the chromodorididae , or chromodorids , are a taxonomic family of colorful , sea slugs ; dorid nudibranchs , marine gastropod mollusks in the superfamily doridoidea .

“ chromodorid nudibranchs are among the most gorgeously colored of all animals . ” the over 360 described species are primarily found in tropical and subtropical waters , as members of coral reef communities , specifically associated with their sponge prey .

the chromodorids are the most speciose family of opisthobranchs .

they range in size from < 1 mm to over 30 cm , although most species are approximately 2 – 3 cm in size .

although , they have a worldwide distribution , most species are found in the indo-pacific region .

a scientific paper published in 2007 , found the most widespread chromodorid genera , ( mexichromis , chromodoris , glossodoris and hypselodoris ) to be paraphyletic or polyphyletic .

the family cadlinidae bergh , 1891 has been considered a synonym of the chromodorididae . "

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 .

the chromodorid nudibranchs without cadlina are now monophyletic and turn out to be a possible sister to the family actinocyclidae .

cadlina and aldisa are the only two genera currently classified in the cadlinidae .

a comprehensive phylogeny of the chromodorid nudibranchs found every one of the 14 traditional chromodorid genera were either non-monophyletic , or rendered another genus paraphyletic .

the authors presented a new classification of the chromodorid nudibranchs , which used molecular data to untangle evolutionary relationships and at the same time retains a historical connection to traditional systematics by using generic names attached to type species as clade names .

all chromodorid nudibranchs feed on sponges .

chromodorididae

kento furui added the japanese common name
イロウミウシ科
to
chromodorididae
.
( opisthobranchia , chromodorididae ) . journal of molluscan studies 72 ( 2 ) : 214–216 .
bergh , 1898 ( doridoidea : chromodorididae ) . journal of natural history 35 : 1143–1171 .
edmunds m ( 1981 ) opishtobranchiate mollusca from ghana : chromodorididae . zoological journal of the linnean society 71 : 175–201 .
bergh , 1875 ( nudibranchia : chromodorididae ) in light of phylogenetic analysis . journal of molluscan studies 65 : 33–45 .
bergh , 1898 ( opisthobranchia : chromodorididae ) from patagonia , argentina . journal of molluscan studies 62 ( 3 ) : 265–273 .
ortea ja ( 1988 ) molluscos opisthobranquios del archipelago de cabo verde : chromodorididae . publicações ocasionais da sociedade portuguesa de malacologia 11 : 1–16 .
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 .
colour group ( mollusca , nudibranchia , chromodorididae ) , with remarks on the genus brachychlanis ehrenberg , 1831 . journal of natural history 35 : 1371–1398 .
stimpson , 1855 ( nudibranchia , chromodorididae ) with the description of a new species from the caribbean sea . journal of molluscan studies 72 : 189–198 .
turner lm , wilson ng ( 2008 ) polyphyly across oceans : a molecular phylogeny of the chromodorididae ( mollusca , nudibranchia ) . zoologica scripta 37 : 23–42 .
rudman wb , berquist pr ( 2007 ) a review of feeding specificity in the sponge - feeding chromodorididae ( nudibranchia : mollusca ) . molluscan research 27 ( 2 ) : 60–88 .
( nudibranchia : chromodorididae ) with a review of the monophyletic clade of indo - pacific species , including descriptions of twelve new species . zoological journal of the linnean society 125 : 1–125 .
rudman wb ( 1984 ) the chromodorididae ( opisthobranchia : mollusca ) of the indo - west pacific : a review of the genera . zoological journal of the linnean society 81 : 115–273 .
ortea j , valdés á , garcia - gomez jc ( 1996 ) review of the atlantic species of the family chromodorididae ( mollusca : nudibanchia ) of the blue chromatic group . avicennia supplement .
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 .
rudman , w . b . ( 1984 ) the chromodorididae ( opisthobranchia : mollusca ) of the indo - west pacific : a review of the genera . zoological journal of the linnean society 81 : 115 - 273 .
rudman , w . b . & bergquist , p . r . ( 2007 ) a review of feeding specificity in the sponge - feeding chromodorididae ( nudibranchia : mollusca ) . molluscan research , 27 ( 2 ) : 60 - 88
rudman , w . b . ( 1986 ) the chromodorididae ( opisthobranchia : mollusca ) of the indo - west pacific : noumea purpurea and chromodoris decora colour groups . zoological journal of the linnean society 86 ( 4 ) : 309 - 353 .
rudman , w . b . ( 1984 ) . the chromodorididae ( opisthobranchia : mollusca ) of the indo - west pacific : a review of the genera . zoological journal of the linnean society . 81 ( 2 ) : 115 - 273 . page ( s ) : 159 [ details ]
rudman , w . b . ( 1986 ) . the chromodorididae ( ophistobranchia : mollusca ) of the indo - west pacific : noumea purpurea and chromodoris decora colour groups . zoological journal of the linnean society . 86 ( 4 ) : 309 - 353 . page ( s ) : 319 [ details ]
gosliner , t . m . & johnson , r . f . ( 1999 ) phylogeny of hypselodoris ( nudibranchia : chromodorididae ) with a review of the monophyletic clade of indo - pacific species , including descriptions of twelve new species . zoological journal of the linnean society , 125 : 1 - 114 .
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 .
the family chromodorididae includes species that are often brightly colored and elaborately patterned , presumably advertising their distastefulness . most chromodorids store toxic chemicals from their sponge food in mantle glands , usually concentrated in a submarginal band on the notum , in order to repel predators . it is the largest indo - pacific nudibranch family , with more than 600 species , and it is well - represented in hawaii with about 43 species in 11 genera (
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 .
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 ] .
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 ) .
bergh [ 90 ] was the first to suggest a separate taxonomic rank for the chromodorid nudibranchs . johnson [ 28 ] showed that the chromodorididae are only monophyletic if cadlina is removed from the family , as cadlina is more closely related to aldisa and other dorid nudibranchs . we expanded on these preliminary results and confirmed the monophyly of the chromodorids in analyses without cadlina and without including as many dorid species . gosliner and johnson [ 101 ] reviewed the genus hallaxa and presented a morphological phylogeny of hallaxa and actinocylcus . they hypothesized that the semi - serial reproductive system found in species of hallaxa and actinocyclus and all chromodorids is a synapomorphy that unites the two groups . the chromodorid nudibranchs are monophyletic in every analysis , as is their sister group relationship with actinocyclidae . these analyses confirm the hypothesis of gosliner & johnson [ 101 ] and the preliminary findings of johnson [ 28 ] and confirm the utility of this morphological synapomorphy .
this paper contains new feeding information on 108 species of the sponge - feeding chromodorid nudibranchs and re - evaluates published information for 63 species . the combined information for 137 species shows a clear pattern of food specificity , at both the species and genus levels . new feeding information on the related actinocyclidae is also presented . species of chromodoris and related genera prefer darwinellids ; hypselodoris and its relatives feed on dysideids , and glossodoris feed exclusively on thorectids . the ' basal ' genera , cadlina , cadlinella are less specialised but both they and species of the anatomically similar family actinocyclidae seem linked by their common choice of halisarca . exceptions to the pattern suggest the genus chromodoris is polyphyletic . the evolution of feeding in the chromodorididae is discussed , and the patterns of food specificity are shown to strongly support prevailing hypotheses on chromodorid evolution . recent taxonomic studies within the sponge orders dictyoceratida and dendroceratida have been essential to this study , enabling the re - identification of many of the food sponges , and the use of marker secondary metabolites in both the nudibranchs and their sponge prey .
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 resulting classification can be used to address questions of interest to a much broader community . a robust phylogeny and corresponding revised classification are necessary to conduct comparative studies in the chromodorididae . evolutionary studies of trophic specialization , color patterns and secondary metabolites , for example , will be much more robust by comparing monophyletic units that are clearly named within a new classification . we have pointed out and highlighted many areas of future research . as we detail above , there are many taxonomic , nomenclatural and species delineation problems that still require refinement within the chromodorid nudibranchs . these questions can best be answered with a detailed examination of morphology together with molecular data . the molecular data needs to be rooted in sound identifications and definitions that are always based on morphology . this phylogeny does not answer all of these issues but it will serve as a framework to more effectively tackle these questions . our classification will serve as a more refined basis for other evolutionary biologists , ecologists and natural products chemists . their results will be more informative in light of a classification based on evolutionary history rather than one based on untested hypotheses . phylogenetic systematics provides a rigorous and repeatable methodology that permits iterative approximations of relationship and our understanding of this diverse and biologically intriguing group of organisms is enhanced by these studies .
traditional taxonomy has obscured the patterns of diversification in the chromodorids . a new classification that properly reflects evolutionary history is required . in the new classification , we only keep existing names , for species not supported in clades if it is not disruptive to the new classification . we also hypothesize the predicted phylogenetic position of taxa that have not yet been included in the phylogenetic analysis . we used the nomenclatural standards set by the international code of zoological nomenclature [ 57 ] . names and dates for genera , families and subfamilies we taken from bouchet et al ' s review of gastropod nomenclature [ 58 ] . every name used is resurrected from synonymy and proposed because the type species of the genus is found in the clade . if more than one generic type species is found in the same clade , the older name has priority . in this way , the history of naming in the chromodorids will be maintained . if the gender of a species ' new genus changes in the new classification , the gender of the specific epithet will be changed . additionally , if the incorrect specific name gender has been used , the proper gender will be used in the new classification . the proposed phylogenetic naming code , the phylocode , recommends naming clades when type species are part of the clade to be named , as we have done here , but does not use or recognize ranks as we have by using generic , subfamily and family names for clades . in many cases there is no conflict between the phylocode and traditional nomenlclature [ 59 ] . the phylocode has not been formally adopted , so there is no official system for naming in accordance with that code . in order to maintain stability and to avoid creating names that may change with the addition of new information , we used a method advocated by dayrat & gosliner [ 60 ] . this method advocates using the most inclusive known clade name as the first part of a species binomial for species that cannot be named without creating a new name . we will use the family name chromodorididae as the name for species that would create instability if the bionomials were unchanged or if new names were given . in our proposed classification we will also include incerte sedis species in the chromodorididae . in clades that are poorly supported ( posterior probabilities below 0 . 85 ) , we have used a generic name for members of those groups with the generic name placed in quotation marks . we prefer this method as an interim solution as it does not leave these taxa in taxonomic limbo and retain the use of single names for polyphyletic groups .
chromodorid nudibranchs ( 16 genera , 300 + species ) are beautiful , brightly colored sea slugs found primarily in tropical coral reef habitats and subtropical coastal waters . the chromodorids are the most speciose family of opisthobranchs and one of the most diverse heterobranch clades . chromodorids have the potential to be a model group with which to study diversification , color pattern evolution , are important source organisms in natural products chemistry and represent a stunning and widely compelling example of marine biodiversity . here , we present the most complete molecular phylogeny of the chromodorid nudibranchs to date , with a broad sample of 244 specimens ( 142 new ) , representing 157 ( 106 new ) chromodorid species , four actinocylcid species and four additional dorid species utilizing two mitochondrial markers ( 16s and coi ) . we confirmed the monophyly of the chromodorididae and its sister group relationship with the actinocyclidae . we were also able to , for the first time , test generic monophyly by including more than one member of all 14 of the non - monotypic chromodorid genera . every one of these 14 traditional chromodorid genera are either non - monophyletic , or render another genus paraphyletic . additionally , both the monotypic genera verconia and diversidoris are nested within clades . based on data shown here , there are three individual species and five clades limited to the eastern pacific and atlantic oceans ( or just one of these ocean regions ) , while the majority of chromodorid clades and species are strictly indo - pacific in distribution . we present a new classification of the chromodorid nudibranchs . we use molecular data to untangle evolutionary relationships and retain a historical connection to traditional systematics by using generic names attached to type species as clade names .
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 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 .
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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 ) .
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 ) .
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 .
we initially used standard phenol - cholorform extractions [ 42 ] , [ 43 ] to extract genomic dna and also used the dneasy spin column extraction method ( qiagen ) to extract genomic dna from the majority of our samples . we used universal primers to amplify , using pcr , double - stranded products from both the cytochrome oxidase 1 ( coi ) and 16s mitochondrial genes . we targeted a 658 bp fragment of coi using folmer et al ' s [ 44 ] universal primers and for 16s sequences , we used palumbi ' s [ 45 ] 16sar and 16sbr primers . we carried out the polymerase chain reaction in 25 µl reactions with one µl of genomic dna template . we used the second ( 200 µl ) elution from my extractions in dneasy ae buffer as the dna template in most reactions . if the amplification was difficult we used one µl of the first elution . for the phenol - cloroform and chelex extractions , we used dilutions of 1∶25 or 1∶50 . no matter the extraction method used , we included 2 . 5 µl of 10× pcr buffer , 0 . 5 µl dntps ( 10 mm stock ) , 0 . 25 µl of each primer ( 25 um stock ) , 0 . 75–0 . 85 µl mgcl ( 50 mm stock ) , 0 . 25 µl taq ( 5 units / µl ) - apex , biolase , usb hotstart - and 19 . 5 µl of ddh2o in each reaction tube . we ran all of the reactions on a biorad mycycler™ thermocycler ( software version 1 . 065 , bio - rad laboratories ) . coi segments were amplified with the following parameters : an initial denaturation at 94°c for three minutes , then , 39 cycles of denaturation at 94°c for 30 seconds ; annealing at 46°c for 30 seconds ; extension at 72°c for 60 seconds , these cycles were followed by extension at 72°c for five minutes . partial 16s sequences were amplified with the following parameters : an initial denaturation at 94°c for three minutes , then 39 cycles of denaturation at 94°c for 30 seconds ; annealing at 50–52°c for 30 seconds ; extension at 72°c for 60 seconds , these cycles were followed by extension at 72°c for five minutes and 25°c for 60 seconds . we used electrophoresis to view pcr products on 0 . 8 % tbe or tae agarose gel stained with ethidium bromide . we cleaned successful pcr products with exosap - it ( usb scientific ) following each product ' s standard protocol .
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 determined the best - fit model of evolution for each codon position for coi ( 1 st , 2 nd 3 rd ) and the 16s fragment using the aic selection from mr . modeltest ver . 2 [ 49 ] for each dataset . we ran bayesian phylogenetic analyses using mr . bayes 3 . 1 . 2 [ 50 ] – [ 52 ] . we ran a monte carlo - metropolis simulation for 50 , 000 , 000 generations for each dataset , with trees sampled every 1000 generations . data was partitioned by gene and by codon position in all combined analyses . we ran one analysis of two runs of six chains for all data sets . all other settings remained in the default and all parameters were unlinked to allow each partition to vary independently ( mr . bayes 3 . 12 manual , urltoken ) all trees saved before convergence of the runs and stationarity of likelihood values were discarded . we determined convergence and stationarity by plotting tree number against likelihood scores for each run to find the point where the likelihood plot leveled off and began to fluctuate around a stable value using tracer 1 . 4 [ 53 ] ( plots not shown ) . in all cases , the conservative estimate of a burnin of 25 % of sampled trees was well into this plateau . the remaining 75000 trees ( 37500 from each run ) were used to construct majority rule consensus trees and calculate posterior probabilities . all clades and support values are shown in the resulting phylogenies . all posterior probabilities are mapped on all trees . as suggested by hulsenbeck [ 50 ] , clades with posterior probabilities of 0 . 95–1 . 00 will be considered to be very well - supported . clades with support values of 0 . 85–0 . 94 will be considered supported . all posterior probabilities lower than 0 . 85 are considered poorly supported and should be viewed with caution , but all posterior probabilities are mapped on all trees . although taxa may appear as sister species , we can only know true sister species relationships if we have complete taxon sampling for the family .
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 .
of the 16 genera in the current chromodorid classficiation , only two , cadlinella and tyrinna , retain the same membership in our new classification . nine more generic names are retained ; ardeadoris , ceratosoma , chromodoris , diversidoris , glossodoris , hypselodoris , mexichromis , noumea and thorunna but their membership has changed . five names are synonymized with the names listed above : digidentis , durvilledoris , pectenodoris , risbecia and verconia . five older names : doriprismatica , felimare , felimida , goniobranchus and miamira , have been rescued from synonymy and are used to describe clades of species previously included in different genera . there are 17 generic names used in our new classification , each of which will be detailed below and in figure s3 . of these , 13 are very well supported with posterior probabilities ≥0 . 95 . mexichromis is supported with a posterior probability of 0 . 87 when variable positions are included and 0 . 95 when they are excluded . noumea consists of two separate clades ( both pp = 1 . 00 ) that are poorly supported as a combined clade in the analysis when variable positions are included ( pp = 0 . 61 ) . although , this support is not sufficient , all of the species in both of these clades are currently named noumea and will retain this name in order to maintain stability . doriprismatica is extremely poorly - supported pp = 0 . 64 with variable positions included and well - supported ( pp = 0 . 92 ) in the analysis with variable positions excluded ( figures s2 , s3 ) . we do not consider this level of support sufficient to definitively name this clade , but because continuing to use the current name , chromodoris , would add greater confusion ( as that name represents a different , well - supported monophyletic group ) we will premilinarily name these species ‘ doriprismatica’ . similarly , a group of species some of which are currently classified as chromodoris and some as glossodoris form a polytomy together with other well supported clades . as these species need a name , but lack appropriate support , they cannot be named definitively . we will preliminaryily name these species ‘felimida’ . naming these clades is much more stable than using names that now represent other well - defined and well supported clades . these names are hypotheses ; with more data the relationships of members of these clades will likely become better resolved . the completed classisifiction is listed in table s2 .
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 )


anarsia choana is a moth of the gelechiidae family .

it was described by park in 1995 .

it is found in taiwan and japan ( ryukyus ) .

the length of the forewings is 4.5 – 4.7 millimetres ( 0.18 – 0.19 in ) .

the forewings are creamy white , speckled with pale brownish grey and scattered with blackish scales .

there is a blackish dot on the extreme base of the costa .

the hindwings are pale brownish grey , hyaline on the basal half and with the veins darker .

anarsia choana

anarsia choana park , 1995 ; tropical lepid . 6 ( 1 ) : 61 ; tl : taipei co . , taiwan
anarsia choana ; ponomarenko , 1997 , far east . ent . 50 : 54 ; ueda , 2010 , trans . lepid . soc . japan 61 : 272 ; park & bae , 2018 , zootaxa 4399 ( 2 ) : ( 272 - 280 )
anarsia chiangmaiensis ; ponomarenko , 1997 , far east . ent . 50 : 54
anarsia conica ; ponomarenko , 1997 , far east . ent . 50 : 54
anarsia lewvanichae ; ponomarenko , 1997 , far east . ent . 50 : 55
anarsia melanoplecta ; ponomarenko , 1997 , far east . ent . 50 : 55
anarsia procera ; ponomarenko , 1997 , far east . ent . 50 : 56
anarsia spatulana ; ponomarenko , 1997 , far east . ent . 50 : 56
anarsia eleagnella kuznetsov , 1957 ; zool . zhurn . 36 ( 7 ) : 1096
anarsia ulneongensis park & ponomarenko , 1996 ; korean j . ent . 26 : 343
figures 52 – 55 . male genitalia : 52 , a . similicampa park , sp . nov . , 52 a , close - up uncus ; 53 , a . choana park ; 53 a , left valva + tegumen of the same species ; 53 b , tegumen + right valva of the same species ; 53 c , ditto , abdominal sternite viii ; 54 , a . kepensis park , sp . nov . , left valva ; 54 a , ditto , right valva + tegumen ; 54 b , ditto , abdominal sternite viii ; 55 , a . didymopa meyrick .
anarsia asymmetrodes park , 2014 ; ent . res . 44 : 18 ; tl : baengnyeongdo
anarsia callicosma janse , 1960 ; moths s . afr . 6 ( 2 ) : 214
anarsia pinnata meyrick , 1931 ; exotic microlep . 4 ( 2 - 4 ) : 69
anarsia pustulata janse , 1949 ; moths s . afr . 5 ( 1 ) : 32
anarsia ulneongensis ; ueda , 2010 , trans . lepid . soc . japan 61 : 275
anarsia amegarta meyrick , 1933 ; exotic microlep . 4 ( 12 ) : 360 ; tl : java
anarsia hippocoma meyrick , 1921 ; exotic microlep . 2 ( 14 ) : 429 ; tl : queensland
anarsia sibirica ; ponomarenko , 1997 , far east . ent . 50 : 56 ; [ fe ]
anarsia vinsonella viette , 1957 ; mém . inst . sci . madagascar ( e ) 8 : 163
anarsia kepensis bae , shin , na & park , 2016 , sp . nov . - plazi treatmentbank
anarsia altercata ; [ nhm card ] ; ponomarenko , 1997 , far east . ent . 50 : 54
anarsia amegarta ; [ nhm card ] ; ponomarenko , 1997 , far east . ent . 50 : 54
anarsia eburnella ; [ nhm card ] ; ponomarenko , 1997 , far east . ent . 50 : 54
anarsia ephippias ; [ nhm card ] ; ponomarenko , 1997 , far east . ent . 50 : 54
anarsia epotias ; [ nhm card ] ; ponomarenko , 1997 , far east . ent . 50 : 55
anarsia eutacta meyrick , 1921 ; zool . meded . leyden 6 : 163 ; tl : java , pekalongan
anarsia eutacta ; [ nhm card ] ; ponomarenko , 1997 , far east . ent . 50 : 55
anarsia geminella ; [ nhm card ] ; ponomarenko , 1997 , far east . ent . 50 : 55
anarsia halimodendri ; [ nhm card ] ; ponomarenko , 1997 , far east . ent . 50 : 55
anarsia libanoticella ; [ nhm card ] ; ponomarenko , 1997 , far east . ent . 50 : 55
anarsia longipalpella rebel , 1907 ; denksch . akad . wiss . wien . 71 ( 2 ) : 124
anarsia melanchropa ; [ nhm card ] ; ponomarenko , 1997 , far east . ent . 50 : 55
anarsia nuristanella ; [ nhm card ] ; ponomarenko , 1997 , far east . ent . 50 : 55
anarsia omoptila ; [ nhm card ] ; ponomarenko , 1997 , far east . ent . 50 : 56
anarsia sthenarota ; [ nhm card ] ; ponomarenko , 1997 , far east . ent . 50 : 56
anarsia veruta ; [ nhm card ] ; ponomarenko , 1997 , far east . ent . 50 : 56
anarsia aleurodes meyrick , 1922 ; exotic microlep . 2 ( 16 ) : 502 ; tl : mesopotamia , museyib
anarsia altercata meyrick , 1918 ; exotic microlep . 2 ( 5 ) : 148 ; tl : bengal , pusa
anarsia crassipalpella legrand , 1966 ; mém . mus . hist . nat . paris ( a ) 37 : 78
anarsia ephippias meyrick , 1908 ; ent . mon . mag . 44 : 197 ; tl : pusa , bengal
anarsia euphorodes meyrick , 1922 ; exotic microlep . 2 ( 16 ) : 503 ; tl : china , shanghai
anarsia inserta [ sic , recte incerta ] ; ponomarenko , 1997 , far east . ent . 50 : 55
anarsia luticostella chrétien , 1915 ; ann . soc . ent . fr . 84 : 332 ; tl : biskra
anarsia nigrimacula janse , 1949 ; moths s . afr . 5 ( 1 ) : 29 ; tl : umkomaas
anarsia reciproca meyrick , 1920 ; exotic microlep . 2 ( 10 ) : 300 ; tl : madras , coimbatore
anarsia retamella chrétien , 1915 ; ann . soc . ent . fr . 84 : 331 ; tl : gafsa
anarsia sthenarota meyrick , 1926 ; sarawak mus . j . 3 : 153 ; tl : mt murud , 6500ft
anarsia triglypta meyrick , 1933 ; exotic microlep . 4 ( 12 ) : 354 ; tl : pusa , bihar
anarsia veruta meyrick , 1918 ; exotic microlep . 2 ( 5 ) : 148 ; tl : bengal , pusa
anarsia didymopa meyrick , 1916 ; exot . microlep . 1 ( 19 ) : 583 ; tl : bengal , pusa
anarsia epotias meyrick , 1916 ; exot . microlep . 1 ( 19 ) : 583 ; tl : bengal , pusa
anarsia idioptila meyrick , 1916 ; exot . microlep . 1 ( 19 ) : 582 ; tl : bengal , pusa
anarsia mitescens meyrick , 1913 ; ann . transv . mus . 3 ( 4 ) : 299 ; tl : barberton
anarsia sagmatica meyrick , 1916 ; exot . microlep . 1 ( 19 ) : 582 ; tl : bengal , pusa
anarsia albibasella janse , 1963 ; moths s . afr . 6 ( 3 ) : 253 ; tl : sw . africa
anarsia amalleuta meyrick , 1913 ; ann . transv . mus . 3 ( 4 ) : 298 ; tl : three sisters
anarsia anthracaula meyrick , 1929 ; exot . microlep . 3 ( 16 ) : 512 ; tl : new hebrides , efate
anarsia beitunica li & zheng , 1997 ; acta zool . hung . 43 ( 2 ) : ( 121 - 132 )
anarsia bimaculata ponomarenko , 1989 ; ent . obozr . 68 ( 3 ) : 635 ; tl : gomotaezhnoe , primorskii krai
anarsia decora li & zheng , 1997 ; acta zool . hung . 43 ( 2 ) : ( 121 - 132 )
anarsia elongata park , 1995 ; tropical lepid . 6 ( 1 ) : 64 ; tl : taichung co . , taiwan
anarsia eximia li & zheng , 1997 ; acta zool . hung . 43 ( 2 ) : ( 121 - 132 )
anarsia largimacularis li & zheng , 1997 ; acta zool . hung . 43 ( 2 ) : ( 121 - 132 )
anarsia magnibimaculata li & zheng , 1997 ; acta zool . hung . 43 ( 2 ) : ( 121 - 132 )
anarsia melanchropa meyrick , 1926 ; exot . microlep . 3 ( 9 ) : 281 ; tl : india , dehra dun
anarsia novitricornis li & zheng , 1997 ; acta zool . hung . 43 ( 2 ) : ( 121 - 132 )
anarsia omoptila meyrick , 1918 ; exotic microlep . 2 ( 5 ) : 147 ; tl : s . india , coimbatore
anarsia sibirica park & ponomarenko , 1996 ; acta zool . hung . 42 ( 1 ) : 78 ; tl : novosibirsk
anarsia squamerecta li & zheng , 1997 ; acta zool . hung . 43 ( 2 ) : ( 121 - 132 )
anarsia balioneura meyrick , 1921 ; ann . transv . mus . 8 ( 2 ) : 79 ; tl : rhodesia , umtali
anarsia eburnella christoph , 1887 ; in romanoff , mém . lépid . 3 : 122 , pl . 5 , f . 14
anarsia libanoticella amsel , 1967 ; beitr . naturk . forsch . südwdtl . 26 ( 3 ) : 21 ; tl : lebanon
anarsia sciotona meyrick , 1927 ; exot . microlep . 3 ( 12 ) : 353 ; tl : cape colony , east london
anarsia spartiella ; [ nhm card ] ; ponomarenko , 1997 , far east . ent . 50 : 56 ; [ fe ]
anarsia spicata meyrick , 1918 ; ann . transv . mus . 6 ( 2 ) : 21 ; tl : transvaal , pretoria
anarsia subfulvescens meyrick , 1918 ; ann . transv . mus . 6 ( 2 ) : 21 ; tl : natal , umkomaas
anarsia acaciae walsingham , 1896 ; proc . zool . soc . lond . 1896 : 278 ; tl : sw . arabia , aden
anarsia anisodonta diakonoff , 1954 ; verh . k . akad . wet . amst . ( 2 ) 50 ( 1 ) : 14
anarsia arachniota meyrick , 1925 ; bull . soc . ent . egypte 9 ( 1 - 3 ) : 210 ; tl : egypt
anarsia carbonaria meyrick , 1913 ; ann . transv . mus . 3 ( 4 ) : 299 ; tl : barberton , waterval onder
anarsia chaonella park , 1995 ; tropical lepid . 6 ( 1 ) : 61 ; tl : taiwan , tapei co . , taihoku
anarsia incerta ueda , 1997 ; trans . lepid . soc . japan 48 ( 2 ) : 80 ; tl : ryukyus , japan
anarsia permissa meyrick , 1926 ; ann . s . afr . mus . 23 : 331 ; tl : sw . africa , windhoek
anarsia triaenota meyrick , 1913 ; j . bombay nat . hist . soc . 22 ( 1 ) : 169 ; tl : gooty
anarsia citromitra meyrick , 1921 ; ann . transv . mus . 8 ( 2 ) : 80 ; tl : portuguese east africa , magude
anarsia geminella amsel , 1967 ; beitr . naturk . forsch . südwdtl . 26 ( 3 ) : 18 ; tl : herat , afghanistan
anarsia nigricana park , 1991 ; jpn . j . ent . 59 ( 3 ) : 494 ; tl : suweon , gyunggi prov .
anarsia vectaria meyrick , 1918 ; ann . transv . mus . 6 ( 2 ) : 21 ; tl : natal , sarnia ; umkomaas
anarsia aspera park , 1995 ; tropical lepid . 6 ( 1 ) : 57 ; tl : taiwan , orchid is . , 4km sw hungta
anarsia isogona meyrick , 1913 ; j . bombay nat . hist . soc . 22 ( 1 ) : 169 ; tl : nilgiris , 3500ft
anarsia melanoplecta meyrick , 1914 ; j . bombay nat . hist . soc . 22 ( 4 ) : 774 ; tl : pusa , bengal
anarsia pensilis meyrick , 1913 ; j . bombay nat . hist . soc . 22 ( 1 ) : 168 ; tl : maskeliya , ceylon
anarsia sagittaria meyrick , 1914 ; j . bombay nat . hist . soc . 22 ( 4 ) : 774 ; tl : pusa , bengal
anarsia nimbosa meyrick , 1913 ; ann . transv . mus . 3 ( 4 ) : 300 ; tl : three sisters , pretoria , waterval onder
anarsia nimbosa ; meyrick , 1921 , ann . transv . mus . 8 ( 2 ) : 79 ; [ nhm card ] ; [ afromoths ]
figures 22 – 40 . labial palpus of male and female : 22 , anarsia trichornis meyrick ; 22 a , ditto , female ; 23 , a . isogona meyrick ; 24 , a . paraisogona park ; 25 , a . incerta ueda ; 26 , a . deuterodes park , sp . nov . ; 27 , a . phortica meyrick ; 28 , a . diversiola park , sp . nov . ; 29 , a . porthmista park , sp . nov . ; 29 a , ditto , female ; 30 . a . acerata meyrick , female ; 31 , a . gryphodes park , sp . nov . ; 32 , a . campestra park , sp . nov . ; 33 , a . similicampa park , sp . nov . , 34 , a . choana park ; 35 , a . kepensis park , sp . nov . ; 36 . a . melanodes park , sp . nov . ; 37 , a . didymopa meyrick ; 38 , a . pusillidia park , sp . nov . ; 39 , a . patulella ( walker ) ; 39 a , ditto , female ; 40 , a . houhunlii park , sp . nov .
anarsia acerata meyrick , 1913 ; j . bombay nat . hist . soc . 22 ( 1 ) : 169 ; tl : n . coorg , 3500ft
anarsia acrotoma meyrick , 1913 ; j . bombay nat . hist . soc . 22 ( 1 ) : 169 ; tl : n . coorg , 3500ft
anarsia lechriosema bradley , 1982 ; j . nat . hist . 16 ( 3 ) : 375 ; tl : norfolk i . , mt bates , 290m
anarsia stylota meyrick , 1913 ; j . bombay nat . hist . soc . 22 ( 1 ) : 168 ; tl : maskeliya and patipola , ceylon
anarsia semnopa meyrick , 1921 ; ann . transv . mus . 8 ( 2 ) : 79 ; tl : rhodesia , umtali ; portuguese east africa , magude
anarsia tortuosella amsel , 1967 ; beitr . naturk . forsch . südwdtl . 26 ( 3 ) : 19 ; tl : chingi , salt range , w pakistan
anarsia psammobia falkovitsh & bidzilya , 2003 ; proc . zool . mus . kiev nat . taras shevchenko univ . 1 ( 1 ) : ( 113 - 147 )
anarsia tricornis meyrick , 1913 ; j . bombay nat . hist . soc . 22 ( 1 ) : 168 ; tl : maskeliya , perdeniya and haldamulla , ceylon
anarsia arsenopa meyrick , 1920 ; in alluaud & jeannel , voyage afr . orientale , ins . lép . 2 : 72 ; tl : british east africa , nairobi forest
anarsia epiula meyrick , 1904 ; proc . linn . soc . n . s . w . 29 ( 2 ) : 418 ; tl : sydney , new south wales
anarsia gajiensis park & ponomarenko , 1996 ; acta zool . hung . 42 ( 1 ) : 75 ; tl : mt gaji - san , gyungnam prov . , korea
anarsia leucophora meyrick , 1904 ; proc . linn . soc . n . s . w . 29 ( 2 ) : 417 ; tl : broken hill , new south wales
anarsia ovula ; ponomarenko , 1997 , far east . ent . 50 : 56 ; park & bae , 2018 , zootaxa 4399 ( 2 ) : ( 272 - 280 )
anarsia paraisogona ; ponomarenko , 1997 , far east . ent . 50 : 56 ; park & bae , 2018 , zootaxa 4399 ( 2 ) : ( 272 - 280 )
anarsia protensa park , 1995 ; tropical lepid . 6 ( 1 ) : 60 ; tl : taiwan , nantou co . , mei - feng , 30km s tayuling , 2200m
anarsia tortuosa ; ueda , 1997 , trans . lepid . soc . japan 48 ( 2 ) : 90 ; ponomarenko , 1997 , far east . ent . 50 : 56
anarsia halimodendri christoph , 1877 ; horae soc . ent . ross . 12 ( 3 ) : 297 , ( 4 ) pl . 8 , f . 69 ; tl : turkmenistan
anarsia inculta walsingham , 1891 ; trans . ent . soc . lond . 1891 ( 1 ) : 112 , pl . 5 , f . 49 ; tl : bathurst , gambia
anarsia nigricana ; ponomarenko , 1997 , far east . ent . 50 : 55 ; bae , lee & park , 2014 , ent . res . 44 : 19 ( list )
anarsia nuristanella amsel , 1967 ; beitr . naturk . forsch . südwdtl . 26 ( 3 ) : 19 , pl . 6 , f . 1 ; tl : nuristan , afghanistan
anarsia silvosa ueda , 1997 ; trans . lepid . soc . japan 48 ( 2 ) : 88 ; tl : japan , honshu , oita pref . , shonai town , shiramiz
anarsia minutella ; sattler , 1976 , bull . br . mus . nat . hist . ( ent . ) 34 ( 2 ) : 140 ( note ) ; [ nhm card ]
anarsia stepposella ponomarenko , 2002 ; far east . ent . 115 : 2 ; tl : russia , tuva republic , 50°44 ' n 93°08 ' e , east tannu ola mts , irbitei , 1000m
anarsia taurella bradley , 1961 ; bull . brit . mus . ( n . h . ) ent . 10 ( 4 ) : 134 , pl . 5 , f . 14 ; tl : guadalcanal , honiara
anarsia ulmarata bradley , 1961 ; bull . brit . mus . ( n . h . ) ent . 10 ( 4 ) : 134 , pl . 5 , f . 15 ; tl : guadalcanal , honiara
anarsia phortica meyrick , 1913 ; j . bombay nat . hist . soc . 22 ( 1 ) : 167 ; tl : maskeliya , kegalle , haldamulla and undugoda , ceylon ; n . coorg ; kuching , borneo
figures 1 – 21 . adults and venations . 1 , anarsia trichornis meyrick , female ; 2 , a . isogona meyrick , male ; 3 , a . paraisogona park , male ; 4 , a . incerta ueda , male ; 5 , a . deuterodes park , sp . nov . , male , holotype ; 6 . a . photica meyrick , male ; 7 , a . diversiola park , sp . nov . , male , holotype ; 8 , a . porthmista park , sp . nov . , female , paratype ; 9 , a . acerata meyrick , male ; 10 , a . gryphodes park , sp . nov . , male , holotype ; 10 a , ditto , hair pencil on underside of forewing , 11 , a . campestra park , sp . nov . , male , holotype ; 12 , a . similicampa park , sp . nov . , male , holotype ; 13 , a . choana park , male ; 14 , a . kepensis park , sp . nov . , male , holotype ; 15 , a . melanodes park , sp . nov . , male . holotype ; 15 a , ditto , hair pencil on underside of hindwing ; 15 b . ditto , female . 16 , a . didymopa meyrick , male ; 17 , a . pusillidia park , sp . nov . , male , holotype ; 18 , a . patulella ( walker ) , male ; 19 , a . houhunlii park , sp . nov . , male , holotype ; 19 a , ditto , hair pencil on underside of forewing ; 20 , hindwing venation of a . patulella walker ; 21 , hindwing venation of a . porthmista park , sp . nov . .
anarsia malagasyella viette , 1968 ; bull . mens . soc . linn . lyon 37 ( 2 ) : 89 ; tl : madagascar , env maroantsetra , forest station farankaraina , route navana , km 16 , 5 , antoroka valley , 100m
anarsia molybdota meyrick , 1904 ; proc . linn . soc . n . s . w . 29 ( 2 ) : 417 ; tl : toowoomba , queensland ; sydney , new south wales ; gisborne , victoria ; carnarvon , perth and york , west australia
anarsia bimaculata ; park , 1991 , jpn . j . ent . 59 ( 3 ) : 496 ; ueda , 1997 , trans . lepid . soc . japan 48 ( 2 ) : 86 ; ponomarenko , 1997 , far east . ent . 50 : 54 ; ponomarenko , park & bae , 2006 , j . asia - pacif . ent . 9 ( 2 ) : 113
bae , yang - seop , shin , young - min , na , sol - moon & park , kyu - tek , 2016 , the genus anarsia in cambodia and the northern vietnam ( lepidoptera , gelechiidae ) , with descriptions of ten new species and a catalogue of the genus in the central - east asia , zootaxa 4061 ( 3 ) , pp . 227 - 252 : 232
bae , yang - seop , shin , young - min , na , sol - moon & park , kyu - tek , 2016 , the genus anarsia in cambodia and the northern vietnam ( lepidoptera , gelechiidae ) , with descriptions of ten new species and a catalogue of the genus in the central - east asia , zootaxa 4061 ( 3 ) , pp . 227 - 252 : 230
bae , yang - seop , shin , young - min , na , sol - moon & park , kyu - tek , 2016 , the genus anarsia in cambodia and the northern vietnam ( lepidoptera , gelechiidae ) , with descriptions of ten new species and a catalogue of the genus in the central - east asia , zootaxa 4061 ( 3 ) , pp . 227 - 252 : 237 - 238
figures 41 – 43 a . male genitalia ( phl : phallus ) : 41 , anarsia trichornis meyrick ; 41 a , ditto , that of the same species ; 41 b , ditto , abdominal sternite viii ; 42 , a . isogona meyrick ; 42 a , ditto , abdominal sternite viii ; 42 b , ditto , left valve of a taiwanese specimen ; 43 , a . paraisogona park ; 43 a , ditto , that of the same species .
larva on ( in seed pods ) acacia edgworthii , a . farnesiana walsingham , 1896 , proc . zool . soc . lond . 1896 : 279
ananarsia acerata ; ponomarenko , 1997 , far east . ent . 50 : 50
ananarsia acrotoma ; ponomarenko , 1997 , far east . ent . 50 : 51
ananarsia aleurodes ; ponomarenko , 1997 , far east . ent . 50 : 51
larva on albizzia sp . ponomarenko , 1997 , far east . ent . 50 : 54
chelaria antisaris meyrick , 1913 ; ann . transv . mus . 3 ( 4 ) : 297 ; tl : barberton
ananarsia arachniota ; ponomarenko , 1997 , far east . ent . 50 : 51
ananarsia aspera ; ponomarenko , 1997 , far east . ent . 50 : 51
chelaria austerodes meyrick , 1918 ; ann . transv . mus . 6 ( 2 ) : 22 ; tl : transvaal , pretoria
ananarsia belutschistanella amsel , 1959 ; stuttgart . beitr . naturk . 28 : 33 ; tl : baluchistan , iran
ananarsia belutschistanella ; ponomarenko , 1997 , far east . ent . 50 : 51
japan , korea , primorye , china ( jilin ) . see [ maps ]
larva on maackia amurensis ponomarenko , 1997 , far east . ent . 50 : 54
chelaria bipinnata meyrick , 1932 ; exotic microlep . 4 ( 7 ) : 200 ; tl : gifu , japan
ananarsia bipinnata ; ponomarenko , 1997 , far east . ent . 50 : 51
larva on elaeagnus multiflora , e . umbellata , acer ginnala , quercus sp . ueda , 1997 , trans . lepid . soc . japan 48 ( 2 ) : 83
nothris centrospila turner , 1919 ; proc . r . soc . qd 31 ( 10 ) : 165 ; tl : qeensland , brisbane
ananarsia didymopa ; ponomarenko , 1997 , far east . ent . 50 : 51
romania , s . ukraine , seeu , altai , transcaucasia , turkmenistan , kazakhstan , afghanistan . see [ maps ]
ananarsia eleagnella ; ponomarenko , 1997 , far east . ent . 50 : 51
ananarsia elongata ; ponomarenko , 1997 , far east . ent . 50 : 51
larva on arachis hypogaea ponomarenko , 1997 , far east . ent . 50 : 54
chelaria eriozona meyrick , 1921 ; ann . transv . mus . 8 ( 2 ) : 80 ; tl : british s . e . africa , bela vista ; portuguese east africa , magude
ananarsia euphorodes ; ponomarenko , 1997 , far east . ent . 50 : 52
ananarsia gajiensis ; ponomarenko , 1997 , far east . ent . 50 : 52 ; bae , lee & park , 2014 , ent . res . 44 : 19 ( list )
guiera bradley , 1969 ; bull . ent . res . 59 ( 1 ) : 79
larva on halimodendron eichvaldii ponomarenko , 1997 , far east . ent . 50 : 55
ananarsia idioptila ; ponomarenko , 1997 , far east . ent . 50 : 52
s . india , laos , china ( zhejiang , yunnan ) , taiwan , japan . see [ maps ]
larva on schima sp . ponomarenko , 1997 , far east . ent . 50 : 52
ceu , seu , asia minor , n . africa , syria , caucasus , transcaucasia , afghanistan , china , india , australia , . . . . see [ maps ]
larva on prunus spp . , p . avium , p . spinosa , p . domestica , p . insititia
larva on prunus spinosa , malus spp . , amerniaca spp . , persica spp . , cerasus spp . , amygdalus spp . , acer tataricum ponomarenko , 1997 , far east . ent . 50 : 52
nothris minutella turati , 1929 ; boll . lab . zool . portici 23 : 124 , f . 4
larva on glycine max park , 1991 , jpn . j . ent . 59 ( 3 ) : 495
larva on cajanus indicus meyrick , 1918 , exotic microlep . 2 ( 5 ) : 147
s . india , ceylon , laos , thailand , shanghai , taiwan , queensland . see [ maps ]
gelechia patulella walker , 1864 ; list spec . lepid . insects colln br . mus . 29 : 635 ; tl : ceylon
larva on prunus salicina , nephelium sp . ponomarenko , 1997 , far east . ent . 50 : 52
ananarsia pensilis ; ponomarenko , 1997 , far east . ent . 50 : 52
s . india , ceylon , thailand , laos , borneo . see [ maps ]
ananarsia protensa ; ponomarenko , 1997 , far east . ent . 50 : 53
larva on elaeagnus pungens ponomarenko , 1997 , far east . ent . 50 : 53
ananarsia reciproca ; ponomarenko , 1997 , far east . ent . 50 : 53
ananarsia sagittaria ; ponomarenko , 1997 , far east . ent . 50 : 53
ananarsia sagmatica ; ponomarenko , 1997 , far east . ent . 50 : 53
chelaria sciograpta meyrick , 1921 ; ann . transv . mus . 8 ( 2 ) : 80 ; tl : transvaal , pretoria
larva on ( in fruit ) mimusops capensis meyrick , 1927 , exot . microlep . 3 ( 12 ) : 353
seu , ceu , sw . siberia , transbaikalia , libya , asia minor , mongolia . see [ maps ]
larva on sarothamnus scoparius , genista tinctoria , lembotropis nigrans , ulex spp . ponomarenko , 1997 , far east . ent . 50 : 56
ananarsia stylota ; ponomarenko , 1997 , far east . ent . 50 : 53
chelaria tortuosa meyrick , 1913 ; j . bombay nat . hist . soc . 22 ( 1 ) : 165 ; tl : matale , ceylon
ananarsia tortuosella ; ponomarenko , 1997 , far east . ent . 50 : 53
ananarsia triaenota ; ponomarenko , 1997 , far east . ent . 50 : 53
ananarsia triglypta ; ponomarenko , 1997 , far east . ent . 50 : 53
larva on inga dulcis ponomarenko , 1997 , far east . ent . 50 : 56
[ afromoths ] de prins , j . & de prins , w . , 2013
sangmi lee , richard brown & sibyl bucheli . gelechioidea - a global framework ;
[ 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 .
microlepidoptera from the solomon islands . additional records and descriptions of microlepidoptera collected in the solomon islands by the rennell island expedition 1953 - 54
microlepidoptera of new guinea , results of the third archbold expedition ( american - netherlands indian expedition 1938 - 1939 ) . part iv
voyage de ch . alluaud et r . jeannel en afrique orientale ( 1911 - 1812 ) . résultats scientifique . insectes lépidoptères . 2 . microlepidoptera in alluaud & jeannel ,
in la faune entomologique de i ' ile de lar réunion . lépidoptères ( excepté les tordeuses et les géométrides )
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 .
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transactions of the lepidopterological society of japan . ( journal , magazine , 1995 ) [ worldcat . org ]
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transactions of the lepidopterological society of japan . / nihon rinshi gakkai . ; ; osaka , japan : lepidopterological society of japan , [ 1995 - 2010 ]
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; clarke , 1969 : 246 ; park and ponomarenko , 1996 : 41 .
; ponomarenko , 1997 : 53 , 2009 : 342 . tl : maskeliya , sri lanka . the type in bmnh .
) : wingspan , 13 . 0 mm . the species is characterized by the extremely large trapezoidal costal patch occupying 1 / 3 the length of costa , with the length of the lower margin nearly same as the upper margin , and an expansible blackish long hairpencil near base on underside of forewing well developed ; the second segment of labial palpus in male ( fig . 27
) quadrate , upturned , strongly angled near base . the forewing is broad , with obtuse apex ; the ground color is grayish , with brownish scales sparsely scattered in basal 1 / 3 , densely covered with brownish scales in middle , then grayish white mottled with dark brown scales in distal part of wing . the hindwing has a broadly developed anterior expansion , abruptly oblique beyond middle . the female is unknown .
a ) : uncus more or less triangular , curved apically , with antero - lateral process , apex pointed . tegumen short , shorter than valvae . left valva roundly expanded in basal 1 / 3 on ventral margin , with slender , heavily sclerotized , s - shaped process , extending to 2 / 3 of valva ; costa slightly concave beyond middle . right valva with a large spatulate lobe on ventral margin near base ; zone with palmately modified scales in distal end . phallus strongly curved before middle , then s - shaped .
material examined . vietnam : 2 ♂ , bac khan prov . , ba be nat . park , 26–28 vii 2006 ( park , chae , & cuong ) , gen . slide no . cis - 6425 .
distribution . vietnam ( new record , vinh phuc prov . ) , n india , sri lanka , thailand ( n & w ) , malaysian borneo ( kutching ) .
figures 44 – 47 a . male genitalia : 44 , a . incerta ueda ; 44 a , ditto , abdominal sternite viii ; 45 , a . deuterodes park , sp . nov . , left and right valva ; 45 a , ditto , tegumen + uncus ; 45 b , ditto , abdominal sternite viii ; 46 . a . phortica meyrick ; 46 a , ditto , phallus ; 47 , a . diversiola park , sp . nov . ; 47 a , ditto , different view of the same species .
no known copyright restrictions apply . see agosti , d . , egloff , w . , 2009 . taxonomic information exchange and copyright : the plazi approach . bmc research notes 2009 , 2 : 53 for further explanation .
holotype : ♂ , cambodia , kep , 11 xi 2011 ( ys bae et al . ) , gen slide no . cis - 6437 .
diagnosis . this new species is easily distinguished from its allies by having very characterized the male labial palpus which is shortly developed and visible , while the third segment atrophied in most known species , and the second segment with an additional , upturned scale tuft arising from base of inner surface .
) . wingspan , 11 mm . head pale brownish gray on dorsal surface , blackish on face , with brownish scales laterally . flagellum of antenna with indistinct brownish annulations . second segment of labial palpus in male ( fig . 35
) very large , quadrate , with scale tuft antero - ventrally , longer than width of head , densely covered with dark brown scales on outer surface ; inner surface black , shiny , except median white zone beyond middle , with an additional upturned scale tuft arising from base , upturned ; third segment shortly developed , visible . tegula covered with blackish scales on anterior half , brownish scales on posterior half . thorax brownish dorsally . forewing elongate ; ground color brownish orange , mottled with dark brown scales ; a small black streak near base of cell ; a longitudinal long blackish streak near base to 2 / 5 along plical line , discal streak black , as long as 1 / 5 the length of forewing , and another blackish streak from end of cell toward costa before apex ; costa nearly straight ; costal patch elongate preceded by the similar , smaller dots and followed by three smaller ones on costa ; long whitish hair pencil on the underside of the forewing absent ; apex more or less obtuse ; termen oblique ; fringe grayish . hindwing elongate , grayish white , hyaline before middle , brownish scales along veins ; broad anterior expansion on costa extended to middle , then oblique ; apex acute .
a–b ) : abdominal sternite viii convex on caudal margin , with pocket - like lateral sacs on anterior margin ( fig . 54
b ) . uncus short , with acute apex , downward apically ; socius elongate . tegumen as long as valve , with almost parallel sides . valvae asymmetrical , but more or less similar ; left valva large , elongate , with a protrusion near basal 1 / 3 on dorsal margin ; ventral margin convex beyond middle , without sclerotized basal process ; patch of modified scales in distal 1 / 6 . right valve slender , narrower beyond 2 / 3 ; patch of modified scales as in left valva . phallus taenioid , terminated with sclerotized spine - like process apically , about 2 / 3 the length of valva .
; meyrick , 1925 : 153 ; caradja & meyrick , 1935 : 69 ; clark , 1969 : 245 ; park , 1995 : 60 ; ueda , 1997 : 79 .
; ponomarenko , 1997 : 52 , 2009 : 341 . tl : india , nilgiris . the holotype in bmnh .
, with the elongated forewing and having a large subtriangular costal patch medially , but can be distinguished by the lack of a long hair pencil on the underside of the forewing . the hindwing is grayish with the anterior expansion developed to beyond middle , and the venation with m 2 close to m 3 basally , m 3 and cua 1 connate and with acute apex . the male genitalia also have quite different characteristics : such as different shapes of valvae , especially the long - stalked , palmately modified scales
a ) : abdominal sternite viii rounded on caudal margin , with long hair pencils laterally . uncus rather short ; socius semiovate . tegumen about as long as valva , slightly expanded laterally beyond middle . left valve broad in basal 3 / 5 , outer margin abruptly truncated , then very narrowed , tapered ; basal process slender , curved , tapered , acute apically ; patch of long - stalked palmately modified scales occupying in distal half . right valve more or less similar to left one , but ventral margin gently rounded , with a very small basal process . phallus slender , tapered , about 2 / 3 the length of left valva .
material examined . vietnam : 1 ♂ , vinh phuc prov . , tam dao nat . park , 750m , 30 vii 2006 ( park , chae , & cuong ) , gen . slide no . cis - 6434 ; 1 ♂ , same locality , 450 m , 15 viii 2006 ( kt park , my kim , & my chae ) .
distribution . india , china ( taiwan : taoyuan co . ) vietnam ( new record ; vinh phuc prov . ) , japan .
remarks . slight morphological differences in male genitalia are found between taiwan ( fig . 42
b ) and japan specimens : vietnam specimen has the ventral margin of valva strongly angled beyond middle and the abdominal sternite viii rounded on caudal margin , not concave at middle . the vietnam specimens are tentatively treated as conspecific in this paper . however , these differences should be re - examined in detail when additional material is available .
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