ 10 ) , which indicates that the effects of temperature and size on wing shape could be effectively separated .

. outbred homokaryotypic flies ) on wing shape fa were tested from the ratio between the traces of the corresponding " individual × side " vcv matrices . notice that the traces of these interaction matrices are equal to the respective mean squares of the procrustes anova as implemented by klingenberg and mcintyre [

coordinates of the corresponding aligned configurations divided by the shape dimension . we performed 10 , 000 randomization runs for each test . inbreeding effects were detected at 18°c but only in females ( 18°c : female

principal component analyses were only implemented for the outbred crosses since they are more representative of the natural situation . the percentages of total shape variation , together with the features of variation associated with the dominant pcs , are graphically plotted in figs .

. for the individual variation several pcs accounted for relatively large amounts of variability . on the contrary , for fa and measurement error pc1 explained almost all total variance ( > 80 % ) . for all levels in the analysis ( i . e . individuals , fa and measurement error ) the dominant pcs were connected to the relatively large variability of landmarks 3 , 6 , 7 and , to a lesser extent , landmark 2 . however , the disproportionate amount of variation associated with these landmarks did not spread to all sources of causal variation because their coefficients were relatively small for the pc1 of karyotype variation ( which explained ~ 60 % of the total variance ; see below ) . furthermore , for the individual variation the first two pcs were also linked to the shift of the anterior ( landmarks 11 and 12 ) and posterior ( landmarks 7 and 13 ) crossveins along the adjoining longitudinal veins .

vectors of the landmarks displacements first two axes of wing shape variation for each effect in the two - way mixed manova ( individuals , individuals × sides interaction , and measurement error ) for females from outbred crosses reared at 18°c . also plotted are the percentages of total wing shape variation explained by the principal components for the corresponding covariance matrices .

vectors of the landmarks displacements same as fig . 3 for males from outbred crosses reared at 18°c .

vectors of the landmarks displacements same as fig . 3 for females from outbred crosses reared at 23°c .

vectors of the landmarks displacements same as fig . 3 for males from outbred crosses reared at 23°c .

permutation tests indicated that vcv matrices were mostly correlated for fa and measurement error effects within samples ( mcs > 0 . 95 ,

) . the individual vcv matrix was significantly correlated with the fa and measurement error matrices only for females at 18°c . between temperatures the vcv matrices were highly correlated for fa and measurement error ( results not shown ) , but loosely correlated for the individual variation ( females mc = 0 . 668 ,

correlations between vcv matrices of landmarks displacements within groups results of the permutation tests used for the analyses within sexes and temperatures .

the angles between the pc1s for fa and measurement error were very much alike ( ranging from angle α = 2 . 1° to α = 3 . 4° ; recall that the 0 . 1 % quantile of the resulting distribution between pairs of random vectors in 22 - dimensional space was 50 . 3° ) , which reflects the similarity due to landmarks 3 , 6 and 7 . however , the first three pcs for interindividual variation were generally distinct to those of fa : the only clear correspondences were between the pc1s for females at 18°c ( α = 21 . 5° ) , and the pc2 of interindividual variation with the pc1 of fa for males at 18°c ( α = 11 . 8° ) . ( the correspondences were qualitatively the same for interindividual variation and measurement error ; results not shown . ) overall , these results seem to suggest that canalization and ds do not generally share the same underlying regulatory mechanisms ( but see below ) .

a potentially important problem with the foregoing approach to compare the patterns of intra - and interindividual variation is to rely on the interaction vcv matrix as the source of variation due to fa . as has been previously argued the uncovering of da is almost ubiquitous for shape data when using the methods of geometry morphometrics , and there was evidence here for statistically significant genetic variation of overall shape da at 18°c ( tables

) . therefore , the vcv matrix from the " individuals × sides " interaction effect gives a biased estimate of developmental stability and cannot be taken as the covariance matrix for fa . in other words , this vcv matrix also includes all causal components due to genetic variation for da , and the corresponding unbiased vcv matrix for fa is that for the within - fly component of the interaction effect ( i . e . , after removing the genetic variation for da [

] ) . in any case , all results were qualitatively similar and , hence , the conclusion that canalization and ds seem to be different mechanisms remains unchanged . however , it is difficult to appraise how this potential problem could have affected the previously published conclusions when comparing interindividual variation and " fa " in fly wings and mouse skulls ( see background section ) .

= 49 . 6° ; recall that the direction of pcs is arbitrary and all the movements in figs .

besides the interindividual variation in the two - way manovas ( which comprises genetic plus environmental covariances due to special environmental effects ) it is important here to asses the patterns of joint displacements of landmarks for each of the causal components of wing shape variation ( figs .

) . for karyotype variation pc1 accounted for ~ 60 % of the total variance and was linked to a great extent with equivalent movements of those landmarks defining the location of the crossveins , which shifted in the same direction . landmarks 4 and 5 tended to move away each other , stretching the wing margin between longitudinal veins iii and iv . landmark 9 budged in the opposite direction to crossveins shifts , thus shaping the relationship between l1 to the total length of longitudinal vein iv ( i . e . shape index l1 wl ; fig .

vectors of the landmarks displacements first two axes of wing shape variation in the two - level nested manova ( karyotypes , crosses nested in karyotypes , and within crosses ) for each causal component effect pertaining to the inter - individual variation in females from outbred crosses reared at 18°c . also plotted are the percentages of total wing shape variation explained by the principal components for the corresponding covariance matrices .

vectors of the landmarks displacements same as fig . 7 for males from outbred crosses reared at 18°c .

vectors of the landmarks displacements same as fig . 7 for females from outbred crosses reared at 23°c .

vectors of the landmarks displacements same as fig . 7 for males from outbred crosses reared at 23°c .

the image shows the thirteen landmarks ( 1 – 13 ) used in this work . i – vi longitudinal veins ; cv - a and cv - p anterior and posterior crossveins ; co costal or marginal veins ; l1 and l2 lengths of the proximal ( euclidian distance between landmarks 9 and 13 ) and distal ( euclidian distance between landmarks 13 and 5 ) segments of longitudinal vein iv , respectively . wing shape index

has been previously used to study shape clines in this species [ 30 ] .

) , but rearing temperature quantitatively modified the shape index ( l1 / wl was lower at the highest temperature ) . however , there was no statistically significant karyotype × temperature interaction . the wing shape index appears to be a purely additive trait since heterokaryotypes were always intermediate to their corresponding homokaryotypes ( fig .

) . actually , none of 12 within - group ( i . e . , sex and temperature ) possible contrasts comparing all three heterokaryotypes with the average of the corresponding homokaryotypes was statistically significant ( the mean square for " crosses " was used as the error term ; see legend in fig .

averages of the relative length ( with 95 % confidence intervals ) of the basal portion of longitudinal vein iv ( l1 ) to the total wing length ( wl = l1 + l2 ) versus karyotype for outbred crosses at the two rearing temperatures . a two - way factorial anova using the shape index as

pc2 for karyotypes was also connected to the variability of landmarks 3 , 6 and 7 . for the crosses component , several pcs explained relatively large amounts of variation , and shifts of crossveins now seem to be independent of each other at 18°c but not at 23°c . finally , for the within - fly variation several pcs accounted for relatively large amounts of variability . pc1s were again connected to the variability of landmarks 3 , 6 and 7 ; and pc2s to shifts in the anterior crossvein .

the large amount of variation of the anterior and posterior crossveins for karyotypes and crosses can be interpreted in terms of developmental processes . the crossveins are determined after the longitudinal veins , and mutations that eliminate crossveins ( e . g .

) do not affect the longitudinal veins ; however , some mutants that affect the longitudinal veins also influence the crossveins ( e . g . the

] ) . however , these shifts do not occur in isolation an also include other landmarks as well ( e . g . , landmarks 9 and 5 on l4 ; landmarks 1 and 2 on l1 ; figs .

) indicated that the vcv matrices of karyotypes and crosses were never significantly correlated with the vcv matrices of fa and measurement error . the high correlation between the vcv matrices of the interindividual and fa effects for females at 18°c was basically due to the ( micro - ) environmental component . also notice that all correlations between the vcv matrices of karyotype and fa effects were close to zero or even negative , which clearly suggests that this genetic component of canalization is unrelated to ds .

in addition , the pc1s of karyotypes and fa were nearly at right angles ( 18°c : females α = 85 . 8° , males α = 77 . 3° ; 23°c : females α = 75 . 6° , males α = 78 . 5° ) . the only matches were between pc2 of karyotypes and pc1 of fa for females at 18°c ( α = 13 . 0° ) and males at 23°c ( α = 31 . 2° ) . the pc1s of crosses and fa were also poorly correlated ; the only exception being females a 23°c ( α = 43 . 3° ) . these results clearly support the hypothesis that genetic canalization and ds are not functionally the same mechanism .

on the other hand , all observed angles involving pc1s between " replicated genotypes " ( i . e . the between - fly component ) and fa were relatively small and highly significant ( 18°c : females

= 36 . 7° ) . ( results were qualitatively the same for all observed angles involving pc1s of the between - fly and measurement error covariance matrices ; results not shown . ) together with the overall comparisons of the covariance matrices ( table

) , these results indicate that ( micro - ) environmental canalization and ds share underlying regulatory mechanisms but are not identical . there was not a complete congruence as pc1 of fa accounted for most part of the variation , while pc1 of between - fly variation usually explained less than 50 % of the total variance ( figs .

in addition , ( vii ) a discrepancy between sexes was observed in some situations ; e . g . overall size fa increased with inbreeding and ( sub - optimal ) temperature effects mainly in females , and the allometric effect on wing shape at both experimental temperatures was similar in females but not in males . it is also interesting to note here that wing size ( measured as wl ; fig .

] . what is not obvious , however , is why there is a difference between the sexes .

] . the strongest evidence in favor of this hypothesis comes from the well - known genotype - phenotype mapping of rna folding . conservation of rna secondary structure is under strong selection , and low structural plasticity is achieved through increasing the thermodynamic independence of any one structural component from the remaining structure [

are adaptive . there are no consistent patterns between latitude and wing shape ( e . g . [

] ) , contrarily to what happens for size - related traits where world - wide latitudinal clines are found with genetically larger individuals derived from higher latitudes ( e . g . [

] ) , and we have shown here that the wing shape index l1 / wl appears to be a purely additive trait since heterokaryotypes were always intermediate to their homokaryotypic counterparts . the wing shape cline in north america colonizing populations of

] , and the small shifts of ( e . g . ) the anterior and posterior crossveins in relation to karyotype variation ( figs .

; notice that the plotted joint variation in landmark positions is an exaggeration of the actual variation in the data set ) are difficult to link with any adaptive response to a better flight capacity . actually , we lack even hypothetical functional explanations for subtle shape variation : gilchrist et al . [

may simply represent drift around an optimum . our present results ( points ( v ) and ( vi ) above ) give some credence to that conjecture . genetic canalization on wing shape does not seem to arise as a by - product of environmental canalization and , therefore , canalization is not a single mechanism to buffer any source of variation as has been suggested [

] the classical linear theory of ds can successfully account for both normally distributed error distributions and leptokurtic distributions caused by the admixture of individuals having different levels of ds , but cannot account for transitions between fa and da . we have previously suggested , however , that a transition from " ideal " fa ( i . e . , a normal distribution of left – right - side scores whose mean is zero ) to a distribution showing da could be made entirely compatible with what it is already known from classical quantitative genetics [

] , but unless the genetic component can be partitioned out the variation in left - right differences cannot be assumed to describe ds . from the results of outbred crosses reared at 18°c ( table

) it is possible to test here for the congruence between patterns of morphological variation with respect to the variation attributable to fa ( i . e . the within - fly environmental component of the interaction term ) and that attributable to genetic variation for da ( the within - fly genetic component due to crosses in karyotypes of the interaction term ) . the corresponding vcv matrices were highly correlated for females ( mc = 0 . 914 ,

= 0 . 0001 ) but not for males ( mc = 0 . 064 ,

= 45 . 9° ) . when considered together , these results clearly suggest that fa and genetic variation for da may or may not be functionally linked .

females , and its arrangement on the wild - type chromosome was identified after four generations of backcrosses . followed by at least another backcross to

) on the o chromosome . the isochromosomal lines were established from the final crosses ♀♀ o

] . the lines were kept at 18°c ( 12 : 12 light / dark cycle ) in 130 - ml bottles with low adult density to standardize the rearing conditions before egg collections .

as previously indicated the experimental flies were obtained from 54 crosses . reciprocal crosses were made for all outbred combinations by mating one - week virgin females and males . after three days the males were discarded and equal numbers of females from each reciprocal cross were placed together in a plastic chamber with a spoon containing non - nutritive agar with a generous smear of live yeast for egg collections . to standardize the experimental conditions , eggs from the inbred ( isogenic ) crosses were also obtained in a similar way ; namely , after mating the flies in bottles and transferring the females to plastic chambers . eggs were placed in six 2 × 8 cm vials with 6 ml of food ( 26 eggs / vial ) ; three vials were kept at 18°c ( optimal temperature ) and the other three at 23°c ( sub - optimal temperature ) . within each experimental temperature the vials were randomly placed on the same incubator shelf . as a result , the total experiment consisted of 324 vials ( 162 vials at each experimental temperature ) , and all eggs were sampled on the same day . emerging flies ( not less than 2 or 3 days old ) were stored in eppendorf tubes with a 3 : 1 mixture of alcohol and glycerol at 4°c before wing measurements .

all fly handling was done at room temperature using co 2 anesthesia on flies not less than 6 h after eclosion .

two randomly sampled females and males emerged from each vial were used for morphometric analyses . both wings were removed from each fly and fixed in dpx under coverslips on microscope slides . bitmap images were captured with a video camera ( sony ccd - iris , tokyo , japan ) connected to a pc computer with mgi videowave software and mounted on a compound microscope ( zeiss axioskop , jena , germany ) , using a 2 . 5 × objective . calibration of the optical system was checked at each session . the images were stored on a dell workstation pws350 . to quantify and minimize measurement error all wings were digitized two times at different sessions as follows : images of both the left and right wings were captured during a given session and after an entire round on all individuals the same process was repeated again . a similar procedure was also used to record the

coordinates of 13 morphological landmarks ( i . e . , labeled geometric points located at the intersections of wing veins or at sites where veins reach the wing margin ; fig .

] . therefore , the process we used guaranteed that the observer was blind with respect to the results from previous measurements .

geometric morphometrics precisely separates morphological variation ( i . e . , variation in form ) into size and shape components [


it is found in spain .

the wingspan is 29 – 31 mm .

adults are on wing from september to october .

the larvae probably feed on biscutella species .

evergestis merceti

evergestis merceti <EOT> evergestis merceti is a species of moth in the crambidae family . it is found in spain .

evergestis renatalis ; rothschild , 1915 , novit . zool . 22 ( 2 ) : 189

evergestis sophialis lupalis zerny , 1928 ; zs . öst . entver . 13 : 50 ; tl : spain

evergestis zernyi schawerda , 1924 ; verh . zool . - bot . ges . wien 73 ( s . b ) : 163

evergestis nolentis heinrich , 1940 ; proc . ent . soc . wash . 42 ( 2 ) : 36 , pl . 6 - 7 , f . 4 - 4b , 8 ; tl : san felipe wash , san diego co . , california

evergestis - species dictionary - global : ispot nature - your place to share nature . ispot is a website aimed at helping anyone identify anything in nature . once you ' ve registered , you can add an observation to the website and suggest an identification yourself or see if anyone else can identify it for you .

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the id resolving service for stable taxon ids is currently under maintenance . meanwhile , please use the name search in order to find the taxon page .

museum für naturkunde leibniz - institut für evolutions - und biodiversitätsforschung invalidenstr . 43 10115 berlin germany e - mail : fauna - eu ( at ) mfn - berlin . de website : urltoken

encyclo . co . uk , online since 2007 , is a search engine for english meanings and definitions . the website aims to publish all wordlists , big and small , on the internet , making it much easier to find the word you need . follow us on facebook

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umbrosalis ( fischer von röslerstamm , 1842 ) ; abbildungen schmettkde : 274 , pl . 92 , f . 2

pyralis desertalis hübner , [ 1813 ] ; samml . eur . schmett . [ 6 ] : f . 171 : europe

noctuelia desertalis ; rothschild , 1915 , novit . zool . 22 ( 2 ) : 190

752x547 ( ~ 90kb ) russia , moscow area , 12 . 06 . 2007 , photo © d . smirnov

phalaena extimalis scopoli , 1763 ; ent . carniolica : 241 , f . 614

523x600 ( ~ 81kb ) russia , moscow area , 16 . 7 . 2006 © d . smirnov

politalis ( denis & schiffermüller , 1775 ) ; ank . syst . schmett . wienergegend : 121

subfuscalis ( staudinger , 1871 ) ; horae soc . ent . ross . 7 ( 1870 ) : 192 , pl . 2 , f . 9

aenealis ( denis & schiffermüller , 1775 ) ; ank . syst . schmett . wienergegend : 123

orobena renatalis oberthür , 1887 ; bull . soc . ent . fr . ( 6 ) 7 : 99 ; tl : bou saada , . . .

funalis ( grote , 1878 ) ; bull . u . s . geol . surv . 4 : 670

obliqualis ( grote , 1883 ) ; trans . kansas acad . sci . 8 : 56

pachyzancloides sexmaculosus matsumura , 1925 ; j . coll . agric . hokkaido imp . univ . 15 : 190 , pl . 11 , f . 27 ♂ ; tl : sakhalin , ichinosawa

this information is not automatically synchronized with globiz and can sometimes be lagging behind .

[ maps ] warning ! the maps are automatically generated from the textual information , and the process does not always produce acceptable result ; see about maps for more info .

histoire naturelle des insectes . species général des lépidoptéres . tome huitiéme . deltoides et pyralites

a revision of the moths of the subfamily pyraustinae and family pyralidae . part 2

walker , 1859 list of the specimens of lepidopterous insects in the collection of the british museum . supplement list spec . lepid . insects colln br . mus . 16 : 1 - 253 ( [ 1859 ] ) , 17 : 255 - 508 ( 1859 ) , 18 : 509 - 798 ( 1859 ) , 19 : 799 - 1036 ( 1859 )

if you have corrections , comments or information to add into these pages , just send mail to markku savela keep in mind that the taxonomic information is copied from various sources , and may include many inaccuracies . expert help is welcome .

2011 - 01 - 26 by & van nieukerken , dr erik j . karsholt , dr ole & by speidel , w . & segerer , a . nuss , m .

this work is licensed under a creative commons attribution - share alike 3 . 0 license

pesi is funded by the european union 7th framework programme within the research infrastructures programme . contract no . ri - 223806 . activity area : capacities . period 2008 - 2011 - website hosted & developed by vliz banner picture : gannet ( morus bassanus ( linnaeus , 1758 ) ) by karl van ginderdeuren - contact pesi

a number of species are pests , including the cross - striped cabbageworm ( e . rimosalis ) , a pest of cole crops such as cabbage .

sparks , a . and d . g . riley . cross - striped cabbageworm . college of agricultural and environmental sciences . the university of georgia . 2008 .

, 2011 : new crambidae from the afrotropical region ( lepidoptera : pyraloidea : crambidae ) .

, 1998 : the scopariinae and heliothelinae stat . rev . ( lepidoptera : pyraloidea : crambidae ) of the oriental region - a revisional synopsis with descriptions of new species from the philippines and sumatra .

( staudinger , 1892 ) comb . rev . from central asia ( pyraloidea : crambidae : evergestinae ) .

this article is issued from wikipedia - version of the 6 / 29 / 2016 . the text is available under the creative commons attribution / share alike but additional terms may apply for the media files .

biokids is sponsored in part by the interagency education research initiative . it is a partnership of the university of michigan school of education , university of michigan museum of zoology , and the detroit public schools . this material is based upon work supported by the national science foundation under grant drl - 0628151 . copyright © 2002 - 2018 , the regents of the university of michigan . all rights reserved .


stauromedusae are the stalked jellyfishes .

they are the sole living members of the class staurozoa , and belong to the medusozoa subphylum of cnidaria .

they are unique among medusa jellyfish in that they do not have an alternation of polyp and medusa life cycle phases but are instead interpreted as an attached medusa stage , with a life style more resembling that of polypoid forms .

they have a generally trumpet-shaped body , oriented upside-down in comparison with other jellyfish , with the tentacles projecting upwards , and the stalk located in the centre of the umbrella .

members of this class are commonly found in relatively cold waters , close to the shoreline .

sexually mature stauromedusae free-spawn eggs or sperm , which fertilize in the sea and form a creeping , unciliated planula larva .

the larvae crawl across the sea floor and find a suitable place , attaching themselves typically to rock or algae , where they eventually develop into a new , attached stauromedusa .

unlike most scyphozoan jellyfish that practice strobilation , or the process of dividing themselves into body segments , which become new individuals , nearly all stauromedusae develop directly into the adult form .

although conventionally considered to be an order in the class scyphozoa , recent genetic studies suggest that stauromedusae should be elevated to a taxon equivalent of scyphozoa and cubozoa , and should therefore be known as the class staurozoa .

stauromedusae

stauromedusae <EOT> stauromedusae uk - an online guide to the stalked jellyfish ( stauromedusae ) . . .

no one has contributed data records for stauromedusae yet . learn how to contribute .

there are a few other nice photographs of stauromedusae available on the web . click here and here to see two nice photographs of stauromedusae representing quite different morphologies than the two photographs on this page .

suborder cleistocarpida accepted as stauromedusae ( group is not monophyletic . see miranda et al . 2016 . )

suborder eleutherocarpida accepted as stauromedusae ( group is not monophyletic . see miranda et al . 2016 . )

what made you want to look up stauromedusae ? please tell us where you read or heard it ( including the quote , if possible ) .

since stauromedusae do not swim , they have rather limited dispersal potential . they can move around , though , often gradually moving their base along the substrate ( like some sea anemones ) ; some species release at the base and use their adhesive tentacles to " somersault " along . rarely , stauromedusae have even been seen drifting free in the water , after having released at the base . the larvae of stauromedusae are simple elongate planulae that lack the cilia of other cnidarian classes . these tiny larvae move by creeping along , thus having much less dispersive ability than the typical planktonic planulae that swim using their cilia . the larvae of most , or perhaps all , stauromedusae , encyst for months at a time , before re - emerging as recognizable tiny stauromedusae .

a new deepwater species of stauromedusae , lucernaria janetae ( cnidaria , staurozoa , lucernariidae ) , and a preliminary investigation of stauromedusan . . . - pubmed - ncbi

a new deepwater species of stauromedusae , lucernaria janetae ( cnidaria , staurozoa , lucernariidae ) , and a preliminary investigation of stauromedusan phylogeny based on nuclear and mitochondrial rdna data .

i have included links to the few images of stauromedusae located elsewhere on the web to which i was comfortable making a positive identification . any additions to these links are welcome .

i have compiled a list of all valid scientific names for stauromedusae . there are about 50 known species worldwide and most are less than 5 cm long . the largest live in far northern latitudes or in the deep - sea and can be more than 15 cm ( 6 " ) tall . the smallest are only a few mm long as adults . stauromedusae typically live in cold water , although they are occasionally found on warm , tropical or subtropical shores . few scientists have studied stauromedusae , so not so much is known about them , although they are not uncommon along many undisturbed rocky shorelines in europe and northern north america and asia . some have also been found in temperate regions in the southern hemisphere . most stauromedusae are colored to blend in well with their surroundings and are seen only by the most careful observers . most are found in the low intertidal to shallow subtidal shoreline area . some of the best - known north - temperate species have become much rarer in recent decades .

mills , c . e . internet 1999 - present . stauromedusae : list of all valid species names . electronic internet document available at urltoken published by the author , web page established october 1999 , last updated ( see date at end of page ) .

in december 2010 , along with several colleagues particularly interested in stauromedusae , i began to write pages for the encyclopedia of life for every species of stauromedusae or staurozoa . the pages will include images , maps , a description , and summary of all that is known for each species . by early 2011 , those pages should start to appear within the encyclopedia of life and until then , some can be found by a general online search under genus and species names . this project could easily take a couple of years to complete .

one of the most interesting recent observations of stauromedusae was made by richard lutz and colleagues , who visit deep - sea hydrothermal vents by submersible and have discovered that a new species of stauromedusa is the dominant organism in at least two particular areas ( see deep - sea research part ii , volume 45 , pages 329 - 334 , 1998 ) . at the first location , a 2605 m deep vent along the east pacific rise , many stauromedusae ( of a single species ) were located within a fissure of an active vent site . the numerous stauromedusae ( lucernaria janetae collins and daly , 2005 ) at the 2750 m " sarah ' s spring " vent site , also on the east pacific rise , are pictured here . it is very exciting to find this typically shallow , shoreline kind of organism as a main player in such a different ecosystem .

order stauromedusae sessile jellyfish that are vase - , goblet - , or trumpet - shaped , and usually bear 8 groups of tentacles . no more than 2–3 cm long . apparently lacking polypoid stage . temperate and cold temperate waters worldwide . the classification of living cnidarians is relatively stable and generally accepted . …

there are about 50 known species of stauromedusae , which range from the intertidal zone to deep hydrothermal vent habitats , and from antarctica and north polar regions to ( much less frequently ) the tropics . where relevant , i have listed synonyms to good species names in parentheses on the line ( s ) following .

the deepwater stauromedusan lucernaria janetae n . sp is described from adult and juvenile specimens collected from the east pacific rise . lucernaria janetae is the first species in the genus recorded from the pacific ocean , and differs from its congeners in size and morphology . mitochondrial ( 16s ) and nuclear ( ssu ) ribosomal gene sequences from l . janetae were analyzed with those of representative stauromedusan taxa to evaluate stauromedusan monophyly . both genes recovered a strongly monophyletic stauromedusae that is the sister group to all other medusozoans . support of these hypotheses is robust to method of phylogenetic reconstruction and to outgroup selection , buttressing the argument that stauromedusae should be recognized as the class staurozoa . the molecular markers used here favor the same topology of relationships among our samples and clearly distinguished between two species , haliclystus sanjuanensis and h . octoradiatus , that have been considered synonymous by many workers . a stable systematic framework for stauromedusae appears achievable through comprehensive study of both morphological and sequence data .

the fourth order , stauromedusae , comprises some 30 described species of nonswimming , stalked jellies . these species occur chiefly in cooler waters . they are goblet - shaped and fixed by a basal stalk ; the mouth is situated at the upper end . ranging from 1 to 10 cm ( 0 . 4 to 4 inches ) in…

stauromedusae are little stalked jellyfishes that spend their entire life attached to the substrate ( rock or algae , usually ) , rather than swimming freely up in the water column like most other jellyfish . they have long been considered an an order ( stauromedusae ) in the class scyphozoa of the phylum cnidaria , but recent morphological and molecular studies ( marques and collins , 2004 ; collins and daly , 2005 ) argued convincingly that they should be elevated to a rank equal to both the scyphozoa and cubozoa , as the staurozoa . for those who prefer to apply taxonomic ranks , these might now all be considered classes , but many scientists are pulling away from the concept of tight adherence to the old hierarchies of rank , in which case just " staurozoa " will do .

having put together a list of what i believe to be all valid scientific names for the ctenophora , i decided to compile a similar list of names for the stauromedusae ( phylum cnidaria : class staurozoa ) that seem to be valid at the present time ; links are included to stauromedusa images located elsewhere on the web . whereas i believe that the list of genera and species is fairly complete , placement of these genera into families and subfamilies seems still to be fairly plastic , with need for further revision indicated . the basic structure of this taxonomy comes from oskar carlgren ( 1935 ) and tohru uchida ( 1929 ) . few living authors ( including myself ) have seen many species of stauromedusae , which has been one problem in interpreting relationships . presenting here an internet webpage rather than a scientific paper , i have tried to be consistent in interpreting previous authors ' work at the family and subfamily levels . dr . yayoi hirano of japan , perhaps the best - informed stauromedusan biologist in the world at this point , has offered some comments and corrections to this list . other scientists have recently taken a new interest in this small group of unusual and beautiful marine animals .

copyright © 1999 - 2018 john wiley & sons , inc . all rights reserved

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haootia quadriformis n . gen . , n . sp . , interpreted as a muscular cnidarian impression from the late ediacaran period ( approx . 560 ma ) . - pubmed - ncbi

warning : the ncbi web site requires javascript to function . more . . .

liu ag 1 , matthews jj 2 , menon lr 2 , mcilroy d 3 , brasier md 4 .

department of earth sciences , university of cambridge , downing street , cambridge cb2 3eq , uk agscl2 @ cam . ac . uk .

department of earth sciences , university of oxford , south parks road , oxford ox1 3an , uk .

department of earth sciences , university of oxford , south parks road , oxford ox1 3an , uk department of earth sciences , memorial university of newfoundland , 300 prince philip drive , st john ' s , newfoundland and labrador , canada a1b 3x5 .

muscle tissue is a fundamentally eumetazoan attribute . the oldest evidence for fossilized muscular tissue before the early cambrian has hitherto remained moot , being reliant upon indirect evidence in the form of late ediacaran ichnofossils . we here report a candidate muscle - bearing organism , haootia quadriformis n . gen . , n . sp . , from approximately 560 ma strata in newfoundland , canada . this taxon exhibits sediment moulds of twisted , superimposed fibrous bundles arranged quadrilaterally , extending into four prominent bifurcating corner branches . haootia is distinct from all previously published contemporaneous ediacaran macrofossils in its symmetrically fibrous , rather than frondose , architecture . its bundled fibres , morphology , and taphonomy compare well with the muscle fibres of fossil and extant cnidaria , particularly the benthic staurozoa . haootia quadriformis thus potentially provides the earliest body fossil evidence for both metazoan musculature , and for eumetazoa , in the geological record .

is haootia quadriformis related to extant staurozoa ( cnidaria ) ? evidence from the muscular system reconsidered .

haootia quadriformis n . gen . , n . sp . , lower fermeuse formation of back cove , bonavista peninsula , newfoundland . ( a ) haootia quadriformis holotype specimen . note the negative - relief central disc , interpreted as a holdfast , and the broadly bilaterally symmetrical bundles of linear ridges , extending into discrete bifurcating branches . inferred current direction indicated by the arrow . ( b ) fibres running along the right - hand margin of haootia ; each fibre is composed of finer , thinner fibres . ( c ) bottom left corner of haootia , detailing the connection between a primary bifurcating branch and the main body . note the twisted fibres along the branch . ( d ) pinching , bundling and superposition of fibres at the base of a subsidiary branch . ( e ) the small circular depression at the centre of the disc , showing mantling parallel fibres forming the base of a short stalk that connects the disc to the body . ( f ) incomplete paratype specimen of h . quadriformis , from the trepassey formation of burnt point , bonavista peninsula . this specimen is preserved on its side , but clearly displays fibres extending up its stem and around the body . a small partially buried holdfast disc is arrowed . scales bars ( a , c , f ) , 10 mm ; ( b , d , e ) , 5 mm .

digitized images of h . quadriformis n . gen . , n . sp . , emphasizing the convergence of fibrous linear features at the corners of the body , and the symmetry of the fossil . ( a ) photograph of the holotype as it appears in situ . ( b ) interpretive sketch of the non - retrodeformed specimen . labels indicate : ( a ) muscle bundles , ( b ) expanded bundles , ( c ) ‘contracted’ bundles , ( d ) twisting fibres , ( e ) superimposed fibres and ( f ) disc . ( c ) digitized overlay of the fossil . symmetrical regions of the organism are colour coded . note the thick bulging of fibres ( indicating muscle contraction ? ) along short axes of the sheet ( light green ) . ( d ) as in ( b ) , but the image has been corrected to account for tectonic deformation on the surface by compressing the disc into a perfectly circular structure ( cf . [ ] , though see [ ] ) . scale bar , 10 mm .

( a ) the extant staurozoan lucernaria quadricornis , exhibiting a body plan similar to that hypothesized for h . quadriformis n . gen . , n . sp . the staurozoa are known from a range of marine depositional environments and water depths [ ] . ( b ) artistic reconstruction of h . quadriformis . scale bars , 10 mm .

the photograph on the left is manania handi , a little - known stauromedusa that lives attached to eelgrass on the shores of some islands in northwestern washington state and southwestern british columbia . it is colored very much like the eelgrass from which it is hanging ( see adjacent photo for eelgrass color ) , making it difficult to see in the field .

the photograph on the right is haliclystus " sanjuanensis " , which occasionally lives in the same eelgrass beds as manania handi , above . this is one representative of a species complex whose members are found in both the north atlantic and pacific oceans in temporal and boreal waters . i have recently been studying the natural history of both of the above species .

this site is maintained by c . e . mills and all photographs are copyrighted by the author unless otherwise stated . * * this page was established june 1999 ; last updated 27 january 2011 * *

claudia e . mills ( email : cemills " at " u . washington . edu )

note that genus and species names are always written in italics . if the author and date following a species name are within parentheses , that is the taxonomists ' code for indicating that the species has been moved from the genus in which it was originally described ; author and date not in parentheses indicates that the genus has not changed from the original description . ( i am not making this format up - it is strictly formalized and governed by an international body of taxonomists . )

if you use this list in your studies , i would appreciate it if you cite it as suggested above as an electronic publication . the list is an ongoing work and is subject to modification at any time . i consider it my intellectual property and do not want to find it dumped wholesale onto someone else ' s web page - links to it are fine . i welcome any additions or corrections that anyone may wish to make - please contact me at the above address . in order to be true to its in - progress nature , i have left my personal shorthand in the electronic manuscript , so any entry preceeded by . means that i ( or yayoi hirano ) have not yet been able to check the original citation for accuracy . although i have chosen to put this list of species names and authors on the web , it will be noted that i have not included the bibliography from which it derives .

kuekenthali ( antipa , 1891 ) - synonymized with l . walteri by thiel , 1928 and kramp , 1961

( = . calicinaria milne edwards , 1860 ) - by james - clark , 1863 p . 547 from sars , 1860

( = . halimocyathus lagena ( o . f . müller , 1776 ) by larson and fautin , 1989

( = . lucernaria fabricii l . agassiz , 1862 ) by mayer , 1910

this site is maintained by c . e . mills and all photographs are copyrighted by the author unless otherwise stated . * * this page was established october 1999 ; last updated 29 march 2012 * *

cleistocarpida ( group is not monophyletic . see miranda et al . 2016 . )

eleutherocarpida ( group is not monophyletic . see miranda et al . 2016 . )

cornelius , p . f . s . ( 2001 ) . cubozoa , in : costello , m . j . et al . ( ed . ) ( 2001 ) . european register of marine species : a check - list of the marine species in europe and a bibliography of guides to their identification . collection patrimoines naturels , 50 : pp . 111 ( look up in imis ) [ details ]

( of eleutherocarpidae ) hayward , p . j . ; ryland , j . s . ( ed . ) . ( 1990 ) . the marine fauna of the british isles and north - west europe : 1 . introduction and protozoans to arthropods . clarendon press : oxford , uk . isbn 0 - 19 - 857356 - 1 . 627 pp . ( look up in imis ) [ details ]

( of cleistocarpidae ) hayward , p . j . ; ryland , j . s . ( ed . ) . ( 1990 ) . the marine fauna of the british isles and north - west europe : 1 . introduction and protozoans to arthropods . clarendon press : oxford , uk . isbn 0 - 19 - 857356 - 1 . 627 pp . ( look up in imis ) [ details ]

( of cleistocarpida ) daly , m . , m . r . brugler , p . cartwright , a . g . collins , m . n . dawson , d . g . fautin , s . c . france , c . s . mcfadden , d . m . opresko , e . rodriquez , s . l . romano , j . l . stake . ( 2007 ) . the phylum cnidaria : a review of phylogenetic patterns and diversity 300 years after linnaeus . zootaxa . ( 1668 ) : 127 - 182 . , available online at urltoken [ details ] available for editors [ request ]

( of cleistocarpida ) miranda , l . s . ; hirano , y . m . ; mills , c . e . ; falconer , a . ; fenwick , d . ; marques , a . c . ; collins , a . g . ( 2016 ) . systematics of stalked jellyfishes ( cnidaria : staurozoa ) . peerj . 4 : e1951 . , available online at urltoken [ details ]

( of eleutherocarpida ) daly , m . , m . r . brugler , p . cartwright , a . g . collins , m . n . dawson , d . g . fautin , s . c . france , c . s . mcfadden , d . m . opresko , e . rodriquez , s . l . romano , j . l . stake . ( 2007 ) . the phylum cnidaria : a review of phylogenetic patterns and diversity 300 years after linnaeus . zootaxa . ( 1668 ) : 127 - 182 . , available online at urltoken [ details ] available for editors [ request ]

( of eleutherocarpida ) miranda , l . s . ; hirano , y . m . ; mills , c . e . ; falconer , a . ; fenwick , d . ; marques , a . c . ; collins , a . g . ( 2016 ) . systematics of stalked jellyfishes ( cnidaria : staurozoa ) . peerj . 4 : e1951 . , available online at urltoken [ details ]

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animal , ( kingdom animalia ) , any of a group of multicellular eukaryotic organisms ( i . e . , as distinct from…

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horse , ( equus caballus ) , a hoofed , herbivorous mammal of the family equidae . it comprises a single species , …

nmfs , national systematics laboratory , national museum of natural history , mrc - 153 , smithsonian institution , p . o . box 37012 , washington , dc 20013 - 7012 , usa . collinsa @ urltoken

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classification from species 2000 & itis catalogue of life : april 2013 selected by dana campbell - see more .

kento furui added the japanese common name " ジュウモンジクラゲ科 " to " kishinouyeidae " .

michelle marshall added the english common name " stalked jellyfish " to " lucernaria quadricornis müller , 1776 " .

david fenwick added a link to " lucernaria quadricornis " on " lucernaria quadricornis müller , 1776 " .

david fenwick added a link to " haliclystus salpinx " on " haliclystus salpinx clark , 1863 " .

david fenwick added a link to " lucernariopsis campanulata " on " lucernariopsis campanulata ( lamouroux , 1815 ) " .

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cairns , stephen d . , dale r . calder , anita brinckmann - voss , clovis b . castro , daphne g . fautin , . . .

full author list : cairns , stephen d . , dale r . calder , anita brinckmann - voss , clovis b . castro , daphne g . fautin , philip r . pugh , claudia e . mills , walter c . jaap , mary n . arai , stephen h . d . haddock , and dennis m . opresko

disclaimer : itis taxonomy is based on the latest scientific consensus available , and is provided as a general reference source for interested parties . however , it is not a legal authority for statutory or regulatory purposes . while every effort has been made to provide the most reliable and up - to - date information available , ultimate legal requirements with respect to species are contained in provisions of treaties to which the united states is a party , wildlife statutes , regulations , and any applicable notices that have been published in the federal register . for further information on u . s . legal requirements with respect to protected taxa , please contact the u . s . fish and wildlife service .