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 .

additionally , both the monotypic genera verconia and diversidoris are nested within clades .

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

chromodorididae <EOT> 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 )


ardozyga nyctias is a species of moth in the family gelechiidae .

it was described by edward meyrick in 1904 .

it is found in australia , where it has been recorded from southern queensland and new south wales .

the wingspan is about 18 mm .

the forewings are dark fuscous irrorated with ashy-grey and with a conspicuous pale ochreous basal dot in the middle .

the stigmata are moderate , dark fuscous , accompanied by some white scales , with the plical somewhat beyond the first discal .

there are some obscure grey-whitish dots along the posterior half of the costa and termen .

the hindwings are fuscous , darker posteriorly .

the larvae feed on the young shoots of corymbia torelliana , living between leaves joined with silk .

ardozyga nyctias

ardozyga nyctias <EOT> protolechia nyctias meyrick , 1904 ; proc . linn . soc . n . s . w . 29 ( 2 ) : 353 ; tl : brisbane , queensland

living between leaves joined with silk . it grew to a length of 1 cm .

it produced a boring little dark brown moth , with a white spot near the centre of each forewing . the wingspan is about 2 cms .

= protolechia ; sattler , 1973 , bull . br . mus . nat . hist . ( ent . ) 28 ( 4 ) : 172 ; [ nhm card ] ; [ sangmi lee ]

protolechia aclera meyrick , 1904 ; proc . linn . soc . n . s . w . 29 ( 2 ) : 348 ; tl : port lincoln , south australia

protolechia acroleuca meyrick , 1904 ; proc . linn . soc . n . s . w . 29 ( 2 ) : 340 ; tl : bathurst , new south wales

protolechia actinota meyrick , 1904 ; proc . linn . soc . n . s . w . 29 ( 2 ) : 352 ; tl : geraldton and perth , west australia

protolechia aeolopis meyrick , 1904 ; proc . linn . soc . n . s . w . 29 ( 2 ) : 369 ; tl : glen innes ( 4500ft ) , new south wales

protolechia amblopis meyrick , 1904 ; proc . linn . soc . n . s . w . 29 ( 2 ) : 368 ; tl : geraldton , west australia

amphiplaca ( meyrick , 1932 ) ( protolechia ) ; exotic microlep . 4 ( 11 ) : 352

protolechia annularia turner , 1919 ; proc . r . soc . qd 31 ( 10 ) : 143 ; tl : queensland , brisbane

protolechia anthracina meyrick , 1904 ; proc . linn . soc . n . s . w . 29 ( 2 ) : 355 ; tl : sydney , new south wales

protolechia arenaria turner , 1933 ; trans . proc . r . soc . s . austr . 57 : 178 ; tl : queensland , national park ( 3000ft )

protolechia arganthes meyrick , 1904 ; proc . linn . soc . n . s . w . 29 ( 2 ) : 377 ; tl : duaringa , queensland

protolechia argocentra meyrick , 1904 ; proc . linn . soc . n . s . w . 29 ( 2 ) : 350 ; tl : albany , west australia

protolechia aspetodes meyrick , 1904 ; proc . linn . soc . n . s . w . 29 ( 2 ) : 386 ; tl : gisborne , victoria

protolechia autopis meyrick , 1904 ; proc . linn . soc . n . s . w . 29 ( 2 ) : 348 ; tl : perth , west australia

gelechia aversella walker , 1864 ; list spec . lepid . insects colln br . mus . 29 : 649 ; tl : moreton bay

protolechia banausodes meyrick , 1904 ; proc . linn . soc . n . s . w . 29 ( 2 ) : 384 ; tl : rosewood , queensland

protolechia bistrigata meyrick , 1921 ; exotic microlep . 2 ( 14 ) : 427 ; tl : queensland , brisbane

protolechia caminopis meyrick , 1904 ; proc . linn . soc . n . s . w . 29 ( 2 ) : 340 ; tl : bathurst , new south wales

catadamanta ( diakonoff , 1954 ) ( prodosiarcha ) ; verh . k . akad . wet . amst . ( 2 ) 50 ( 1 ) : 11

protolechia catarrhacta meyrick , 1904 ; proc . linn . soc . n . s . w . 29 ( 2 ) : 366 ; tl : sydney , new south wales

protolechia celidophora turner , 1919 ; proc . r . soc . qd 31 ( 10 ) : 145 ; tl : n . queensland , cairns distr . ; eumundi , nr nambour

protolechia cephalota meyrick , 1904 ; proc . linn . soc . n . s . w . 29 ( 2 ) : 373 ; tl : albany , west australia

protolechia chalazodes turner , 1919 ; proc . r . soc . qd 31 ( 10 ) : 140 ; tl : queensland , mt tambourine

protolechia chenias meyrick , 1904 ; proc . linn . soc . n . s . w . 29 ( 2 ) : 359 ; tl : glen innes ( 4500ft ) , new south wales

protolechia chionoprora turner , 1927 ; pap . proc . r . soc . tasm . 1926 : 138

protolechia chiradia meyrick , 1904 ; proc . linn . soc . n . s . w . 29 ( 2 ) : 380 ; tl : rosewood , queensland

protolechia cladara meyrick , 1904 ; proc . linn . soc . n . s . w . 29 ( 2 ) : 346 ; tl : hobart , tasmania

protolechia compsochroa meyrick , 1904 ; proc . linn . soc . n . s . w . 29 ( 2 ) : 370 ; tl : sydney , new south wales ; port lincoln , south australia

protolechia cosmotis meyrick , 1904 ; proc . linn . soc . n . s . w . 29 ( 2 ) : 351 ; tl : perth , west australia

protolechia creperrima turner , 1919 ; proc . r . soc . qd 31 ( 10 ) : 137 ; tl : queensland , brisbane

protolechia crotalodes meyrick , 1904 ; proc . linn . soc . n . s . w . 29 ( 2 ) : 385 ; tl : rosewood , queensland

protolechia crypsibatis meyrick , 1904 ; proc . linn . soc . n . s . w . 29 ( 2 ) : 344 ; tl : mt kosciusco ( 4000ft ) , new south wales ; deloraine , tasmania ; mt lofty , south australia

protolechia crypsicneca turner , 1927 ; pap . proc . r . soc . tasm . 1926 : 138

protolechia cryptosperma meyrick , 1921 ; exotic microlep . 2 ( 14 ) : 428 ; tl : victoria , birchip

gelechia decaspila lower , 1899 ; proc . linn . soc . n . s . w . 24 ( 1 ) : 99 ; tl : stawell , victoria

gelechia deltodes lower , 1896 ; trans . proc . r . soc . s . austr . 20 : 169 ; tl : gisborne , victoria

protolechia diplanetis meyrick , 1904 ; proc . linn . soc . n . s . w . 29 ( 2 ) : 373 ; tl : blackheath ( 3500ft ) , new south wales

baryzancla dysclyta turner , 1933 ; proc . linn . soc . n . s . w . 58 : 81 ; tl : western australia , kalamunda , nr perth

eutorna dysphanes turner , 1947 ; proc . linn . soc . n . s . w . 72 ( 3 - 4 ) : 147 ; tl : queensland , bunya mts

protolechia diplonesa meyrick , 1904 ; proc . linn . soc . n . s . w . 29 ( 2 ) : 344 ; tl : geraldton , west australia

protolechia elassopis turner , 1919 ; proc . r . soc . qd 31 ( 10 ) : 136 ; tl : queensland , caloundra

protolechia elpistis meyrick , 1904 ; proc . linn . soc . n . s . w . 29 ( 2 ) : 349 ; tl : perth , west australia

protolechia emmeles turner , 1933 ; trans . proc . r . soc . s . austr . 57 : 178 ; tl : queensland , national park ( 3000ft )

protolechia enchotypa turner , 1919 ; proc . r . soc . qd 31 ( 10 ) : 138 ; tl : victoria , gisborne

protolechia englypta meyrick , 1904 ; proc . linn . soc . n . s . w . 29 ( 2 ) : 357 ; tl : gisborne , victoria

cleodora eumela lower , 1897 ; trans . r . soc . s . aust . 21 : 59 ; tl : stawell , victoria

protolechia euprepta turner , 1933 ; trans . proc . r . soc . s . austr . 57 : 177 ; tl : queensland , national park ( 3000ft )

protolechia euryarga turner , 1919 ; proc . r . soc . qd 31 ( 10 ) : 135 ; tl : queensland , brisbane

protolechia eustephana turner , 1919 ; proc . r . soc . qd 31 ( 10 ) : 146 ; tl : queensland , brisbane

protolechia exarista meyrick , 1904 ; proc . linn . soc . n . s . w . 29 ( 2 ) : 339 ; tl : york , west australia

protolechia flexilis meyrick , 1904 ; proc . linn . soc . n . s . w . 29 ( 2 ) : 345 ; tl : sydney , new south wales

protolechia frugalis meyrick , 1904 ; proc . linn . soc . n . s . w . 29 ( 2 ) : 345 ; tl : sydney and blackheath ( 3500ft ) , new south wales ; perth , west australia

protolechia furcifera turner , 1919 ; proc . r . soc . qd 31 ( 10 ) : 139 ; tl : victoria , gisborne

prodosiarcha glagera turner , 1919 ; proc . r . soc . qd 31 ( 10 ) : 129 ; tl : n . queensland , cairns

protolechia gorgonias meyrick , 1904 ; proc . linn . soc . n . s . w . 29 ( 2 ) : 380 ; tl : rosewood , queensland

protolechia gypsocrana turner , 1919 ; proc . r . soc . qd 31 ( 10 ) : 143 ; tl : queensland , brisbane ; toowoomba ; stradbroke i .

protolechia hedana turner , 1919 ; proc . r . soc . qd 31 ( 10 ) : 145 ; tl : queensland , brisbane

protolechia hilara turner , 1919 ; proc . r . soc . qd 31 ( 10 ) : 135 ; tl : victoria , gisborne

protolechia hormodes meyrick , 1904 ; proc . linn . soc . n . s . w . 29 ( 2 ) : 363 ; tl : sydney , new south wales

protolechia hylias meyrick , 1904 ; proc . linn . soc . n . s . w . 29 ( 2 ) : 369 ; tl : mt kosciusko ( 5000ft ) , new south wales

protolechia hypocneca turner , 1919 ; proc . r . soc . qd 31 ( 10 ) : 137 ; tl : queensland , warwick

protolechia hypoleuca meyrick , 1904 ; proc . linn . soc . n . s . w . 29 ( 2 ) : 351 ; tl : sydney , new south wales

semocharista idiospila meyrick , 1922 ; ark . zool . 14 ( 15 ) : 4 ; tl : nw . australia , kimberley district

protolechia invalida meyrick , 1904 ; proc . linn . soc . n . s . w . 29 ( 2 ) : 346 ; tl : brisbane , queensland ; sydney , new south wales

protolechia involuta turner , 1919 ; proc . r . soc . qd 31 ( 10 ) : 142 ; tl : n . queensland , townsville

protolechia iochlaema [ = iochlaena ] meyrick , 1904 ; proc . linn . soc . n . s . w . 29 ( 2 ) : 353 ; tl : perth , west australia

baryzancla ithygramma turner , 1933 ; proc . linn . soc . n . s . w . 58 : 81 ; tl : western australia , mt dale

protolechia liota meyrick , 1904 ; proc . linn . soc . n . s . w . 29 ( 2 ) : 350 ; tl : albany , west australia

gelechia lithina lower , 1899 ; proc . linn . soc . n . s . w . 24 ( 1 ) : 98 ; tl : broken hill , new south wales

protolechia loemias meyrick , 1904 ; proc . linn . soc . n . s . w . 29 ( 2 ) : 383 ; tl : murrurundi , new south wales ; gisborne , victoria

prodosiarcha loxodesma meyrick , 1904 ; proc . linn . soc . n . s . w . 29 ( 2 ) : 330 ; tl : adelaide , south australia

protolechia mechanistis meyrick , 1904 ; proc . linn . soc . n . s . w . 29 ( 2 ) : 360 ; tl : evandale and hobart , tasmania

protolechia megalommata meyrick , 1904 ; proc . linn . soc . n . s . w . 29 ( 2 ) : 384 ; tl : rosewood , queensland

protolechia megalosticta turner , 1919 ; proc . r . soc . qd 31 ( 10 ) : 141 ; tl : new south wales , sydney

protolechia melicrata turner , 1919 ; proc . r . soc . qd 31 ( 10 ) : 137 ; tl : n . queensland , cairns ; lucinda point , nr ingham

new south wales , s . australia , victoria , tasmania . see [ maps ]

gelechia mesochra lower , 1894 ; trans . proc . r . soc . s . aust . 18 : 107 ; tl : parkside , south australia

protolechia mesopsamma turner , 1919 ; proc . r . soc . qd 31 ( 10 ) : 144 ; tl : queensland , rosewood

protolechia microdora meyrick , 1904 ; proc . linn . soc . n . s . w . 29 ( 2 ) : 346 ; tl : geraldton , west australia

protolechia molyntis meyrick , 1904 ; proc . linn . soc . n . s . w . 29 ( 2 ) : 385 ; tl : gisborne , victoria ; mt lofty , south australia

protolechia nephelota meyrick , 1904 ; proc . linn . soc . n . s . w . 29 ( 2 ) : 354 ; tl : mt kosciusko ( 3000ft ) , new south wales ; gisborne , victoria

protolechia neurosticha turner , 1933 ; trans . proc . r . soc . s . austr . 57 : 177 ; tl : queensland , brisbane

protolechia nothrodes meyrick , 1921 ; exotic microlep . 2 ( 14 ) : 429 ; tl : queensland , brisbane

protolechia obscura turner , 1933 ; trans . proc . r . soc . s . austr . 57 : 177 ; tl : queensland , national park ( 2 - 2500ft )

protolechia ochrobathra turner , 1933 ; trans . proc . r . soc . s . austr . 57 : 177 ; tl : queensland , bunya mtns ( 3500ft )

protolechia odorifera meyrick , 1904 ; proc . linn . soc . n . s . w . 29 ( 2 ) : 371 ; tl : sydney and shoalhaven , new south wales

gelechia orthanotos lower , 1900 ; proc . linn . soc . n . s . w . 25 ( 1 ) : 50 ; tl : stawell , victoria

protolechia penthicodes meyrick , 1921 ; exotic microlep . 2 ( 14 ) : 427 ; tl : queensland , brisbane

protolechia phasianis meyrick , 1904 ; proc . linn . soc . n . s . w . 29 ( 2 ) : 382 ; tl : mt kosciusko ( 4700ft ) , new south wales ; melbourne and gisborne , victoria ; launceston , tasmania

protolechia plinthactis meyrick , 1904 ; proc . linn . soc . n . s . w . 29 ( 2 ) : 363 ; tl : sydney , new south wales

brachyzancla poenicea turner , 1947 ; proc . linn . soc . n . s . w . 72 ( 3 - 4 ) : 154 ; tl : queensland , bunya

protolechia polioxysta turner , 1933 ; trans . proc . r . soc . s . austr . 57 : 177 ; tl : queensland , springbrook ( 3000ft ) , macpherson range

protolechia prisca meyrick , 1904 ; proc . linn . soc . n . s . w . 29 ( 2 ) : 343 ; tl : sydney , west australia

protolechia proscripta meyrick , 1921 ; exotic microlep . 2 ( 14 ) : 429 ; tl : queensland , brisbane

protolechia psephias meyrick , 1904 ; proc . linn . soc . n . s . w . 29 ( 2 ) : 352 ; tl : geraldton , west australia

protolechia pyrrhica turner , 1919 ; proc . r . soc . qd 31 ( 10 ) : 141 ; tl : queensland , coolangatta

protolechia sarisias meyrick , 1904 ; proc . linn . soc . n . s . w . 29 ( 2 ) : 356 ; tl : brisbane , queensland

protolechia sciodes meyrick , 1904 ; proc . linn . soc . n . s . w . 29 ( 2 ) : 347 ; tl : geraldton , west australia

protolechia scytina meyrick , 1904 ; proc . linn . soc . n . s . w . 29 ( 2 ) : 374 ; tl : bathurst , new south wales ; port lincoln , south australia

protolechia secta meyrick , 1921 ; exotic microlep . 2 ( 14 ) : 428 ; tl : queensland , brisbane

protolechia semiographa turner , 1919 ; proc . r . soc . qd 31 ( 10 ) : 146 ; tl : queensland , mt tambourine

protolechia sisyraea meyrick , 1904 ; proc . linn . soc . n . s . w . 29 ( 2 ) : 359 ; tl : gisborne , victoria

protolechia sporodeta turner , 1919 ; proc . r . soc . qd 31 ( 10 ) : 144 ; tl : queensland , killarney

protolechia tabulata meyrick , 1904 ; proc . linn . soc . n . s . w . 29 ( 2 ) : 367 ; tl : sydney , new south wales ; port lincoln , south australia

protolechia taracta turner , 1919 ; proc . r . soc . qd 31 ( 10 ) : 143 ; tl : queensland , montville ( 1500ft ) , nr nambour ; mt tambourine ; toowoomba

protolechia telopis meyrick , 1904 ; proc . linn . soc . n . s . w . 29 ( 2 ) : 338 ; tl : sydney , new south wales

protolechia temenitis meyrick , 1904 ; proc . linn . soc . n . s . w . 29 ( 2 ) : 341 ; tl : rosewood , queensland

protolechia tetraploa meyrick , 1904 ; proc . linn . soc . n . s . w . 29 ( 2 ) : 338 ; tl : melbourne , victoria

gelechia thanatodes lower , 1893 ; trans . proc . r . soc . s . austr . 17 ( 1 ) : 170 ; tl : parkside

protolechia thyridota meyrick , 1904 ; proc . linn . soc . n . s . w . 29 ( 2 ) : 355 ; tl : glen innes ( 4500ft ) , new south wales ; york , west australia

protolechia thyrsoptera meyrick , 1904 ; proc . linn . soc . n . s . w . 29 ( 2 ) : 381 ; tl : rosewood , queensland

protolechia trachyphanes meyrick , 1904 ; proc . linn . soc . n . s . w . 29 ( 2 ) : 343 ; tl : geraldton , west australia

protolechia trichalina meyrick , 1904 ; proc . linn . soc . n . s . w . 29 ( 2 ) : 377 ; tl : geraldton and perth , west australia

protolechia trichosema meyrick , 1904 ; proc . linn . soc . n . s . w . 29 ( 2 ) : 347 ; tl : albany , west australia

protolechia trichroma turner , 1933 ; trans . proc . r . soc . s . austr . 57 : 176 ; tl : queensland , national park ( 3000ft )

gelechia tridecta lower , 1900 ; proc . linn . soc . n . s . w . 25 ( 1 ) : 48 ; tl : parkside , south australia

protolechia trochias meyrick , 1921 ; exotic microlep . 2 ( 14 ) : 428 ; tl : queensland , cairns

protolechia tyroessa turner , 1933 ; trans . proc . r . soc . s . austr . 57 : 176 ; tl : ; queensland , national park ( 3500ft )

gelechia vacatella walker , 1864 ; list spec . lepid . insects colln br . mus . 30 : 1026 ; tl : australia ?

protolechia voluta meyrick , 1904 ; proc . linn . soc . n . s . w . 29 ( 2 ) : 341 ; tl : newcastel and sydney , new south wales

protolechia xanthocephala meyrick , 1904 ; proc . linn . soc . n . s . w . 29 ( 2 ) : 376 ; tl : brisnabe , queensland ; sydney and blackheath ( 3500ft ) , new south wales

protolechia xestolitha meyrick , 1904 ; proc . linn . soc . n . s . w . 29 ( 2 ) : 354 ; tl : sydney , new south wales

protolechia xuthias meyrick , 1904 ; proc . linn . soc . n . s . w . 29 ( 2 ) : 361 ; tl : wirrabara , south australia

brachyzancla sporima turner , 1947 ; proc . linn . soc . n . s . w . 72 ( 3 - 4 ) : 154 ; tl : queensland , stanthorpe

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 of new guinea , results of the third archbold expedition ( american - netherlands indian expedition 1938 - 1939 ) . part iv

results of dr . e . mjöberg ' s swedish scientific expedition to australia 1910 - 1913 . microlepidoptera

walker , 1864 list of the specimens of lepidopterous insects in the collection of the british museum list spec . lepid . insects colln br . mus . 27 : 1 - 286 ( 1863 ) , 28 : 287 - 562 ( 1863 ) , 29 : 563 - 835 ( 1864 ) , 30 : 837 - 1096 ( 1864 )

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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the kerguelen tern ( sterna virgata ) is a tern of the southern hemisphere .

this seabird mainly breeds colonially in the kerguelen islands , as its common name implies .

however , smaller colonies are also found in the prince edward islands ( i.e. prince edward and marion ) and crozet islands .

the total number of individuals is from 3,500 – 6,500 birds , although there is no recent data from the main colony at kerguelen .

these birds do not inhabit kerguelen proper , instead nesting on islets free of feral cats .

during bad weather , they are known to abandon their colonies .

kerguelen terns are among the least-ranging of all typical terns .

they generally do not reach far into the seas near their breeding grounds .

these birds eat fish and marine invertebrates , especially those found in beds of the seaweed macrocystis spp .

they sometimes also hunt insects on land and catch fish from rivers on kerguelen .

there are two subspecies : s. v. mercuri ( voisin , 1971 ) – crozet and prince edward islands s. v. virgata ( cabanis , 1875 ) – kerguelen island

kerguelen tern

kerguelen tern <EOT> is sister to elegant tern , not sandwich tern of europe ( efe et al . 2009 , collinson et al 2017 ) ;

aspects of the breeding and feeding of kerguelen and antarctic terns at the kerguelen islands by p . m . sagar

the kerguelen tern is sedentary and remains all year round in vicinity of the breeding islands . the flight is fast and graceful .

calls and songs : sounds by xeno - canto the kerguelen tern utters high - pitched screams . while foraging , it produces churrs and squeaks .

introduction : the kerguelen tern occurs only on four islands in the indian ocean , and has very restricted range . this species is one of the rarest terns in the world . very similar to the antarctic tern ( s . vittata ) , which is present in the same range , the kerguelen tern has darker plumage and the species can be differentiated in the field by the greyish underwing ( white in s . vittata ) . the kerguelen tern defends aggressively it nest , including eggs and / or chicks , against larger seabirds and even humans .

cabanis , 1875 – kerguelen is ( reported breeding on heard i erroneous ) .

the introduction of salmonid fish into rivers on kerguelen has provided a new source of food .

kerguelen tern and antarctic tern are probably competing on their breeding islands , and coexistence of these closely related species has involved a parallel reduction of clutch size and the displacement of the laying period . competition for the limited food resources is reduced and allows both species to feed the chicks . ( weimerskirch & stahl – 1988 )

protection / threats / status : the kerguelen tern has reduced range and small population . the main threat probably comes from hard weather conditions and strong winds that reduce the food sources and the breeding success . predation by skuas still occurs on the islands , but exotic predators have been eradicated from the islands , except a large population of feral cats on kerguelen . the global population is estimated at 3 , 500 / 6 , 500 individuals , equivalent to 2 , 300 / 4 , 300 mature individuals . this population appears to be stable . but currently , the kerguelen tern is classified as near threatened

the kerguelen tern is colonial nester and breeds two months before the antarctic tern . the colonies include up to 30 pairs , but some of them breed solitary . the nests are widely scattered , about 35 - 38 metres apart . the displays are similar to that of s . vittata , and include aerial displays and courtship feeding by male to female . they are generally monogamous .

subspecies and range : the kerguelen tern has two subspecies . s . v . mercuri occurs on prince edward and marion islands , and on crozet islands . this race has more restricted grey area on forehead in non - breeder birds , and some subtle differences on the bare parts . s . v . virgata ( described above ) is found on kerguelen islands .

habitat : the kerguelen tern is not exclusively marine , and also frequents inland freshwaters . it can be found near ponds , marshes and beaches of all islands . it breeds on flat ground with sparse vegetation on rocky or volcanic islands , on cliff tops , near rivers and sometimes close to beach . the non - breeding birds feed in the vicinity of the islands .

the kerguelen tern adult has smoky - grey upperparts , including the uppertail - coverts , whereas the rump is white . the tail is greyish with grey outer streamers . the underparts are smoky - grey too , but the undertail - coverts are mostly white . the underwing is grey with narrow , black trailing edge on the primaries . on the head , forehead , crown and nape are black . cheeks and moustachial area are white and contrast strongly with the black above , and the grey of chin and throat below . the bill is deep red . the eyes are dark brown . legs and webbed feet are bright orange ( crozet ) and dull red ( kerguelen ) .

gochfeld , m . , burger , j . & garcia , e . f . j . ( 2018 ) . kerguelen tern ( sterna virgata ) . in : del hoyo , j . , elliott , a . , sargatal , j . , christie , d . a . & de juana , e . ( eds . ) . handbook of the birds of the world alive . lynx edicions , barcelona . ( retrieved from urltoken on 9 july 2018 ) .

31 cm . very dark , small tern . adult overall very dark grey with black cap and narrow white cheek stripe separating black cap from grey neck . non breeding adult has grizzled forehead and paler underparts . immature heavily barred buff on mantle and has all brown cap .

behaviour in the wild : the kerguelen tern feeds on both aquatic and terrestrial invertebrates , according to the season . at sea , it feeds on fish , crustaceans and molluscs caught by plunge - diving and contact - dipping . they forage in flocks of up to 20 birds on inshore waters or by walking on kelp . on land , it feeds on earthworms , insects and spiders found in the vegetation , taken both on the ground and in flight . flocks may contain up to 50 individuals . they can be aggressive towards each other while feeding , displacing another by flying towards it while churring .

1988 , thibault and guyst 1993 ) . the south african population is listed in the regional red data book as critically endangered [ taylor and wanless 2015 ] , due to the apparent loss of the prince edward island population . there are no recent counts from the main breeding area on kerguelen ( v . bretagnolle

the non - breeding adult has dull reddish bill ( crozet ) or black ( kerguelen ) . the forehead becomes white by february while the bird is moulting during the incubation . the juvenile is finely vermiculated grey , brown and buff , with broad pale and dark bars . it is similar to the juvenile s . vittata .

2009 ] and no breeding observed in 2011 [ taylor and wanless , 2015 ] ) , crozet islands ( french southern territories ) ( 150 - 200 pairs over 1980 - 1982 ) and kerguelen islands ( also french southern territories ) ( 1 , 000 - 2 , 000 pairs during 1982 - 1985 ) . the total population is estimated at 3 , 500 - 6 , 500 individuals ( jouventin

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del hoyo , j . , collar , n . j . , christie , d . a . , elliott , a . and fishpool , l . d . c . 2014 . hbw and birdlife international illustrated checklist of the birds of the world . volume 1 : non - passerines . lynx edicions birdlife international , barcelona , spain and cambridge , uk .

from which it differs by being darker , smaller with shirt wings and tail and with much shorter , weaker and spikey shaped bill .

anderson , o . , butchart , s . , moreno , r . , o ' brien , a . , shutes , s . , stattersfield , a . , taylor , j . , van der merwe , n .

this species is listed as near threatened owing to its small population . it is not thought to be undergoing a decline at present , but any indication of a decline would result in the species being reclassified as threatened .

1999 ) and thus population trends are unknown , but it is assumed that the species is not undergoing any significant decline .

the total population has been estimated at 3 , 500 - 6 , 500 individuals , roughly equivalent to 2 , 300 - 4 , 300 mature individuals .

the population is suspected to be stable in the absence of any significant threats .

. egg - laying commences in mid - october , and continues until january , with a peak in breeding from early november to mid - december . one or two eggs are laid . the incubation period is 24 days , followed by a fledging period of 31 - 39 days and then 20 days of dependence ( del hoyo

inhabits predator - free islets around the main island and therefore predation is not considered a major threat ( t . micol

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ioc _ names _ file _ plus - 8 . 2g : 8 . 2

and terns are embedded among the gulls in a monophyletic clade ; separation of the families rynchopidae and sternidae would render the gull family laridae paraphyletic relative to these other two groups ( baker et al . 2008 )

red sea , indian ocean east through the pacific to hawaiian is . and easter i .

clipperton i . ( off w mexico ) and cocos i . ( off w costa rica )

marcus ( japan ) and n marshall is . ( micronesia ) to nw hawaiian is .

henderson , easter , and sala y gómez is . ( west of chile )

may be a basal noddy ( baker et al . 2007 , ödeen et al . 2010 , cibois et al . 2016 ) ; so " white noddy " if confirmed ? see yeung et al . 2009 re taxonomy of pacific ocean subspecies

seychelles and mascarene is . to the central pacific ( except ranges of next two ssp . )

eu , af : iberian pen . through the middle east to pakistan and india ; mauritania , senegal and gambia

n and ne australia , new caledonia , loyalty is . ( sw pacific )

red - billed gull originally split by schodde et al . 1983 , sibley & monroe 1990 , burger & gochfeld 1996 , and given et al . 2005 , but now lumped by h & m4 , hbw alive , gill et al . 2010 .

by some ( h & m4 , hbw / bli ) based on pons et al . 2005 poorly resolved phylogeny

genetically distinct short - billed gull previously split from common / mew gull ( aou sibley 1996 , zink et al 1995 , olsen & larsson 2003 , bna ) . bna makes the point that zink’s paper only looked at mtdna differences between

is needed to round out the picture and because some have proposed ( e . g . s & m , 1990 ) that the species break might be between

) has long been a controversial taxon . critical review favors treatment as subspecies of iceland gull

( sangster et al . 2005 ) ; recognized by bou , but not by aou

as ssp following collinson et al . 2008 , yésou ( dutch birding 24 : 271 - 298 , 2002 )

( sangster et al . 2005 , aou 2007 ) ; recognized by bou , aou . this species retains the english name yellow - legged gull of the former , unsplit species . .

as subspecies following collinson , j . m . , parkin , d . t . , knox , a . g . , sangster , g . , and l . svensson . 2008 . species boundaries in the herring and lesser black - backed gull complex . british birds 101 ( 7 ) : 340 - 363 .

e china to w , n and e australia , ryukyu and tuamotu is .

w and n australia and new guinea through lord howe , norfolk and kermadec is and the s pacific to easter i .

bonin is . ( japan ) to the hawaiian is . south to kiritimati ( line is . )

sa : coastal peru and se brazil to tierra del fuego , falkland is .

stewart , snares , auckland , bounty , antipodes , campbell and macquarie is .

( aou 2000 , bou / sangster et al . , 2004 , chu et al . 2009 )

frans josef land ( russia ) to st . lawrence i . ( alaska )

na , ma : channel is . ( california , usa ) and islands off w baja california ( mexico )

scripps ' s murrelet is split from guadalupe [ xantus ' s ] murrelet ( sibley and monroe 1993 , nettleship 1996 , birt et al . 2012 , nacc 2011 - 14 - c )

sandgrouse ( pterocliformes ) are sister to the malagasy mesites ( hackett et al . 2008 , jarvis et al . 2014 , prum et al . 2015 ) .

sympatric with s . vittata , but different phenology and feeding niche . race mercuri separated principally on basis of more restricted grey area on forehead ( non - breeders ) and bare - part differences , but distinctions subtle ; species often treated as monotypic . two subspecies recognized .

33 cm ; wingspan 71 cm . forehead to nape black ; upperparts , including uppertail - coverts , smoky grey , with contrasting white rump ; outer tail streamers grey , tail greyish ; . . .

calls include a harsh " chittick " and a long drawn out " keeeaaar " .

rocky , volcanic islands . breeds on sparsely vegetated flat ground , either cliff tops or river flats . . .

fish , crustaceans , molluscs , earthworms , insects and spiders . diet changes seasonally from mainly crustaceans in sept , fish in nov , insects . . .

virtually unstudied . returns to colony in sept , and begins laying mid - oct , continuing until jan ; peak breeding early nov to mid - dec , 2 . . .

not globally threatened . currently considered near threatened , and previously , vulnerable . the global population has been estimated at 3 , 500–6 , 500 individuals . . . .

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sometimes treated as including one or more of the genera onychoprion , sternula , gelochelidon , hydroprogne , chlidonias and thalasseus .

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this map displays aggregated data from ibc and my birding ( anonymously ) ; markers do not indicate precise localities .

recommended citation birdlife international ( 2018 ) species factsheet : sterna virgata . downloaded from urltoken on 09 / 07 / 2018 . recommended citation for factsheets for more than one species : birdlife international ( 2018 ) iucn red list for birds . downloaded from urltoken on 09 / 07 / 2018 .

current time at palmer station , antarctica : current temperature : cold . damn cold .

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howard and moore 4th edition ( incl . corrigenda vol . 1 - 2 ) :

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reproduction of this species : the laying occurs between mid - october and january , with peak in early november to mid - december . the nest is placed on the ground , often on slopes above the beach or near the high tide line , occasionally well inland . it is a scrape made with stones and twigs , and lined with plant material such as moss and grass .

the female lays 1 - 2 eggs and both adults incubate during 24 days . the chicks are brooded for five days . then , they leave the nest but they hide in vegetation and rocks near the nest . they fledge 31 - 39 days after hatching , but they still depend on adults for food for about 3 weeks .

a complete guide to antarctic wildlife by hadoram shirihai and illustrated by brett jarrett - edited by guy m . kirwan - alul . a press oy , finland - isbn 9519894705

sonogram images © xeno - canto foundation . sonogram images share the same license terms as the recording they depict .

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english - german online dictionary developed to help you share your knowledge with others . more information ! contains translations by tu chemnitz and mr honey ' s business dictionary ( german - english ) . thanks on that account ! links to this dictionary or to single translations are very welcome ! questions and answers