cadlinidae is a family of sea slugs , dorid nudibranchs , marine gastropod mollusks in the superfamily doridoidea .

this family is within the clade euctenidiacea .

molecular phylogenetic studies and their taxon-sampling schemes can have a strong influence of the resulting phylogeny .

research by r.f. johnson in 2011 has shown that cadlina does not belong to the family chromodorididae .

she has therefore brought back the name cadlinidae from synonymy with chromodorididae .

she also included the genus aldisa bergh , 1878 in the family cadlinidae .

cadlinidae

cadlinidae <EOT> which taxonomic groups does the family cadlinidae belong to and what are the different cadlinidae genus ? below , you will find the taxonomic groups the family cadlinidae belongs to and the taxonomic tree with all the different genus .

which are the most common photographed cadlinidae genus ? below , you will find the list of genus commonly photographed by underwater photographers .

clade euthyneura clade nudipleura clade nudibranchia clade euctenidiacea clade doridacea | superfamilia = doridoidea | familia = cadlinidae | familia _ authority = bergh , 1891 | diversity _ ref = | diversity _ link = diversity of gastropods | diversity = 2 genera | type _ genus = cadlina bergh , 1878 | synonyms _ ref = | synonyms = cadlinidae is a family of . . .

aldisa bergh , 1878 ( taxonomy source ) aldisidae odhner , 1939 accepted as cadlinidae bergh , 1891 ( source of synonymy ) cadlina luarna ( er . marcus & ev . marcus , 1967 ) ( basis of record ) cadlina rumia er . marcus , 1955 ( additional source ) cadlinidae bergh , 1891 ( status source ) chromodorididae bergh , 1891 ( taxonomy source ) chromodoris magnifica ( quoy & gaimard , 1832 ) ( additional source ) doridoidea rafinesque , 1815 ( taxonomy source ) inuda er . marcus & ev . marcus , 1967 accepted as cadlina bergh , 1879 ( source of synonymy ) inuda luarna er . marcus & ev . marcus , 1967 accepted as cadlina luarna ( er . marcus & ev . marcus , 1967 ) ( source of synonymy ) inudinae er . marcus & ev . marcus , 1967 accepted as cadlinidae bergh , 1891 ( source of synonymy ) miamira bergh , 1874 ( source of synonymy ) orodoris bergh , 1875 accepted as miamira bergh , 1874 ( basis of record ) orodoris miamirana bergh , 1875 accepted as miamira miamirana ( bergh , 1875 ) ( source of synonymy )

a . ‘phylogenetic scenario’ for the chromodorid genera modified from [ 5 ] , [ 26 ] . b . morphological phylogeny of generic representatives for the chromodorididae [ 13 ] . c . combined 16s and coi phylogram of the chromodorididae from [ 27 ] . d . combined 16s and coi phylogram of the chromodorididae and cadlinidae from [ 28 ] . rudman ' s ‘ chromodoris group’ in red , ‘ hypselodoris group’ in blue , cadlinella in yellow , diversidoris ( not included in [ 5 ] , [ 13 ] , [ 26 ] ) , cadlina in grey and other dorids in black .

genus acanthochila mörch , 1868 accepted as cadlina bergh , 1879 ( placed on the official index of rejected names by iczn opinion 812 ( bulletin of zoological nomenclature ( 1967 ) 24 : 91 . ) )

genus echinochila mörch , 1869 accepted as cadlina bergh , 1879 ( placed on the official index of rejected names by iczn opinion 812 . ( bulletin of zoological nomenclature ( 1967 ) 24 : 91 . ) )

bouchet p . & rocroi j . - p . ( 2005 ) . classification and nomenclator of gastropod families . malacologia . 47 ( 1 - 2 ) : 1 - 397 isbn 3 - 925919 - 72 - 4 . page ( s ) : 41 [ details ]

johnson r . f . ( 2011 ) breaking family ties : taxon sampling and molecular phylogeny of chromodorid nudibranchs ( mollusca , gastropoda ) . zoologica scripta 40 ( 2 ) : 137 - 157 . [ details ]

( of inudinae er . marcus & ev . marcus , 1967 ) bouchet p . & rocroi j . - p . ( 2005 ) . classification and nomenclator of gastropod families . malacologia . 47 ( 1 - 2 ) : 1 - 397 isbn 3 - 925919 - 72 - 4 . page ( s ) : 92 [ details ]

( of inudinae er . marcus & ev . marcus , 1967 ) johnson r . f . ( 2011 ) breaking family ties : taxon sampling and molecular phylogeny of chromodorid nudibranchs ( mollusca , gastropoda ) . zoologica scripta 40 ( 2 ) : 137 - 157 . [ details ]

( of echinochilidae odhner , 1968 ) bouchet p . & rocroi j . - p . ( 2005 ) . classification and nomenclator of gastropod families . malacologia . 47 ( 1 - 2 ) : 1 - 397 isbn 3 - 925919 - 72 - 4 . page ( s ) : 261 [ details ]

( of aldisidae odhner , 1939 ) bouchet p . & rocroi j . - p . ( 2005 ) . classification and nomenclator of gastropod families . malacologia . 47 ( 1 - 2 ) : 1 - 397 isbn 3 - 925919 - 72 - 4 . page ( s ) : 22 [ details ]

( of aldisidae odhner , 1939 ) johnson r . f . ( 2011 ) breaking family ties : taxon sampling and molecular phylogeny of chromodorid nudibranchs ( mollusca , gastropoda ) . zoologica scripta 40 ( 2 ) : 137 - 157 . [ details ]

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cadlina bergh , l . s . r . , 1878 type species : cadlina repanda alder , j . & a . hancock , 1842

cadlinella thiele , j . , 1931 type species : cadlinella ornatissima risbec , j . , 1928

chromocadlina odhner , n . h . j . in franc , a . , 1968 type species : chromocadlina panamensis odhner , n . h . j . in franc , a . , 1968

hans - martin braun added the english common name " yellow - margin dorid " to " cadlina luteomarginata mcfarland , 1966 " .

hans - martin braun added the german common name " glatte prachtsternschnecke " to " cadlina laevis ( linnaeus , 1767 ) " .

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genus : aldisa l . s . r . bergh , 1878 ( syn : adisa - db : 22 sp , 13 img )

genus : cadlina l . s . r . bergh , 1878 ( syn : acanthochila , echinochila , ectinochila , juanella , inuda - db : 25 sp , 10 img )

cadlina ( sea - slugs ) ( e . g . figures a - b )

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johnson r . f . ( 2011 ) breaking family ties : taxon sampling and molecular phylogeny of chromodorid nudibranchs ( mollusca , gastropoda ) .

family ceratosomatidae gray , 1857 accepted as chromodorididae bergh , 1891 ( under art . 23 . 9 , declared nomen oblitum by bouchet & rocroi , 2005 )

subfamily doriprismaticinae h . adams & a . adams , 1858 accepted as chromodorididae bergh , 1891 ( under art . 23 . 9 , declared nomen oblitum by bouchet & rocroi , 2005 )

bouchet p . & rocroi j . - p . ( 2005 ) . classification and nomenclator of gastropod families . malacologia . 47 ( 1 - 2 ) : 1 - 397 isbn 3 - 925919 - 72 - 4 . [ details ]

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citation : johnson rf , gosliner tm ( 2012 ) traditional taxonomic groupings mask evolutionary history : a molecular phylogeny and new classification of the chromodorid nudibranchs . plos one 7 ( 4 ) : e33479 . urltoken

editor : jonathan h . badger , j . craig venter institute , united states of america

copyright : © 2012 johnson , gosliner . this is an open - access article distributed under the terms of the creative commons attribution license , which permits unrestricted use , distribution , and reproduction in any medium , provided the original author and source are credited .

funding : this work was supported by the california academy of sciences , the university of california santa cruz - department of ecology and evolutionary biology , the national science foundation deb 9978155 and deb 0329054 to tg , and an encyclopedia of life rubenstein fellowship to rj . the funders had no role in study design , data collection and analysis , decision to publish , or preparation of the manuscript .

new collections ( from this contribution and [ 28 ] in blue . genbank specimens in red . size of circle represents number of specimens collected in each region . specimen details in supplementary table s1 ) .

the goals of this contribution are : ( 1 ) generate a phylogeny that tests the species level relationships of the chromodorid nudibranchs and confirms the monophyly of the chromodorididae , ( 2 ) assess the phylogenetic validity of the chromodorid genera , and ( 3 ) propose a new classification for the chromodorid nudibranchs that reflects their relationships .

in this study and a companion study [ 28 ] , thanks to targeted collecting trips , dedicated collectors and dna extracted from museum collections , we were able to include specimens from throughout the indo - pacific ( ip ) , the eastern pacific ( ep ) and west atlantic ( wa ) ( figure 2 and table s1 ) . we use the term indo - pacific to define the biogeographic region including the tropical and subtropical regions of the indian ocean ( from the red sea to the east coast of south africa ) and both the western and central pacific , but not the tropical eastern pacific [ 40 ] . museum collections are an invaluable resource for biodiversity studies [ 41 ] . we have found existing natural history collections can reduce the need for additional collecting . our study , combined with data from [ 28 ] and genbank , is unique in its wide taxonomic and geographic sampling . because we have included both the type species of every genus and additional species of all 14 of the non - monotypic genera , we can test the monophyly every genus in the family ( table s1 ) .

we directly sequenced 142 specimens representing 106 species . we combined these new data with all available sequences on genbank ( table s1 ) . specimens and data from johnson [ 28 ] , genbank accession numbers beginning with eu , are included with new data for figure 2 , but are not treated as new in the numbers of specimens sequenced for this study . in total , we analyzed data from 244 chromodorid specimens , four actinocyclid species and four additional dorid nudibranch species for a total of 165 species and 252 individual specimens . we used doris kerguelensis as the outgroup based on preliminary analyses [ 28 ] . the chromodorid species include at least one species from all of the genera currently classified in the family chromodorididae . the number of species included in this analysis compared to the number of described species per genus is as follows : ardeadoris ( 2 / 2 ) , cadlinella ( 2 / 3 ) , ceratosoma ( 9 / 13 , two undescribed ) , chromodoris ( 50 / 88 , two undescribed ) , digidentis ( 3 / 4 ) , diversidoris ( 1 / 1 ) , durvilledoris ( 3 / 4 ) , glossodoris ( 17 / 30 , two undescribed ) , hypselodoris ( 30 / 59 , two undescribed ) , mexichromis ( 7 / 12 ) , noumea ( 12 / 22 ) , pectenodoris ( 2 / 2 ) , risbecia ( 3 / 5 ) , thorunna ( 8 / 12 ) , tyrinna ( 2 / 2 ) and verconia ( 1 / 1 ) ( s1 ) . all sequences taken from genbank are listed with gb following the species name . we also included coi sequence from two specimens from the moorea biocode project in our analyses ( urltoken ) . we have examined all of the new specimens included here and they are deposited in natural history museums , as indicated by catalog numbers . we never combined sequences from different individuals into chimeras representing one species ; specimens included in these analyses are treated as individuals .

the majority of the specimens used in this study are part of the california academy of sciences invertebrate zoology ( casiz ) collection . we had the permission of casiz to take tissue samples from specimens for dna analysis . as stated in the casiz collections policy : ‘no specimens will be accessioned without adequate labeling , collection notes , field notes , or other locality information , nor without appropriate legal documentation ( collecting permits , export permits from country of origin , etc . ) when applicable . ’ we also included dna extracted for five specimens currently deposited in the museum national d ' histoire naturelle ( paris museum ) and the western australian museum . these tissues samples were collected during joint field trips under the agreement that the tissue could be sequenced at the california academy of sciences , while the specimens would remain at the respective museum . all other data used is from genbank or the moorea biocode database .

most of our samples were collected especially for molecular work and were preserved accordingly , either in 95 % etoh , sed buffer ( saturated nacl solution with edta and dmso ) or frozen . in addition to the specimens collected specifically for molecular study , we were also able to use museum material that was , either preserved in 70–75 % etoh or the original fixation method is unknown .

the cleaned , pcr products were copied and labeled with fluorescently dye - terminators ( big dye 3 . 1 abi ) in 10 µl reactions . each reaction contained 0 . 5–2 µl of cleaned pcr product , 1 . 63 µl of 5× reaction buffer , . 5 µl of primer ( 10 mm stock ) , 0 . 5 µl–0 . 75 µl of big dye and water to 10 µl . these reactions were run on a perkin elmer 9600 - geneamp pcr system or a biorad mycycler™ thermocycler ( software version 1 . 065 , bio - rad laboratories ) . the resulting labeled , single stranded dna was precipitated by addition of 2 . 5 µl of edta and sequential washing and pelleting in ( centrifuge details ) with 100 % and then 70 % etoh . the pelleted dna was denautured for two minutes at 94°c in 13–15 µl of hidi formamide ( applied biosystems ) . the denatured , labeled dna fragments were sequenced in both directions on the abi 3100 and 3130 genetic analyzer in the center for comparative genomics ( formerly the osher laboratory for molecular systematics ) at the california academy of sciences .

we assembled , edited and removed primer strands from forward and reverse strands for each gene fragment sequenced using sequencher ( ver . 4 . 7 . genecodes corporation ) and genious 3 . 0 - 5 . 3 . 3 ( biomatters ) . we aligned the coi sequences by eye and translated the base pair data into amino acids to using macclade 4 . 08 [ 46 ] to confirm alignment accuracy . we aligned 16s sequences with muscle [ 47 ] . we then further optimized the alignments by eye using both macclade [ 46 ] and genious 3 . 0 - 5 . 3 . 3 ( biomatters ) .

we tested for saturation or multiple substitutions at the same site by plotting the absolute number of transitions and transversions at each codon position ( 1 st , 2 nd , 3 rd ) for coi and at each base pair for 16s against both uncorrected p distance and log det using paup [ 48 ] and excel ( plots not shown ) .

sequence data for both genes was not obtained for every specimen we studied . we worked with two main data sets , because we wanted to test the effect of missing data on the resulting phylogeny : the two data sets were : 1 ) combined 16s and coi for specimens with sequence data for both genes , 2 ) all 16s and coi data for all specimens ( table s1 . ) both of these data sets were analyzed both including and excluding variable characters in the 16s alignment . for all of these analyses we used doris kerguelensis as the outgroup .

we know from the discovery of polyphyletic and paraphyletic generic groupings in chromodoris , glossodoris , hypsleodoris , mexichromis and noumea [ 27 ] , [ 28 ] , that the current classification of the chromodorididae does not reflect the evolutionary history of the group . we cannot continue to use this current classification . we will use the resulting phylogenies to propose a new classification of the chromodorid nudibranchs . the proposed new classification is based on several fundamental tnets of phylogenetic classification . only clades are named , with two exceptions described below . each clade contains the type species of the name - bearing clade . exisiting , available names are utilized wherever possible to minimize the disruption to nomenclature , while simultaneously reflecting relationship . we will identify clades that include the type species of each chromodorid genus and delineate genera to minimize conflict with current classification and support recognition of interesting morphology . the translation of phylogenetic hypotheses into classifications is the best way to communicate results to a larger community , but even as the number of molecular phylogenies increases , the number of new classifications is decreasing [ 54 ] – [ 56 ] . the growing phylogeny / classification gap is troubling . phylogenies are hypotheses of relationship and communicating these new hypotheses is one of the main contributions systematics can make to the scientific community .

the sequenced coi fragment is 658 base pairs ( bp ) long . the edited 16s sequences are 531 bp long . the combined data sets with gaps introduced for alignment are 1189 base pairs long . all sequences are available from genbank coi ( jq727822–jq727914 ) , 16s ( jq727689–jq727821 ) and aligned data matrices are available upon request from the corresponding author . excluded variable 16s regions are identified as character sets in all nexus files . saturation was not found in the 16s fragment or the first or second positions of the coi fragment . there is slight saturation in the third position transitions in the coi data set ( not shown ) . the third positions were included in the bayesian analysis as the partitioning allows the parameters of this position to be estimated separately and the inclusion of the third positions did not change the resulting trees . the recommended model of evolution ( aic form mr . model test ) was used to set parameters in mr . bayes for each partition . the resulting best - fit model of evolution for each partition using the aic selection from mr . modeltest ver . 2 [ 78 ] were coi 1 st : gtr + g , coi 2nd : trn + i + g , coi 3 rd : gtr + i + g and 16s : gtr + i + g . these models correspond to the following settings in mr . bayes ; all partitions set to nst = 6 and rates = invgamma except for the coi second codon position partition which was set rates = gamma .

the figured trees are the resulting consensus phylograms from the bayesian analyses ( figures s1 , s2 ) . all posterior probabilities are shown above the branches on the bayesian phylograms . tree topology was not altered with the inclusion or exclusion or the 16s fragment ' s variable regions ( see figure s2 for comparison of trees with and without variable regions ) . the resulting phylogenetic hypotheses for each dataset are summarized below . we will discuss relationships in terms of posterior probabilities .

coi and 16s combined analysis : including only specimens with data for both genes ( figure s1 ) .

this data set included 164 individual chromodorids , representing 123 species , three species of actinocyclidae , four other dorids . the outgroup was doris kerguelensis . the data set included was 1189 bases long included gaps introduced to aid in alignment of variable regions . all bases are included . in the majority rule consensus phylogram resulting from the bayesian analysis , the chromodorids are monophyletic ( pp = 1 . 00 ) . they are sister ( pp = 0 . 98 ) to the monophyletic actinocyclids ( pp = 1 . 00 ) . cadlinella ornatissima is sister to the rest of the chromodorids ( pp = 1 . 00 ) . the monophyletic tyrinna ( pp = 1 . 00 ) is poorly supported as sister to the main clade of chromodorids ( pp = 0 . 82 ) . the main clade of all chromodorids , except cadlinella and tyrinna is supported ( pp = 0 . 85 ) . two clades of noumea ( both pp = 1 . 00 ) are part of a basal polytomy with the clade including the remaining chromodorid species ( pp = 0 . 89 ) . a well - supported clade ( pp = 1 . 00 ) containing some species of glossodoris is poorly supported at the base of the chromodorid grade . within this main clade of chromodorids , there is one very well supported clade , which includes all species of ceratosoma , durvilledoris , hypselodoris , mexichromis , pectenodoris , risbecia , thorunna and some digidentis ( pp = 1 . 00 ) diversidoris aurantionodulosa and noumea crocea are sister species ( pp = 1 . 00 ) and poorly supported ( pp = 0 . 78 ) as sister to a poorly supported clade ( pp = 0 . 66 ) that includes this well - supported clade and chromodoris alternata and chromodoris ambiguus ( pp = 1 . 00 ) . there is also a poorly supported clade ( or grade if the clade is collapsed ) of smaller clades made up of species of ardeadoris , chromodoris , diversidoris , glossodoris , noumea , verconia and one species of digidentis ( pp = 0 . 67 ) . of the other 12 non - monotypic traditional genera , seven ( ceratosoma , chromodoris , digidentis , glossodoris , hypselodoris , mexichromis , noumea ) are non - monophyletic and three ( durvilledoris , pectenodoris , risbecia ) are monophyletic but render another genus paraphyletic . both ardeadoris and thorunna are made paraphyletic by nested members of other genera ( noumea , glossodoris - within ardeadoris and digidentis - within thorunna ) . there are three species and five clades of eastern pacific and / or atlantic species .

this data set included 244 individual chromodorids , representing 157 species , four species of actinocyclidae , four other dorids . the outgroup was doris kerguelensis .

the complete data set included 1189 bases . the chromodorids are monophyletic ( pp = 0 . 94 ) . they are sister to the monophyletic actinocyclids ( pp = 0 . 98 ) . a clade including both species of cadlinella is sister to the rest of the chromodorids ( pp = 1 . 00 ) . the monophyletic tyrinna ( pp = 1 . 00 ) is poorly supported as sister to the main clade ( pp = 0 . 83 ) . there are two clades containing species of noumea and the one species of verconia ( pp = 1 . 00 and pp = 1 . 00 ) that form a polytomy with the clade of all of the remaining chromodorids ( pp = 0 . 85 ) . within the main clade of chromodorids , there is one very well supported clade , which includes all species of ceratosoma , durvilledoris , hypselodoris , mexichromis , pectenodoris , risbecia , thorunna and some digidentis ( pp = 1 . 00 ) and a grade of clades of species ardeadoris , chromodoris , diversidoris , glossodoris , noumea , verconia and one species of digidentis . of the other 12 non - monotypic traditional genera , seven ( ceratosoma , chromodoris , digidentis , glossodoris , hypselodoris , mexichromis , noumea ) are non - monophyletic and three ( durvilledoris , pectenodoris , risbecia ) are monophyletic but render another genus paraphyletic . both ardeadoris and thorunna are made paraphyletic by nested members of other genera ( noumea , glossodoris and digidentis ) ( figure 3 ) . more detailed results found within each clade will be discussed below . there are three individual species and five clades of eastern pacific or atlantic species ( figure 4 ) . the data set without variable regions included 1108 bases . there are only slight changes to the tree topology , including slight losses of support and changes to branching pattern in the species relationships within four clades containing species of noumea , glossodoris , chromodoris and thorunna . additionally , some clades are more well - supported in the phylogram without variable regions , most notably there is some support for the two noumea clades as sisters . all of these differences are mapped in mirrored trees ( figure s2 ) .

tree is the same bayesian phylogram as figured in s3a . all specimens , both genes and all characters included .

tree is the same bayesian phylogram as figured in s3a . all specimens , both genes and all characters included . blue = indo - pacific , red = atlantic and caribbean , gold = eastern pacific , green = sister group , black = outgroups . dark grey = solely eastern pacific and atlantic clades . light grey = primarily indo - pacific clades with eastern pacific members .

our classification is based on our coi and 16s combined phylogeny with all specimens included ( figure s2a ) . the older name that describes this clade , ceratosomatidae gray 1857 [ 61 ] was declared nomen oblitum under art . 23 . 9 of the international code of zoological nomenclature [ 58 ] . even though it is older than the name in current usage , chromodorididae , it had not been used in over fifty years . in the phylogeny of the chromodorid nudibranchs , there are five basal clades : cadlinella , tyrinna , two clades made up of some species of noumea and verconia and one clade made up of some species of glossodoris . there is one , main , well - supported clade including species of ceratosoma , hypselodoris , and grade of clades . we will briefly introduce each clade and its member species in the context of a new classification for chromodorid nudibranchs ( figure 5 , s2a , s3 ) .

the two species of cadlinella included here , cadlinella ornatissima and cadlinellla subornatissima form a clade and are sister to the rest of the chromodorid species ( pp = 1 . 00 ) . these findings support previous results [ 27 ] , [ 28 ] and rudman ' s evolutionary scenario [ 5 ] , [ 26 ] . the widespread indo - pacific genus , cadlinella is an enigmatic taxon . it has at different times been considered it own separate family [ 64 ] , a part of the cadlininae [ 65 ] and a member of the chromodoridinae / chromodorididae [ 26 ] , [ 66 ] .

cadlina burnayi ortea , 1988 [ 68 ] = t . nobilis [ 69 ]

the only two species of tyrinna : t . evelinae and t . nobilis are included here . tyrinna is always monophyletic ( pp = 1 . 00 ) . after the split from cadlinella , this clade is poorly - supported as the sister group to the main group of chromodorids ( pp = 0 . 83 ) . rudman [ 26 ] suggested that tyrinna , cadlinella and cadlina form a basal grade of primitive chromodorids . cadlina had been shown not to be a chromodorid [ 28 ] , but our results support rudman ' s suggestion that tyrinna and cadlinella are basal to the rest of the chromodorids . muniain et al [ 70 ] and schrödl and millen [ 71 ] extensively reviewed the morphology of the two species in this clade .

verconia verconis is well supported as part of a clade that includes n . haliclona , n . laboutei , n . romeri and n . simplex ( pp = 1 . 00 ) . noumea varians , n . purpurea and n . norba form a well - supported clade ( pp = 1 . 00 ) that is not part of a name bearing clade , but is one branch of the polytomy that includes the ‘noumea sensu stricto’ and the branch leading to the rest of the family ( pp = 0 . 88 ) . the monotypic genus verconia is nested within the noumea clade as suggested by rudman [ 75 ] and weakly supported as the sister species to another south australian species , n . haliclona , as found in the preliminary results shown by turner & wilson [ 27 ] .

type species : doris xantholeuca ehrenberg , 1831 [ 76 ] = g . pallida ( by subsequent designation )

the glossodoris clade ( pp = 1 . 00 ) includes species g . pallida and g . rufomargninata . in an important , but often overlooked detailed examination of the relationships of the species classified in the genus glossodoris , rudman identified five subgroups of this genus based on morphology [ 77 ] . the species in this glossodoris clade were considered by rudman [ 77 ] to be members of the ‘ glossodoris pallida subgroup’ . this clade also includes two species he did not include in any subgroup , g . cincta and g . hikuerenesis .

lissodoris odhner , 1934 [ 80 ] . type species : l . mollis odhner , 1934 ( = c . aureomarginata cheeseman , 1881 [ 86 ] ( by monotypy )

this clade includes all of the indo - pacific species of chromodoris that are not part of the black - lined , planar egg mass clade ( pp = 1 . 00 ) , except chromodoris alternata and chromodoris ambiguus . this phylogeny is the first to find definitive support for a clade of chromodorids , first suggested by wilson [ 16 ] and turner and wilson [ 27 ] known to lay egg masses with extra - capsular yolk . when pease designated doris vibrata as the type species for the new genus goniobranchus , he should have changed the ending of vibrata to vibratus to reflect the masculine gender of the –us ending . we have made that correction here and changed the gender of all of the species names that require changing ( names derived from adjectives ) in goniobranchus .

casella h . & a . adams , 1858 : 57 [ 84 ] . type species : c . gouldii h . & a . adams , 1858 [ 84 ] ( by monotypy )

species included in the glossodoris atromarginata subgroup [ 77 ] are recovered in this clade , with the addition of g . sedna and digidentis kulonba ( pp = 0 . 95 ) .

this name will be used for all eastern pacific and atlantic species of chromodoris and glossodoris ( except glossodoris sedna ) . these species form a polytomy including glossodoris baumanni and three clades of atlantic and eastern pacific chromodorids .

chromodoris clenchi , c . norrisi and c . sphoni ( pp = 1 . 00 )

chromodoris krohni , c . luteorosea and c . purpurea ( pp = 0 . 78 )

these exclusively eastern pacific and atlantic clades do not form a monophyletic group , but we will provisionally name all of these species ‘felimida’ . this is the most conservative choice , the choice that requires the fewest name changes and is the least disruptive pending further information and broader taxon sampling .

the ardeadoris clade includes both species of ardeadoris : a . egretta and a . scottjohnsoni , five species of glossodoris ( g . averni , g . pullata , g . rubroannulata , g . tomsmithi and glossodoris undaurum ) and noumea angustolutea ( pp = 1 . 00 ) . . based on their analysis , turner and wilson [ 27 ] suggested that with more sampling it would be come clear if ardeadoris should be synonmized with glossodoris . by sampling more broadly within the family , we found the converse . four species of glossodoris and noumea angustolutea need to be included within ardeardoris because they are strongly supported as part of the clade including ardeadoris egretta and not the type species of glossodoris . three of the species , g . averni , g . undaurum and g . rubroannulata , found in this clade were part of rudman ' s glossodoris sedna subgroup [ 77 ]

this clade includes all of the black - lined species of chromodoris and chromodoris aspersa ( pp = 1 . 00 ) . this clade was identified by , both wilson & lee [ 17 ] and turner & wilson [ 27 ] , as the planar spawning or black - lined chromodoris clade . all of the members of this clade lay flat egg masses .

type species : diversidoris aurantionodulosa rudman , 1987 [ 89 ] ( by original designation ) .

the diversidoris includes , diversidoris aurantionodulosa , two yellow species of noumea , n . crocea and n . flava , and a new species from moorea , french polynesia - chromodoridae biocode 2937 ( pp = 0 . 95 ) .

the miamirinae clade includes all of the species currently classified as ceratosoma , durvilledoris , hypselodoris , mexichromis , pectenodoris , risbecia , thorunna and two species of digidentis ( pp = 1 . 00 )

this clade was first predicted by rudman [ 26 ] based on morphological similarities and then confirmed by rudman & berquist ' s [ 5 ] finding that all of the species in this clade feed exclusively on sponges of the family dysideaidae , although they assumed all of the genera to be monophyletic . miamirinae bergh 1891 is the oldest appropriate and available subfamily or family name for this clade . the remaining six genera ; miamira , ceratosoma , felimare , mexichromis , thorunna and hypselodoris make up the miamirinae .

the miamira clade includes the following species ( as currently classified ) ceratosoma alleni , ceratosoma magnificum , ceratosoma miamiranum , ceratosoma sinuatum . miamira is part of a grade with ceratosoma . the morphological phylogeny of species of ceratosoma and classified as miamira and orodoris , that was used as justification for their synonomy , predicted a sister group relationship between species of miamira and ceratosoma alleni [ 93 ] . our results confirm that c . alleni is more closely related to species of miamira , but do not find support for synonymy of miamira and ceratosoma . although , it is possible this relationship will be recovered with further sampling and by including molecular markers that will help resolve basal branches on the phylogeny .

the ceratosoma clade includes c . amoenum , c . gracillimum , c . ingozi , c . tenue , c . trilobatum and a new species . ( pp = 1 . 00 )

the felimare clade includes all eastern pacific , atlantic and mediterranean species of hypselodoris and two species of mexichromis , m . porterae and m . kempfi from the eastern pacific and caribbean respectively ( pp = 1 . 00 ) . both gosliner and johnson [ 13 ] and alejandrino and valdés [ 31 ] hypothesized a sister group relationship between the indo - pacific and eastern pacific / atlantic species of hypselodoris . turner and wilson [ 27 ] did not recover that relationship , but instead found the same relationships shown here .

this clade includes the type species of mexichromis , m . antonii , known only from the eastern pacific and the three included species of durvilledoris , d . lemniscata , d . pusilla and d . similaris , the two described species of pectenodoris , p . aurora and p . trilineata and all of the indo - pacific species currently considered mexichromis , m . festiva , m . macropus , m . mariei and m . mutituberculata ( pp = 1 . 00 ) . there are two well - supported clades within the mexichromis clade . the clade including mexichromis antonii and the species of durvilledoris is sister to the clade including pectenodoris and indo - pacific mexichromis . these clades could be given two names , but it is much less disruptive and confusing to maintain the name mexichromis for all clade members . the clade including p . aurora and p . trilineata can be called the ‘ pectenodoris ’ clade of mexichromis .

the thorunna clade includes all species of thorunna and two species of digidentis , d . arbutus and d . perplexa . all of species currently classified as thorunna are found in the indo - pacific and the species of digidentis are limited to southern australia . as suggested by rudman [ 26 ] , the only species within thorunna with mantle glands , t . australis and the species of digidentis ( all of which have mantle glands ) form a clade .

this clade includes all of the indo - pacific species of hypeslodoris and risbecia ( pp = 1 . 00 ) .

species of risbecia s . s forms a well - supported clade nested within hypselodoris and can be referred to as the risbecia clade of hypselodoris . risbecia aplogema is not part of this risbecia clade and was previously considered a species of hypselodoris . including all of the members of the risbecia and hypselodoris bullocki clade in risbecia is not an option because this would render hypsleodoris paraphyletic . the second clade includes , h . bennetti , h , maritima , h . bertschi , h . paulinae , h . kaname , h . bollandi , h . obscura , h . infucata , h . zephrya and one or two new species . the third clade includes h . reidi , h . krakatoa , h . jacksoni and one new species . this clade was also recovered in gosliner & johnson ' s [ 13 ] morphological phylogeny of hypselodoris .

the enigmatic south australian species , chromodoris alternata and c . ambiguus are very different than other chromodorids . they are two of the five chromodorid species with a plesiomorphic serial reproductive system ( c . loringi , c . thompsoni , c . woddwardae ) [ 26 ] , [ 28 ] , [ 89 ] . all five of these species are found only in southeastern australia . these species were found to be more closely related to cadlina than chromodoris by wilson & lee [ 17 ] , but as part of the chromodorid grade in turner & wilson [ 27 ] . clearly further work on this group and its relationship to all cryptobranchs is needed . the addition of specimens of c . loringi , c . thompsoni and c . woodwardae [ 26 ] , [ 89 ] , [ 99 ] , the only other chromodorid species known to have a serial reproductive system may help solve this problem . these two species are always each other ' s closest relatives and are sister to the rest of the miamirainae in the all analyses . as suggested by dayrat & gosliner [ 60 ] they should be considered chromodorididae , because they are not included in a named clade . until the ambiguity of the relationship of these taxa to other chromodorids can be resolved , they should be considered chromodorididae alternata and chromodorididae ambiguous .

in future contribtuions , we will work out synapomorphies for the clades identified here , but because of the amount of homoplasy and number of incomplete descriptions , this is a huge undertaking and not appropriate here .

in summary , with the most comprehensive sampling of chromodorid species to date , we confirmed that the chromodorids are monophyletic and are sister to the monophyletic actinocyclids . we also found that the majority , 12 / 14 non - monotypic traditional genera , were not monophyletic or make another clade paraphyletc . seven traditional genera , ceratosoma , chromodoris , digidentis , glossodoris , hypselodoris , mexichromis , noumea are non - monophyletic and three ( durvilledoris , pectenodoris , risbecia ) are monophyletic but render another genus paraphyletic . both ardeadoris and thorunna are made paraphyletic by nested members of other genera ( noumea , glossodoris and digidentis ) . the two monotypic genera , diversidoris and verconia are nested within clades . only tyrinna and cadlinella are monophyletic and without disruption to any other clades ( figure 3 , s1 , s2 ) . the classification proposed here and discussed at length above renames clades and is more consisitent with evolutionary history ( figure s3 ) .

the most speciose chromodorid genera : chromodoris , glossodoris and hypselodoris were originally created to describe indo - pacific species . it wasn ' t until some time after these names were created that previously described , similar , brightly colored cryptobranch dorid species found in the eastern pacific , western atlantic and mediterranean were added to these genera [ 1 ] , [ 26 ] , [ 65 ] , [ 95 ] , [ 102 ] , [ 103 ] . in mexichromis the opposite is true . the type species , mexichromis antonii , was described from the eastern pacific and indo - pacific species were included later included in this genus [ 26 ] , [ 95 ] . other eastern pacific “ mexichromis ” are shown here to belong to felimare .

this new classification clarifies our view of biogeographic patterns in the chromodorid nudibranchs . instead of taxonomy obscuring patterns of diversification in this group , this taxonomy reflects and reinforces evolutionary history . it gives us a much better framework for exploring evolutionary questions .

the majority of chromodorid nudibranchs are found in the indo - pacific , but there are three individual species and five clades of solely atlantic and / or eastern pacific species ( figure 4 ) . the sister group to the rest of the chromodorids , cadlinella is found only in the indo - pacific , while the sister to the chromodorididae , the actinocyclidae is found in most temperate and tropical waters . although there are other possibly scenarios , such as trans - pacific dispersal and migration around africa , the pattern uncovered here , strongly supports the simplest hypothesis that the chromodorids diversified rapidly from the tropical tethyan realm . this pattern has been found in other gastropod groups [ 104 ] – [ 108 ] ( figure 4 ) . the chromodorids were likely widely distributed and different lineages diversified in isolation following vicariant events . this scenario is further supported by the fact that goniobranchis is sister to ‘ doriprismatica ’ and its closest realitves and that all the memebers of goniobranchus are indo - pacific . also in this scenario , the specimens identified as d . sedna from the atlantic and eastern pacific appear to be distinct species as indicated by coi pairwise distances of 11 . 7–11 . 0 % between eastern pacific and altantic specimens while the three eastern pacific specimens are 0–0 . 7 % different from each other . this scenario clearly supports vicariance between the indo - pacific and eastern pacific and atlantic preceding the vicariance between the eastern pacific and atlantic . in the main chromodorid grade of clades there are two individual species and three clades that are exclusively atlantic and / or eastern pacific . specimens identified as ‘ doriprismatica’ sedna found both in the eastern pacific and the western atlantic , are always sisters and are nested within a clade of exclusively indo - pacific species . this is most likely a radiation into the eastern pacific and atlantic from the indo - pacific . the remainder of the atlantic and eastern pacific species , not included in the miamirinae , are part of a polytomy including five clades , three containing only eastern pacific and atlantic species of ‘felimida’ and the indo - pacific ardeadoris and chromodoris clades . ‘felimida’ baumanni , found in the eastern pacific , is also part of this polytomy . the relationships in this grade need to be examined more closely with the addition of more specimens and more genes .

relationships within the miramirinae clade are more resolved . there are two clades that include eastern pacific and atlantic species . the felimare clade is exclusively eastern pacific and atlantic . there are two eastern pacific and atlantic splits in this clade , the eastern pacific f . porterae and caribbean f . kempfi are potentially geminate species and are sister to a larger clade of eastern pacific , caribbean and eastern atlantic felimare species . more sampling is needed in this clade to further untangle the emergent biogeographic patterns within felimare . additionally , within the miamirinae , the eastern pacific species , mexichromis antonii , is sister to the exclusively indo - pacific m . lemniscata , m . pusilla and m . similaris and this clade is sister to the rest of the indo - pacific mexichromis . within the miamirinae , it appears that there had been more than one dispersal event from the indo - pacific , to eastern pacific and atlantic . ceratosoma and miamira species are only found in the indo - pacific and adjacent temperate regions . the sister taxon , the clade including : felimare , mexichromis , thorunna and hypselodoris , has a wider distribution . within this sister taxon , felilmare is exclusively eastern pacific and atlantic while its sister species in its sister taxon mexichromis , thorunna and hypselodoris are almost exclusively indo - pacific and adjacent temperate regions . mexichromis antonii , which is found in the eastern pacific , is the only speices in this clade not found in the indo - pacific or adjacent regions . thus , felimare and mexichromis antonii represent two distrinct invasions of the eastern pacific from the indo - pacific .

we hope the work presented here will serve as a starting point for further research into the evolutionary history of the chromodorid nudibranchs . this phylogeny is based only on mitochondrial genes , one of our first next steps will be to include sequences from nuclear genes for all of the species included here . the addition of more slowly evolving unlinked markers should help resolve some poorly supported node at the base of the phylogenies presented here and will add a separate line of evidence to this hypothesis . addtionally , morphological synapomorphies need to be found for the clades recoverd in this phyogeny . there is still much work to do in order to untangle the evolutionary history of this group . this phylogeny and classification is a start that will allow us to use names that represent monophyetic groups as the starting point for future discovery .

the electronic version of this document does not represent a published work according to the international code of zoological nomenclature ( iczn ) , and hence the nomenclatural acts contained in the electronic version are not available under that code from the electronic edition . therefore , a separate edition of this document was produced by a method that assures numerous identical and durable copies , and those copies were simultaneously obtainable ( from the publication date noted on the first page of this article ) for the purpose of providing a public and permanent scientific record , in accordance with article 8 . 1 of the code . the separate print - only edition is available on request from plos by sending a request to plos one , public library of science , 1160 battery street , suite 100 , san francisco , ca 94111 , usa along with a check for $ 10 ( to cover printing and postage ) payable to “public library of science” .

table of specimens used in this study . specimens used in this study listed by family . the names in this table reflect current classification not proposed classification ( new names are listed in the text ) . abbreviations are as follows : casiz = california academy of sciences , sam = south australian museum , wam = western australian museum , am = australian museum , zsm = zoologische staatssammlung münchen , sio - bic = scripps institute of oceanography , biocode = moorea biocode project .

new classification of the chromodorididae with synonyms . generic names and type species in bold and the most recent genus membership follows . listing order follows phylogeny .

bayesian consensus phylogram including all specimens with data for both genes . posterior probabilities are listed above branches . doris kerguelensis is the outgroup . this phylogram is the consensus of 50 , 000 , 000 generations with trees sampled every 1000 generations with a burnin of 25 % . data was partitioned by gene and by codon position .

bayesian consensus phylograms including all specimens . a . phylogram resulting from the inclusion of all characters b . phylogram resulting from excluding hard to align characters . doris kerguelensis is the outgroup . these phylograms are the consensuses of 50 , 000 , 000 generations with trees sampled every 1000 generations with a burnin of 25 % . data was partitioned by gene and by codon position . dotted lined indicate areas of disagreement .

ángel valdés , don potts , nudibranch central at cas , especially marta pola , yolanda camacho , jamie chan and benoit dayrat , the potts lab ucsc , the center for comparative genomics at cas , especially josh hallas , boni cruz , susie lockhart , monica medina , kristy deiner , kristen cella , alessandra lopez , anna sellas and brian simison . a work of this scope is a true collaboration and would have been impossible with the dedication and knowledge of : cory pittman , pauline fiene , robert bolland , scott johnson , inbio and yolanda camacho , lucas cervera , larry harris , alicia hermosillo , yvonne valles , benoit dayrat , mike miller , hans bertsch , marta pola , goncalo caladao , shireen fahey , patrick krug , manuel malquias , correy whisson of wam , lindsay groves of lacm , moorea biocode project , philippe bouchet and census of marine life - pangolo and santo expeditions .

conceived and designed the experiments : rfj tmg . performed the experiments : rfj . analyzed the data : rfj . contributed reagents / materials / analysis tools : tmg . wrote the paper : rfj . conceptualized and wrote the new classification : tmg rfj .

edmunds m ( 1981 ) opishtobranchiate mollusca from ghana : chromodorididae . zoological journal of the linnean society 71 : 175–201 .

species ( gastropoda : nudibranchia ) ultrastructure and chemical analysis of mantle dermal formations ( mdfs ) . marine biology 106 : 245–250 .

( opisthobranchia , chromodorididae ) . journal of molluscan studies 72 ( 2 ) : 214–216 .

rudman wb , berquist pr ( 2007 ) a review of feeding specificity in the sponge - feeding chromodorididae ( nudibranchia : mollusca ) . molluscan research 27 ( 2 ) : 60–88 .


perithemis is a genus of dragonflies commonly known as amberwings .

they are characterized by their small size and the amber wings of the male .

perithemis includes the following species : perithemis bella kirby , 1889 perithemis capixaba costa , de souza & muzón , 2006 perithemis cornelia ris , 1910 - orange amberwing perithemis domitia ( drury , 1773 ) - slough amberwing perithemis electra ris , 1930 - golden amberwing perithemis icteroptera ( selys in sagra , 1857 ) perithemis intensa kirby , 1889 - mexican amberwing perithemis lais ( perty , 1834 ) - fine-banded amberwing perithemis mooma kirby , 1889 perithemis parzefalli hoffmann , 1991 - clear-tipped amberwing perithemis rubita dunkle , 1982 - ruby amberwing perithemis tenera ( say , 1840 ) - eastern amberwing perithemis thais kirby , 1889

perithemis

perithemis <EOT> eastern amberwing perithemis tenera female balmorhea lake , reeves co . texas 21 august 2013

eastern amberwing perithemis tenera female dripping springs , hays co . , texas 30 june 2017

eastern amberwing perithemis tenera male southeast metro park , travis co . , texas 16 may 2014

eastern amberwing perithemis tenera male barkeley meadows park , travis co . , texas 8 september 2015

eastern amberwing perithemis tenera male southeast metro park , travis co . , texas 9 july 2016

eastern amberwing perithemis tenera male hornsby bend , austin , travis co . , texas 30 may 2013

eastern amberwing perithemis tenera female hornsby bend , austin , travis co . , texas 7 june 2013

eastern amberwing perithemis tenera female hornsby bend , austin , travis co . , texas 11 may 2014

eastern amberwing perithemis tenera male hornsby bend , austin , travis co . , texas 2 august 2014

eastern amberwing perithemis tenera male hornsby bend , austin , travis co . , texas 14 august 2014

eastern amberwing perithemis tenera female hornsby bend , austin , travis co . , texas 27 april 2015

eastern amberwing perithemis tenera female hornsby bend , austin , travis co . , texas 9 june 2015

eastern amberwing perithemis tenera male hornsby bend , austin , travis co . , texas 11 may 2016

eastern amberwing perithemis tenera male hornsby bend , austin , travis co . , texas 26 april 2017

eastern amberwing perithemis tenera male chub sandhill natural area preserve , sussex co . , virginia 7 june 2017

eastern amberwing perithemis tenera male roger hanks park , dripping springs , hays co . , texas 17 june 2015

eastern amberwing perithemis tenera male city of rocks state park , grant co . , new mexico 27 august 2015

eastern amberwing perithemis tenera male harrison lake national fish hatchery , charles city co . , virginia 7 june 2017

eastern amberwing perithemis tenera female pedernales river nature park , johnson city , blanco co . , texas 13 june 2013

eastern amberwing perithemis tenera female pedernales river nature park , johnson city , blanco co . , texas 15 august 2013

eastern amberwing perithemis tenera female lakes regional park , ft . myers , lee co . , florida 8 october 2014

eastern amberwing perithemis tenera teneral male , just emerged pocahontas state park , chesterfield co . , virginia 8 june 2017

eastern amberwing perithemis tenera male san marcos river , ca 3 mi w of luling , caldwell co . , texas 21 july 2017

eastern amberwing perithemis tenera female yett creek park , austin , travis co . , texas 16 july 2015 lat 30 . 429596 , lon - 97 . 737002

the eastern amberwing ( perithemis tenera ) is a small , thick - bodied dragon that utilizes many habitats from streams to lakes to ponds and marshes . it is one of the smallest dragons in north america and it is sometimes called a " wasp mimic " since it can look very wasp - like in flight . the first image shown here was photographed in austin , travis co . , texas in may , 2003 . the image was a digital capture with a canon eos 10d and ef 300 mm f4 l is lens with a 1 . 4 extender and 550 ex flash . the next image , to illustrate the dorsal abdominal pattern in comparison to the slough amberwing , was shot in eagle pass , maverick co . , texas in september , 2003 with a canon eos 10d and ef 70 - 200 mm f / 2 . 8 l is lens with 2x extender and 550 ex flash .

amphibians & reptiles birds mammals dragonflies fishes plants world biomes bird wing & tail images library resources publications pacific nw moths ( external site ) bug guide ( external site ) a catalogue of butterflies of the united states and canada , j . pelham , 2012 usfws feather atlas an identification manual to the small mammals of british coumbia tags silphidae of washington state

this list , including english names originally proposed in 1978 and now revised several times , is offered as both a current north american check - list and a list of english names . the english names were generated by dennis r . paulson and sidney w . dunkle ( a few were already in use ) and approved , with some modification , by the membership of the dragonfly society of the americas . common names for species recently added to the list were coined by the common names committe and then the checklist committee of the dsa or by the original describer . the list is kept up to date with taxonomic changes , name changes , and species newly added to the fauna .

slater museum of natural history 1500 n . warner st . # 1088 tacoma , wa 98416 253 . 879 . 3356

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justification : extent of occurrence is 3 , 955 , 500 km² and the species is known from 32 locations , several within protected areas .

found from honduras south to bolivia and brazil ( ris 1930 , rácenis 1953 , dunkle 1982 , donnelly 1992 , louton et al . 1996 , d . r . paulson pers . comm . 2006 , r . w . garrison 2006 ) .

the species is known from 32 localities , several situated within a protected area and thought to be widespread although no data is available on population numbers .

adults perch on marginal vegetation of ponds , lakes , pools , sloughs , swamps and streams ; female oviposits on twigs 1 - 2 inches above water ( ris 1930 ) . larva described by santos ( 1970 ) .

present in protected areas : reserva biologica la selva , heredia province , costa rica , parque nacional soberania , canal zone , panama , parque nacional manu , madre de dios department , peru , reserva de limoncocha , napo province , ecuador ( dunkle 1982 , donnelly 1992 , louton et al . 1996 , d . r . paulson pers . comm . 2006 ) .

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upcoming events 2018 bugguide gathering in virginia july 27 - 29 : registration and discussion photos of insects and people from the 2015 gathering in wisconsin , july 10 - 12 photos of insects and people from the 2014 gathering in virginia , june 4 - 7 . photos of insects and people from the 2013 gathering in arizona , july 25 - 28 photos of insects and people from the 2012 gathering in alabama photos of insects and people from the 2011 gathering in iowa photos from the 2010 workshop in grinnell , iowa photos from the 2009 gathering in washington

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this small , colorful skimmer is quick and alert . the length of the body varies from 0 . 8 to 1 . 0 inches . the thorax generally is brown with short , thin dorsal stripes and yellowish side spots . the abdomen is short and stout with variable patterns , either not striped or with narrow brown stripes . the abdomen for the male has an orange tint to it . the male ' s wings are tinted amber with yellow veins and red stigmas ( near the tips of wings ) . the female ' s wings are variable with brown spots and red stigmas . the face is yellowish . the legs are yellowish with black spines .

this species is common throughout eastern and central united states , from maine to arizona . eastern amberwings like permanent still or slowly moving waters such as ponds , lakes , ditches , and stream pools , but not bogs . it is widely distributed and fairly common throughout much of wisconsin .

shading illustrates monthly percentages of the total flight season records for the species . each flight season record is a unique date / location / observer combination where one or more adult or an exuvia was recorded ( excludes nymphs ) . the actual number of flight season records for each month is shown in parentheses . flight seasons begin earlier in the southern part of the state , often by a week or more . also , flight charts may not be accurate for rare species because of few data available .

this site is produced in conjunction with the wisconsin aquatic and terrestrial resources inventory and sponsored by the wisconsin department of natural resources . the information presented on this site is subject to the wisconsin department of natural resources ' legal notices , disclaimers , and terms of use .

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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 .

confused by a class within a class or an order within an order ? please see our brief essay .

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 .

this material is based upon work supported by the national science foundation grants drl 0089283 , drl 0628151 , due 0633095 , drl 0918590 , and due 1122742 . additional support has come from the marisla foundation , um college of literature , science , and the arts , museum of zoology , and information and technology services .

this is a small species with a brown thorax . there are two wide greenish stripes laterally and middorsally on the thorax . males have orange or amber wings that usually develop a brown spot above the triangles . females have variously shaped brown spots or stripes through the amber areas . both sexes have red pterostigmata . the abdomen is narrowed basally , but thick thereafter and brown with a dark row of chevrons dorsally .

total length : 19 - 25 mm ; abdomen : 12 - 16 mm ; hindwing : 16 - 21 mm .

slough amberwing ( p . domitia ) has dark brown stripes , not chevrons , dorsally on the abdomen . other skimmers with amber in wings are significantly larger .

this small dragonfly has been well studied . it has an elaborate courtship behavior . males come to the water ' s edge early in the morning in search of a territory . they then patrol and defend these territories , as potential egg laying sites , where they regularly perch on emergent sticks or twigs . these small territories , less than 5 square m , are only accepted by the male if he is not disturbed and there is no competition from other males . females appear and are courted by the male . he will fly out to her and lead her back to his prospective oviposition site , hovering with his abdomen turned up . upon acceptance by the female , signaled by a slower wing beat , the pair perch on a twig and mate , taking 20 - 30 sec . females then la y eggs either accompanied by the male or alone and guarded . females tap the abdomen against sticks or twigs within the oviposition area , attaching to it a gelatinous mass just above the waterline . the clumps of eggs released into water seem to explode into individual eggs as the clump drifts downward through the water . both sexes of this group mimic wasps by perching at the ends of grasses or weeds and , while beating their wings , pump the abdomen up and down . females also fly with the hindwings held together vertically with the abdomen bent up . females with more darkly pigmented wings tend to select the more favorable oviposition sites such as logs or sticks , and the lighter pigmented females select less favorable patches of floating vegetation . andromorphic females , with diffusely amber wings , are occasionally reported . one study found that males that were prevented from mating were much more likely to change potential oviposition sites the following day than males that were allowed t o mate , possibly implying that males use their reproductive success to determine the quality of oviposition sites .

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the male eastern amberwing shown here was in the davis mountains , jeff davis co . , texas , in june , 2005 . this shot was taken with a canon eos 1d mark ii and ef 600mm f / 4 l is lens and 2x extender and 580 flash .

the next three shots show male eastern amberwings at austin , travis co . , texas , in august , 2007 . these shots were taken with a canon eos 1d mark iii and the same lens and flash as the previous images .

the next two shots of a male eastern amberwing were taken in austin , texas , in july , 2008 , with the same gear as previously described .

the shot shown here shows a teneral female eastern amberwing that has just recently emerged . this image was taken in may , 2009 , at buescher state park , bastrop co . , texas with a canon eos 1d mark iii and ef 300mm f / 4 l is lens and 2x extender and 580 flash .

the female eastern amberwing shown here was in austin , travis co . , texas , in august , 2009 . this image was taken with a canon eos 1d mark iii and an ef 300mm f / 4 l is lens and 2x extender and 580 flash .

the female eastern amberwing shown here was in victoria , victoria co . , texas , in november , 2009 . a canon eos 1d mark iii and an ef 300mm f / 4 l is lens and 2x extender and 580 flash was used for this image .

the female eastern amberwing on the right was at a small pond on the caddo national grasslands , fannin co . , texas , in june , 2011 .

the male eastern amberwing on the right was in cameron , milam co . , texas , in september , 2011 . this shot was taken with a canon eos 1d mark iv and a sigma 50 - 500mm lens and canon 580 flash .

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