the lime-speck pug ( eupithecia centaureata ) is a moth of the family geometridae .

it is a common species throughout the palearctic region ( where it is found in europe , central asia , mongolia , southern siberia , eastern china ( guangdong ) and taiwan ) , the near east and north africa .

lime - speck pug

lime - speck pug ( eupithecia centaureata ) - norfolk moths - the macro and micro moths of norfolk .
riley , a . m . , prior , g . ( 2003 ) british and irish pug moths . 1 - 145 : 57
riley , a . m . , prior , g . ( 2003 ) british and irish pug moths . 1 - 145 : 98
riley , a . m . , prior , g . ( 2003 ) british and irish pug moths . 1 - 145 : 105
riley , a . m . , prior , g . ( 2003 ) british and irish pug moths . 1 - 145 : 122
riley , a . m . , prior , g . ( 2003 ) british and irish pug moths . 1 - 145 : 120
riley , a . m . , prior , g . ( 2003 ) british and irish pug moths . 1 - 145 : 59
riley , a . m . , prior , g . ( 2003 ) british and irish pug moths . 1 - 145 : 114
riley , a . m . , prior , g . ( 2003 ) british and irish pug moths . 1 - 145 : 83
riley , a . m . , prior , g . ( 2003 ) british and irish pug moths . 1 - 145 : 49
riley , a . m . , prior , g . ( 2003 ) british and irish pug moths . 1 - 145 : 126
riley , a . m . , prior , g . ( 2003 ) british and irish pug moths . 1 - 145 : 142
riley , a . m . , prior , g . ( 2003 ) british and irish pug moths . 1 - 145 : 79
riley , a . m . , prior , g . ( 2003 ) british and irish pug moths . 1 - 145 : 76
riley , a . m . , prior , g . ( 2003 ) british and irish pug moths . 1 - 145 : 62
riley , a . m . , prior , g . ( 2003 ) british and irish pug moths . 1 - 145 : 96
riley , a . m . , prior , g . ( 2003 ) british and irish pug moths . 1 - 145 : 102
riley , a . m . , prior , g . ( 2003 ) british and irish pug moths . 1 - 145 : 39
riley , a . m . , prior , g . ( 2003 ) british and irish pug moths . 1 - 145 : 101
riley , a . m . , prior , g . ( 2003 ) british and irish pug moths . 1 - 145 : 52
riley , a . m . , prior , g . ( 2003 ) british and irish pug moths . 1 - 145 : 37
riley , a . m . , prior , g . ( 2003 ) british and irish pug moths . 1 - 145 : 81
riley , a . m . , prior , g . ( 2003 ) british and irish pug moths . 1 - 145 : 53
riley , a . m . , prior , g . ( 2003 ) british and irish pug moths . 1 - 145 : 43
riley , a . m . , prior , g . ( 2003 ) british and irish pug moths . 1 - 145 : 143
riley , a . m . , prior , g . ( 2003 ) british and irish pug moths . 1 - 145 : 129
riley , a . m . , prior , g . ( 2003 ) british and irish pug moths . 1 - 145 : 45
riley , a . m . , prior , g . ( 2003 ) british and irish pug moths . 1 - 145 : 88
riley , a . m . , prior , g . ( 2003 ) british and irish pug moths . 1 - 145 , 86
riley , a . m . , prior , g . ( 2003 ) british and irish pug moths . 1 - 145 : 124
riley , a . m . , prior , g . ( 2003 ) british and irish pug moths . 1 - 145 : 117
riley , a . m . , prior , g . ( 2003 ) british and irish pug moths . 1 - 145 : 131
riley , a . m . , prior , g . ( 2003 ) british and irish pug moths . 1 - 145 : 64
riley , a . m . , prior , g . ( 2003 ) british and irish pug moths . 1 - 145 : 132
riley , a . m . , prior , g . ( 2003 ) british and irish pug moths . 1 - 145 : 55
riley , a . m . , prior , g . ( 2003 ) british and irish pug moths . 1 - 145 : 141
riley , a . m . , prior , g . ( 2003 ) british and irish pug moths . 1 - 145 : 138
riley , a . m . , prior , g . ( 2003 ) british and irish pug moths . 1 - 145 : 91
riley , a . m . , prior , g . ( 2003 ) british and irish pug moths . 1 - 145 : 67
riley , a . m . , prior , g . ( 2003 ) british and irish pug moths . 1 - 145 : 75
riley , a . m . , prior , g . ( 2003 ) british and irish pug moths . 1 - 145 : 50
riley , a . m . , prior , g . ( 2003 ) british and irish pug moths . 1 - 145 : 41
riley , a . m . , prior , g . ( 2003 ) british and irish pug moths . 1 - 145 : 72
riley , a . m . , prior , g . ( 2003 ) british and irish pug moths . 1 - 145 : 110
riley , a . m . , prior , g . ( 2003 ) british and irish pug moths . 1 - 145 : 137
riley , a . m . , prior , g . ( 2003 ) british and irish pug moths . 1 - 145 : 47
riley , a . m . , prior , g . ( 2003 ) british and irish pug moths . 1 - 145 : 103
riley , a . m . , prior , g . ( 2003 ) british and irish pug moths . 1 - 145 : 112
riley , a . m . , prior , g . ( 2003 ) british and irish pug moths . 1 - 145 : 127
riley , a . m . , prior , g . ( 2003 ) british and irish pug moths . 1 - 145 : 106
riley , a . m . , prior , g . ( 2003 ) british and irish pug moths . 1 - 145 : 94
riley , a . m . , prior , g . ( 2003 ) british and irish pug moths . 1 - 145 : 99
riley , a . m . , prior , g . ( 2003 ) british and irish pug moths . 1 - 145 : 119
riley , a . m . , prior , g . ( 2003 ) british and irish pug moths . 1 - 145 , 33
riley , a . m . , prior , g . ( 2003 ) british and irish pug moths . 1 - 145 : 135
a distinctive and slender - winged pug , it is thought that this species may resemble a bird - dropping and thus reduce attraction to predators .
wingspan 16 - 20 mm . a distinctive and slender - winged pug , it is thought that this species may resemble a bird - dropping and thus reduce attraction to predators .
a taxon identifier is composed of name , author , year and attribute , all separated by a blank . these are all extracted from the original publication .
the name is reproduced exactly as proposed in the original publication . the name of a genus is made up of one word and species made up of two words ( genus and species ) separated by a blank .
the author ' s name is made up of a string of letters , with no blanks , and multiple authors ' names are separated by a comma . spelling of author ' s name is based on the original publication . if there are more than three authors , only the names of the first two authors are shown , followed by
, +
and the number of omitted authors .
attribute is enclosed in square brackets . this is rarely needed , but to differentiate homo - identifiers , this will contain the page , line or plate number of original publication .
all diacritic marks , hyphens , and apostrophes are eliminated , thus only the following characters are used : a to z , a to z , 0 to 9 , blank , comma , and opening and closing square brackets . although upper and lower cases are used for the convenience of human recognition , it is not case sensitive .
created by dicky sick ki yu 1997 - 2012 please send me information about errors and omissions ( contact information ) with supporting references , possibly with pdf or hard copy .
fairly common throughout britain , occurring in a wide range of habitats , the species has one or two broods , being found on the wing from april to september .
the caterpillars feed on the flowers of a range of low - growing plants .
ukmoths is built , run and maintained by ian kimber , with thanks to the many kind contributors who provide photos and information .
the ukmoths facebook page is a great place to post your identification queries . more often than not you ' ll get a positive id on most photos fairly quickly .
looking for a specific moth species ? enter just part of the name below .
procache : v317 render date : 2018 - 07 - 07 06 : 05 : 08 page render time : 0 . 3618s total w / procache : 0 . 4299s
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recorded in 66 ( 96 % ) of 69 10k squares . first recorded in 1834 . last recorded in 2018 .
underlying maps using digital map data © norfolk online lepidoptera archive - nola™ 2018 . © james wheeler - n o r f o l k m o t h s 2007 - 2018 . data © nola™ 2018
click here to support naturespot by making a donation - small or large - your gift is very much appreciated . thank you .
the species has one or two broods , being found on the wing from april to september .
fairly common throughout britain . in a recent survey to determine the status of all macro moths in britain this species was classified as common .
fairly common in leicestershire and rutland . l & r moth group status = a ( common and resident )
the large dark splodge on each forewing together with smaller black round - bractet is highly indicative of this moth . when the insect is resting it could easily be mistaken for a bird dropping . has many habitats in open land . is not fussy and can be found on a wide range of herbaceous plants . 20 - 24mm across when opened .
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
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for the county , we have a total of 5098 records from 377 sites . first recorded in 1880 .
: sparsely recorded from all five vice - counties . most frequent in the south , although not seen near selby ( vc61 / 64 ) for many years ( smj pers . comm . ) .
: doing well in the county particularly in lowland areas . two overlapping broods , the second being larger .
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notes : common in gardens , hedgerows , scrub and saltmarshes , and on roadside verges throughout the british isles . in hampshire and on the isle of wight widespread and common . wingspan 20 - 24 mm . unmistakable . larva feeds on such plants as ragwort , mugwort , sea wormwood , burnet - saxifrage , knapweed , scabious , hemp - agrimony , yarrow , traveller ' s - joy , goldenrod and wild angelica , over - wintering as a pupa .
description : wingspan 20 - 24 mm . adults resemble a bird dropping when at rest . forewings greyish white with a dark brown blotch that merges with the black discal spot ; the size of the blotch can vary between individuals . there are also darker blotches running along the dorsum of the forewing . hindwings pale white with a small black discal spot and chequered fringe .
flight period : end of may to late august . skinner gives from summer to early autumn as the normal flight period in britain .
status : common and fairly widespread in eastern counties with the majority of records coming from well - recorded sites along the down coastline . it has also been recorded from several inland sites in north armagh at loughgall , aughinlig and navan and also from rathlin island , antrim .
ecology : a polyphagous species found in a variety of habitats in small numbers . adults come frequently to light and will occasionally visit flowers . the larvae feed during june and july on the flowers of mouse - ears cerastium spp . it overwinters as a pupa .
world distribution : eurasiatic ; common throughout europe as far east as asia and also north africa .
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weed of the week burdock description : . . . - thunderbird outdoor adventures | facebook
these pages show a variety of moth caterpillars that can be found in typical habitats . this is unlike the organisation of adult moths as they tend to have a recognised and restricted flight period , often only a couple of months . caterpillars , on the other hand , are around much longer . the larval stage of many species lasts maybe ten months , with part - grown larvae hibernating over the winter . there are caterpillars you would be almost guaranteed to find on heather moorland but would never see in your garden , and vice versa . other categories include coast , scrub , carr & hedgerows , woodland , roadsides & waste ground . some species overlap habitats , inevitably . a selection of the most likely species to be found in each habitat are shown .
it should be emphasised just how hard it is to find most larvae , and then how difficult it is to identify many without rearing them . comments alongside many photographs provide guidance on identification and confusion species ( many of which make then indistiguishable ) and also the likelyhood of finding them and when they might be found .
butterfly conservation is a registered charity and non - profit - making company , limited by guarantee . registered office : manor yard , east lulworth , wareham , dorset , bh20 5qp . charity registered in england and wales ( 254937 ) and in scotland ( sco39268 ) .
taken at musselburgh on 13th august 2017 using panasonic lumix lx5 in macro mode .
a variety of flowers including yarrow , burdock , heather . nectar rich flowers . all the wings being largely white except for a black blotch on the costa of the forewing . 20 - 24 mm . woodland . summer - autumn .
the larvae are found in various colours with diagonal patches down the sides finishing on top of the larva . yellow form - with brownish red markings . green form - dark green markings . pinkish form - deep pink markings .
the moth has a white to whitish brown narrow forewing with a black discal spot located on a light or darker black patch adjacent to the costa , which makes this moth unmistakeable .
if disturbed they tend to hide beneath a leaf . the moth is attracted to light and is a regular visitor to the light trap .
website where further information like photos , physical characteristics , habitats , edible uses , medicinal uses , cultivation , propagation , range , height etc . are clearly listed .
plant families - in bold red below takes the visitor to the relevant
lepi - plants
page where other butterflies & moths using the plants below are listed .
a sub - family of more than 150 species . typically feeding on the foliage of a wide range of dicot plants , although many species consume the host ' s flowers and seeds . particular genera display affinities for certain hosts , e . g . scotopteryx on leguminous herbs and shrubs ( fabaceae ) ; catarhoe on bedstraws ( rubiaceae ) ; chloroclysta on shrubs and trees ; and thera on conifers .
old data currently unassessed by a specialist for changes in nomenclature , new species to the british list , new interactions , errors in original sources , and misspellings . new material added from waring , townsend & lewington , 2003 . field guide to the moths of great britain and ireland . british wildlife publishing , uk . ; ;
below is a list of all sources for this family / sub - family contained within dbif . click on source name for a list of all records for this source .
brit . ent . and nat . hist . soc . ( 1981 ) an identification guide to the british pugs . lepidoptera geometridae 1 - 42 : 26
waring , p . townsend , m ( 2003 ) field guide to the moths of great britain and ireland . 1 - 432 , 130
waring , p . townsend , m ( 2003 ) field guide to the moths of great britain and ireland . 1 - 432 , 122
brit . ent . and nat . hist . soc . ( 1981 ) an identification guide to the british pugs . lepidoptera geometridae 1 - 42 : 15
barbour , d . a . ( 1985 ) entomologist ' s rec . j . var . 97 146 - 146 : 146
noble , k . ( 1975 ) working draft of
the natural foodplants of macrolepidoptera larvae in britain , part ii .
field studies council : 25
brit . ent . and nat . hist . soc . ( 1981 ) an identification guide to the british pugs . lepidoptera geometridae 1 - 42 : 20
costen , p . d . m . & peet , t . n . d . ( 1986 ) entomologist ' s rec . j . var . 0098 0217 - 0218 : 0217
waring , p . townsend , m ( 2003 ) field guide to the moths of great britain and ireland . 1 - 432 , 103
waring , p . townsend , m ( 2003 ) field guide to the moths of great britain and ireland . 1 - 432 , 88
haggett , g . m . ( 1980 ) entomologist ' s gaz . 31 221 - 226 :
riley , a . m . ( 1986 ) entomologist ' s rec . j . var . 0098 0207 - 00208 : 0208
corley , m . f . v . ( 1984 ) entomologist ' s gaz . 35 : 2 76 - 77 : 77
porter , j . ( 1997 ) the colour identification guide to caterpillars of the british isles . 1 - 154 , 34
waring , p . townsend , m ( 2003 ) field guide to the moths of great britain and ireland . 1 - 432 , 128
porter , j . ( 1997 ) the colour identification guide to caterpillars of the british isles . 1 - 154 , 40
waring , p . townsend , m ( 2003 ) field guide to the moths of great britain and ireland . 1 - 432 , 98
waring , p . townsend , m ( 2003 ) field guide to the moths of great britain and ireland . 1 - 432 , 129
waring , p . townsend , m ( 2003 ) field guide to the moths of great britain and ireland . 1 - 432 , 116
noble , k . ( 1975 ) working draft of
the natural foodplants of macrolepidoptera larvae in britain , part ii .
field studies council : 41
noble , k . ( 1975 ) working draft of
the natural foodplants of macrolepidoptera larvae in britain , part ii .
field studies council : 24
waring , p . townsend , m ( 2003 ) field guide to the moths of great britain and ireland . 1 - 432 , 92
waring , p . townsend , m ( 2003 ) field guide to the moths of great britain and ireland . 1 - 432 , 119
porter , j . ( 1997 ) the colour identification guide to caterpillars of the british isles . 1 - 154 , 41
waring , p . townsend , m ( 2003 ) field guide to the moths of great britain and ireland . 1 - 432 , 82
noble , k . ( 1975 ) working draft of
the natural foodplants of macrolepidoptera larvae in britain , part ii .
field studies council : 26
waring , p . townsend , m ( 2003 ) field guide to the moths of great britain and ireland . 1 - 432 , 117
arnold , v . w . ( 1979 ) entomologist ' s rec . j . var . 91 : 11 289 - 332 : 322
waring , p . townsend , m ( 2003 ) field guide to the moths of great britain and ireland . 1 - 432 , 115
brit . ent . and nat . hist . soc . ( 1981 ) an identification guide to the british pugs . lepidoptera geometridae 1 - 42 : 34
brit . ent . and nat . hist . soc . ( 1981 ) an identification guide to the british pugs . lepidoptera geometridae 1 - 42 : 16
waring , p . townsend , m ( 2003 ) field guide to the moths of great britain and ireland . 1 - 432 , 126
waring , p . townsend , m ( 2003 ) field guide to the moths of great britain and ireland . 1 - 432 , 76
waring , p . townsend , m ( 2003 ) field guide to the moths of great britain and ireland . 1 - 432 , 95
noble , k . ( 1975 ) working draft of
the natural foodplants of macrolepidoptera larvae in britain , part ii .
field studies council : 40
brit . ent . and nat . hist . soc . ( 1981 ) an identification guide to the british pugs . lepidoptera geometridae 1 - 42 : 18
noble , k . ( 1975 ) working draft of
the natural foodplants of macrolepidoptera larvae in britain , part ii .
field studies council : 32
owen , d . f . ( 1982 ) entomologist ' s rec . j . var . 94 52 - 52 : 52
waring , p . townsend , m ( 2003 ) field guide to the moths of great britain and ireland . 1 - 432 , 107
noble , k . ( 1975 ) working draft of
the natural foodplants of macrolepidoptera larvae in britain , part ii .
field studies council : 33
brit . ent . and nat . hist . soc . ( 1981 ) an identification guide to the british pugs . lepidoptera geometridae 1 - 42 : 31
waring , p . townsend , m ( 2003 ) field guide to the moths of great britain and ireland . 1 - 432 , 108
brit . ent . and nat . hist . soc . ( 1981 ) an identification guide to the british pugs . lepidoptera geometridae 1 - 42 : 10
owen , d . f . ( 1987 ) entomologist ' s gaz . 38 209 - 213 : 210
waring , p . townsend , m ( 2003 ) field guide to the moths of great britain and ireland . 1 - 432 , 124
waring , p . townsend , m ( 2003 ) field guide to the moths of great britain and ireland . 1 - 432 , 85
noble , k . ( 1975 ) working draft of
the natural foodplants of macrolepidoptera larvae in britain , part ii .
field studies council : 23
porter , j . ( 1997 ) the colour identification guide to caterpillars of the british isles . 1 - 154 , 33
brit . ent . and nat . hist . soc . ( 1981 ) an identification guide to the british pugs . lepidoptera geometridae 1 - 42 : 24
owen , d . f . ( 1987 ) entomologist ' s gaz . 38 209 - 213 : 212
waring , p . townsend , m ( 2003 ) field guide to the moths of great britain and ireland . 1 - 432 , 96
waring , p . townsend , m ( 2003 ) field guide to the moths of great britain and ireland . 1 - 432 , 123
waring , p . townsend , m ( 2003 ) field guide to the moths of great britain and ireland . 1 - 432 , 83
waring , p . townsend , m ( 2003 ) field guide to the moths of great britain and ireland . 1 - 432 , 120
knill - jones , r . p . ( 1982 ) entomologist ' s rec . j . var . 94 77 - 77 : 77
noble , k . ( 1975 ) working draft of
the natural foodplants of macrolepidoptera larvae in britain , part ii .
field studies council : 42
noble , k . ( 1975 ) working draft of
the natural foodplants of macrolepidoptera larvae in britain , part ii .
field studies council : 38
waring , p . townsend , m ( 2003 ) field guide to the moths of great britain and ireland . 1 - 432 , 89
waring , p . townsend , m ( 2003 ) field guide to the moths of great britain and ireland . 1 - 432 , 110
waring , p . townsend , m ( 2003 ) field guide to the moths of great britain and ireland . 1 - 432 , 121
brit . ent . and nat . hist . soc . ( 1981 ) an identification guide to the british pugs . lepidoptera geometridae 1 - 42 : 33
waring , p . townsend , m ( 2003 ) field guide to the moths of great britain and ireland . 1 - 432 , 102
waring , p . townsend , m ( 2003 ) field guide to the moths of great britain and ireland . 1 - 432 , 75
waring , p . townsend , m ( 2003 ) field guide to the moths of great britain and ireland . 1 - 432 , 117
waring , p . townsend , m ( 2003 ) field guide to the moths of great britain and ireland . 1 - 432 , 111
waring , p . townsend , m ( 2003 ) field guide to the moths of great britain and ireland . 1 - 432 , 100
brit . ent . and nat . hist . soc . ( 1981 ) an identification guide to the british pugs . lepidoptera geometridae 1 - 42 : 27
brit . ent . and nat . hist . soc . ( 1981 ) an identification guide to the british pugs . lepidoptera geometridae 1 - 42 : 13
waring , p . townsend , m ( 2003 ) field guide to the moths of great britain and ireland . 1 - 432 , 90
porter , j . ( 1997 ) the colour identification guide to caterpillars of the british isles . 1 - 154 , 39
waring , p . townsend , m ( 2003 ) field guide to the moths of great britain and ireland . 1 - 432 , 78
brit . ent . and nat . hist . soc . ( 1981 ) an identification guide to the british pugs . lepidoptera geometridae 1 - 42 : 21
brit . ent . and nat . hist . soc . ( 1981 ) an identification guide to the british pugs . lepidoptera geometridae 1 - 42 : 30
porter , j . ( 1997 ) the colour identification guide to caterpillars of the british isles . 1 - 154 , 37
waring , p . townsend , m ( 2003 ) field guide to the moths of great britain and ireland . 1 - 432 , 127
waring , p . townsend , m ( 2003 ) field guide to the moths of great britain and ireland . 1 - 432 , 104
noble , k . ( 1975 ) working draft of
the natural foodplants of macrolepidoptera larvae in britain , part ii .
field studies council : 29
waring , p . townsend , m ( 2003 ) field guide to the moths of great britain and ireland . 1 - 432 , 112
brit . ent . and nat . hist . soc . ( 1981 ) an identification guide to the british pugs . lepidoptera geometridae 1 - 42 : 14
waring , p . townsend , m ( 2003 ) field guide to the moths of great britain and ireland . 1 - 432 , 84
waring , p . townsend , m ( 2003 ) field guide to the moths of great britain and ireland . 1 - 432 , 91
waring , p . townsend , m ( 2003 ) field guide to the moths of great britain and ireland . 1 - 432 , 72
owen , d . f . ( 1983 ) entomologist ' s rec . j . var . 95 20 - 20 : 20
noble , k . ( 1975 ) working draft of
the natural foodplants of macrolepidoptera larvae in britain , part ii .
field studies council : 37
noble , k . ( 1975 ) working draft of
the natural foodplants of macrolepidoptera larvae in britain , part ii .
field studies council : 36
waring , p . townsend , m ( 2003 ) field guide to the moths of great britain and ireland . 1 - 432 , 87
porter , j . ( 1997 ) the colour identification guide to caterpillars of the british isles . 1 - 154 , 36
waring , p . townsend , m ( 2003 ) field guide to the moths of great britain and ireland . 1 - 432 , 116
brit . ent . and nat . hist . soc . ( 1981 ) an identification guide to the british pugs . lepidoptera geometridae 1 - 42 : 19
waring , p . townsend , m ( 2003 ) field guide to the moths of great britain and ireland . 1 - 432 , 101
brit . ent . and nat . hist . soc . ( 1981 ) an identification guide to the british pugs . lepidoptera geometridae 1 - 42 : 22
riley , a . m . ( 1986 ) entomologist ' s mon . mag . 37 : 2 68 - 68 : 68
michaelis , h . n . ( 1981 ) entomologist ' s rec . j . var . 93 40 - 40 : 40
noble , k . ( 1975 ) working draft of
the natural foodplants of macrolepidoptera larvae in britain , part ii .
field studies council : 35
waring , p . townsend , m ( 2003 ) field guide to the moths of great britain and ireland . 1 - 432 , 118
waring , p . townsend , m ( 2003 ) field guide to the moths of great britain and ireland . 1 - 432 , 79
noble , k . ( 1975 ) working draft of
the natural foodplants of macrolepidoptera larvae in britain , part ii .
field studies council : 30
noble , k . ( 1975 ) working draft of
the natural foodplants of macrolepidoptera larvae in britain , part ii .
field studies council : 31
waring , p . townsend , m ( 2003 ) field guide to the moths of great britain and ireland . 1 - 432 , 97
waring , p . townsend , m ( 2003 ) field guide to the moths of great britain and ireland . 1 - 432 , 73
brit . ent . and nat . hist . soc . ( 1981 ) an identification guide to the british pugs . lepidoptera geometridae 1 - 42 : 12
brit . ent . and nat . hist . soc . ( 1981 ) an identification guide to the british pugs . lepidoptera geometridae 1 - 42 : 29
viart , m . ( ed . ) ( 1979 ) poplars and willows in wood production and land use . fao forestry series 10 1 - 328 : 230
waring , p . townsend , m ( 2003 ) field guide to the moths of great britain and ireland . 1 - 432 , 81
waring , p . townsend , m ( 2003 ) field guide to the moths of great britain and ireland . 1 - 432 , 114
riley , a . m . ( 1985 ) entomologist ' s gaz . 36 : 4 259 - 261 : 260
brit . ent . and nat . hist . soc . ( 1981 ) an identification guide to the british pugs . lepidoptera geometridae 1 - 42 : 11
west , b . k . ( 1981 ) entomologist ' s rec . j . var . 93 198 - 198 : 198
porter , j . ( 1997 ) the colour identification guide to caterpillars of the british isles . 1 - 154 , 38
waring , p . townsend , m ( 2003 ) field guide to the moths of great britain and ireland . 1 - 432 , 113
haggett , g . m . ( 1981 ) entomologist ' s rec . j . var . 93 236 - 236 : 236
sterling , m . ( 1985 ) entomologist ' s rec . j . var . 97 27 - 28 : 27
waring , p . townsend , m ( 2003 ) field guide to the moths of great britain and ireland . 1 - 432 , 77
waring , p . townsend , m ( 2003 ) field guide to the moths of great britain and ireland . 1 - 432 , 80
noble , k . ( 1975 ) working draft of
the natural foodplants of macrolepidoptera larvae in britain , part ii .
field studies council : 34
noble , k . ( 1975 ) working draft of
the natural foodplants of macrolepidoptera larvae in britain , part ii .
field studies council : 28
agassiz , d . & skinner , b . ( 1980 ) entomologist ' s rec . j . var . 92 162 - 166 : 164
noble , k . ( 1975 ) working draft of
the natural foodplants of macrolepidoptera larvae in britain , part ii .
field studies council : 39
brit . ent . and nat . hist . soc . ( 1981 ) an identification guide to the british pugs . lepidoptera geometridae 1 - 42 : 17
waring , p . townsend , m ( 2003 ) field guide to the moths of great britain and ireland . 1 - 432 , 109
waring , p . townsend , m ( 2003 ) field guide to the moths of great britain and ireland . 1 - 432 , 93
brit . ent . and nat . hist . soc . ( 1981 ) an identification guide to the british pugs . lepidoptera geometridae 1 - 42 : 28
waring , p . townsend , m ( 2003 ) field guide to the moths of great britain and ireland . 1 - 432 , 106
noble , k . ( 1975 ) working draft of
the natural foodplants of macrolepidoptera larvae in britain , part ii .
field studies council : 27
waring , p . townsend , m ( 2003 ) field guide to the moths of great britain and ireland . 1 - 432 , 125
riley , a . m . ( 1986 ) entomologist ' s rec . j . var . 98 85 - 89 : 85
brit . ent . and nat . hist . soc . ( 1981 ) an identification guide to the british pugs . lepidoptera geometridae 1 - 42 : 23
waring , p . townsend , m ( 2003 ) field guide to the moths of great britain and ireland . 1 - 432 , 74
waring , p . townsend , m ( 2003 ) field guide to the moths of great britain and ireland . 1 - 432 , 86
noble , k . ( 1975 ) working draft of
the natural foodplants of macrolepidoptera larvae in britain , part ii .
field studies council :
porter , j . ( 1997 ) the colour identification guide to caterpillars of the british isles . 1 - 154 , 35
waring , p . townsend , m ( 2003 ) field guide to the moths of great britain and ireland . 1 - 432 , 99
skinner , b . ( 1985 ) entomologist ' s rec . j . var . 97 185 - 186 : 186
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we supply yarrow in our selected wildflower trays or seed mixes but we also sell it here on its own , as seed , 55cc plug plants , or in more generous half litre pots .
supplier : british wildflower plants / all things rural our yarrow plants and seeds have guaranteed uk provenance and are supplied by signatories to the flora locale code of practice .
welcome to our online shop , where you will find top quality plants , trees and seeds , delivered straight to you from our community of specialist growers and harvesters around britain . we donate half of our profits from sales through the shop to uk conservation charities .
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butterfly conservation ' s rough bank reserve is a site known by some local butterfly enthusiasts for many years , but until 2012 ( the year when bc purchased rough bank ) there had been few reports of moths found there apart from daytime observations on a few occasions . one such record in 1993 was of a larva of narrow - bordered bee hawk - moth found feeding on devil ' s - bit scabious . five nationally scarce micro - moth species had also been recorded at the site in the past .
since 2012 a total of 628 moth species have been recorded on the site ( plus 4 other species only recorded in earlier years ) . three of these were the first records for the species in gloucestershire : trifurcula headleyella and coleophora therinella in 2012 , and aphelia viburnana in 2014 .
3 nationally rare species have been found at rough bank ( species given provisional red data book status in the 2012 review of the status of microlepidoptera in britain ) , and 43 nationally scarce species : 3 in the
notable a
category , 40
notable b
. for more information about these species see the rough bank rare and scarce moths page .
this page lists the moth species recorded at the site , mostly on the unimproved limestone grassland slopes of rough bank sssi but also any additional species recorded in the two pasture fields which were included in bc ' s purchase . these fields both have wetland areas with plants associated with such habitat and supporting colonies of some rush - feeding moth species .
for each species the table below lists the agassiz , beavan & heckford checklist number , the bradley list number , scientific and common names , national status , earliest and most recent years recorded and typical uk larval food . the list will be updated at least annually using records sent to the current vc33 ( east gloucestershire ) moth recorder , robert homan . it is updated more frequently with my own records from the site , and sometimes with additional species reported to me . records prior to 2012 are from the former gloucestershire moth recorder roger gaunt ' s database .
following the publication in september 2015 of a paper showing that delplanqueia inscriptella occurs in the uk and d . dilutella seems to be very restricted in its range , all previous gloucestershire records of d . dilutella are doubtful . from reexamination of a few kept specimens ( all looking too dark for d . dilutella ) , d . inscriptella has been found at several gloucestershire sites including rough bank .
the species name links in the table below are to the species entries on the uk moths website . please note that the gloucestershire moth distribution maps are not updated frequently , so some species listed below may not yet be shown as recorded in the rough bank tetrad so90e . also some informal reports of species from rough bank are listed below , but such records will only be mapped if the recorder sends them to the vc33 ( east gloucestershire ) moth recorder - for contact details see the distribution maps page .
perforate & other st . john ' s - worts , bird ' s - foot trefoil
bird ' s - foot trefoil ( f . palustrella ) ; greater b - f . tref . ( f . decreta )
page last revised june 2018 butterfly conservation . company limited by guarantee , registered in england ( 2206468 ) . registered office : manor yard , east lulworth , wareham , dorset , bh20 5qp . charity registered in england and wales ( 254937 ) and in scotland ( sco39268 ) . vat no gb 991 2771 89
upto four 125wmv traps two 6w and a 15w actinic trap were placed over a wide area of the bog on four separate dates in may , june , july and august , usually covering the bog myrtle / heather communities and the birch / pine habitats . a further daytime search was made in october for leaf mines . the uk status is common
unless stated , although some of the micro - moths ( especially the leaf mines , nepticulidae & gracillariidae ) usually comes under the ‘unknown’ category as very little is known and studied with this neglected group . all larval food plants are in brackets . the numbers on the left hand side are the bradley & fletcher numbers .
1313 catoptria pinella ( grasses & sedges esp . tufted hair - grass ) ( local )
2130 dotted clay ( various trees & plants inc . heather and bog myrtle )
2196 striped wainscot ( common reed & various grasses inc . purple moor - grass ) ( local )
2286 light knot grass ( various plants inc . heather and bog myrtle ) ( local )
2305 small angle shades ( various trees & plants inc . bracken and birch )
2412 silver hook ( sedges & grasses inc . tufted - hair grass ) ( local )
2484 pinion - streaked snout ( unknown , but probably inc . heather’s and rushes ) ( local )
2493 dotted fan - foot ( various marsh grasses inc . wood - rush & wood - sedge ) ( notable n / b )
if we were to split the food plants into three main categories which are associated with dersingham bog , heathland , birch and pines then this gives more of a clue as to what species is using what habitat .
striped wainscot ( grasses inc . purple - moor grass / common reed ) local
bog myrtle would be widely used by many species under the larval food plant heading of ‘various plants’ ( as would heather ) . this would include the local pinion - streaked snout .
mention must be made of the light knot grass trapped here in may and june . in norfolk it has been recorded from roydon common for some years now and this was the first record away from this site . its nearest occurrence to norfolk is the staffordshire moorlands .
other food plants associated with the wetter areas of dersingham bog such as phragmites , juncus etc , deserve special mention too .
a summary of the above gives over 45 local species associated with the bog , one notable na and three notable nb .
is a brightly - coloured geometer moth that is usually seen more in daylight , lightly flying over the wetter areas of the heathland , it will also come to light . the food plant is marsh cinquefoil (
is only a tiny noctuid 15 - 18 mm long , very easily overlooked and may be mistaken for a microlepidopteron . upto 100 were noted around the bog myrtle during the july visit , with fewer in the august visit . the larval food plant is not known but orhant ( 1977 ) considers it likely to be juncus spp .
usually found in reed - beds and fens in east anglia and the larva feed on sallow and various marsh sedges and grasses . it seems to becoming more widespread over the last few years .


the euteleostei or euteleosts is a clade of bony fishes within the teleostei that evolved some 240 million years ago .

it is divided into the protacanthopterygii ( including the salmon and dragonfish ) and the neoteleostei ( including the lanternfish , lizardfish , oarfish , and the acanthopterygii ) .

the cladogram is based on near et al. ( 2012 ) and betancur-rodriguez et al. 2016 .

they explored the phylogeny and divergence times of every major lineage , analysing the dna sequences of 9 unlinked genes .

they calibrated ( set actual values for ) branching times in this tree from 36 reliable measurements of absolute time from the fossil record .

euteleostei

( 27 ) euteleostei . mrca : lepidogalaxias , takifugu . hard minimum age : 149 ma . 95 % soft maximum age : 260 ma . prior setting : uniform distribution ( crown calibration ) . calibration source : broughton et al . [ 2 ] .
davis mp . evolutionary relationships of the aulopiformes ( euteleostei : cyclosquamata ) : a molecular and total evidence approach . in : nelson js , schultze hp , wilson mvh , editors . origin and phylogenetic interrelationships of teleosts . 2010 . münchen , germany : verlag dr . friedrich pfeil . pp . 317 - 336 .
with regard to the relationships of early euteleosts , our phylogenetic analyses support several results from previous molecular studies and a new result that places galaxiidae as the sister lineage of neoteleostei ( without stomiiforms ) [ bs = 95 % , bpp = 1 . 00 ( fig . 1 , and figs . s1 and s2 ) ] . lineages previously treated as “protacanthopterygians” ( 3 ) are polyphyletic in the molecular phylogeny because the alepocephaliforms ( slickheads ) are resolved in a clade containing clupeomorphs ( anchovies and herrings ) and ostariophysians ( catfish and minnows ) [ bs = 94 % , bpp = 1 . 00 ( 21 , 33 ) ] , the enigmatic freshwater australian species lepidogalaxias salamandroides is the sister lineage to all other euteleostei ( 15 , 23 ) [ bs = 100 % , bpp = 1 . 00 ( fig . 1 , and figs . s1 and s2 ) ] , salmonids ( trouts and salmon ) and esociforms ( pikes and mudminnows ) are resolved as a clade [ bs = 100 % , bpp = 1 . 00 ( 21 , 23 ) ] , and there is strong support for a clade containing stomiiforms ( dragonfishes ) , osmeriforms ( smelts ) , and retropinnids ( southern smelts ) [ bs = 100 % , bpp = 1 . 00 ( 23 ) ] . although most of these relationships were reflected in the species tree , lepidogalaxias was resolved as the sister lineage of galaxiidae ( fig . s3 ) . however , only one of the two gene trees ( rag1 ) that sampled both lepidogalaxias and galaxiidae resolved these lineages as sharing a common ancestor . the phylogenetic resolution of these early euteleost lineages using morphology is thought to have been hampered by a mosaic of highly modified and ancestral character states ( 3 , 13 ) . the relationships inferred in our trees provide a phylogenetic framework to investigate the evolution of morphological character state changes , which have proven difficult to use in the inference of relationships among early diverging euteleost lineages ( e . g . , ref . 34 ) .
edited by david m . hillis , university of texas , austin , tx , and approved july 19 , 2012 ( received for review april 22 , 2012 )
ray - finned fishes make up half of all living vertebrate species . nearly all ray - finned fishes are teleosts , which include most commercially important fish species , several model organisms for genomics and developmental biology , and the dominant component of marine and freshwater vertebrate faunas . despite the economic and scientific importance of ray - finned fishes , the lack of a single comprehensive phylogeny with corresponding divergence - time estimates has limited our understanding of the evolution and diversification of this radiation . our analyses , which use multiple nuclear gene sequences in conjunction with 36 fossil age constraints , result in a well - supported phylogeny of all major ray - finned fish lineages and molecular age estimates that are generally consistent with the fossil record . this phylogeny informs three long - standing problems : specifically identifying elopomorphs ( eels and tarpons ) as the sister lineage of all other teleosts , providing a unique hypothesis on the radiation of early euteleosts , and offering a promising strategy for resolution of the “bush at the top of the tree” that includes percomorphs and other spiny - finned teleosts . contrasting our divergence time estimates with studies using a single nuclear gene or whole mitochondrial genomes , we find that the former underestimates ages of the oldest ray - finned fish divergences , but the latter dramatically overestimates ages for derived teleost lineages . our time - calibrated phylogeny reveals that much of the diversification leading to extant groups of teleosts occurred between the late mesozoic and early cenozoic , identifying this period as the “second age of fishes . ”
ray - finned fishes ( actinopterygii ) are one of the most successful radiations in the long evolutionary history of vertebrates , yet despite the rapid progress toward reconstructing the vertebrate tree of life , only 5 % of the ray - finned fish phylogeny is resolved with strong support ( 1 ) . actinopterygii contains more than 30 , 000 species ( 2 ) , with all but 50 being teleosts ( 3 ) . compared with other large vertebrate radiations , such as mammals ( 4 ) or birds ( 5 ) , a general consensus on the phylogenetic relationships and timing of diversification among the major actinopterygian and teleost lineages is lacking ( 3 , 6 , 7 ) . this uncertainty about relationships has prevented the development of a comprehensive time - calibrated phylogeny of ray - finned fishes , which is necessary to understand macroevolutionary processes that underlie their diversity .
) , and unlike other clades of vertebrates , there has been no comprehensive effort to resolve the phylogeny of actinopterygians and teleosts using molecular data that sample multiple nuclear genes and include taxa that span the major lineages . despite the long history of using morphological data in the phylogenetics of ray - finned fishes , there are several areas of uncertainty and disagreement regarding some of the most fundamental relationships . first , there are two competing hypotheses on the phylogenetic relationships that reflect the earliest diversification of teleosts : either the osteoglossomorpha [ bony tongues (
) ] are the sister lineage of all other teleosts . second , the relationships of lower euteleosts ( e . g . , salmons , smelts , pikes , slickheads , and galaxiids ) , or “protacanthopterygians , ” has changed frequently as a result of phylogenetic analyses of different morphological datasets (
) to label the percomorpha as the “bush at the top of the [ teleost ] tree . ”
applications of molecular data to these three long - standing questions in teleost phylogenetics have yielded mixed results . for example , analyses of nuclear and mtdna gene sequences have supported all three possible relationships among osteoglossomorphs , elopomorphs , and all other teleosts [ i . e . , clupeocephalans (
we investigated phylogenetic relationships and divergence times of all major lineages of actinopterygii and teleostei using dna sequences of nine unlinked protein - coding nuclear genes sampled from 232 species . we used 36 well - justified absolute time calibrations from the fossil record of ray - finned fishes in relaxed - molecular clock analyses to estimate divergence times . phylogenies resulting from these analyses were well resolved , the majority of phylogenetic inferences were supported with strong node support values , were robust to inferences using new “species tree” methods , and provide a comprehensive molecular perspective on areas of long - standing disagreement and uncertainty in the relationships of teleost fishes . divergence times estimated from relaxed - molecular clock analyses yield a comprehensive time - scale of actinopterygian diversification that is remarkably close to ages inferred from the fossil record .
maximum - likelihood analyses of the nine nuclear gene dataset resolved 89 % of the 232 nodes in the actinopterygian phylogeny with bootstrap replicate scores ( bs ) ≥70 % and the phylogenies inferred using the bayesian method had 91 % of the nodes strongly supported posterior probabilities ( bpp ) ≥ 0 . 95 ( fig . 1 , and figs . s1 and s2 ) . relationships of nonteleostean actinopterygians were consistent with traditional morphologically - based inferences ( 6 ) with polypterids ( bichirs and ropefish ) resolved as the sister lineage of all other actinopterygians ( actinopteri ) in the relaxed - clock analysis ( fig . 1 ) . in addition , acipenseriformes ( sturgeons and paddlefishes ) were the sister lineage of neopterygii with strong support ( bs = 100 % , bpp = 1 . 00 ) , and holostei ( bowfin and gars ) was resolved as the sister lineage of teleosts [ bs = 100 % , bpp = 1 . 00 ( fig . 1 , and figs . s1 and s2 ) ] . these results contrast with earlier molecular studies that either resolved acipenseriforms and holosteans as an “ancient - fish” clade ( 31 ) or acipenseriforms and polypteriforms as a weakly supported clade ( 32 ) .
actinopterygian time - calibrated phylogeny based on nine nuclear genes and 36 fossil age constraints . bars represent the posterior distribution of divergence - time estimates . gray bars identify nodes supported with bpp ≥ 0 . 95 , and white bars mark nodes with bpp < 0 . 95 . nodes with age priors taken from the fossil record are marked with a “c . ” for full details on calibration see materials and methods and fig . s2 . the time - calibrated tree is scaled to the geological time scale with absolute time given in millions of years .
our results provide resolution to three of the most compelling questions in teleost phylogenetics . the molecular phylogeny resulted in the strongly supported position ( bs = 97 % , bpp = 1 . 00 ) of elopomorphs as the sister lineage of all other teleosts ( fig . 1 , and figs . s1 and s2 ) . this result is also strongly supported in a species tree analysis , which accounts for potential discordance among individual gene histories , with a bootstrap proportion of 100 % ( fig . s3 ) . evidence for osteoglossomorpha as the sister lineage of all other teleosts was based on the presence of a single character state in the caudal fin skeleton ( 9 , 10 ) . on the other hand , the hypothesis that elopomorpha is the sister lineage of all other teleosts was based on eight derived character - state changes identified from optimization of a matrix containing 135 discretely coded morphological characters ( 11 ) . our results strongly support the latter hypothesis , illustrating agreement between phylogenetic inferences from a robust morphological data matrix and our densely sampled nuclear gene dna sequence dataset .
one of the most important problems in vertebrate phylogenetics is the resolution of the major lineages of percomorpha . the phylogeny confirms several results presented in previous molecular analyses , including the resolution of ophidiiforms ( cusk eels ) and batrachoidids ( toadfish ) as early diverging percomorphs (
posterior distribution of molecular age estimates and patterns of calibration sharing across studies of ray - finned fish phylogeny . ( a ) posterior distribution of molecular age estimates , in millions of years , for 14 actinopterygian lineages , resulting from analyses of whole mtdna genomes ( blue ) , the rag1 nuclear gene ( orange ) , the rag1 nuclear gene using the calibrations from this study ( yellow ) , and the nine nuclear gene dataset presented in this study ( green ) . the circle represents the mean of the posterior estimate and the whiskers mark the upper and lower 95 % highest posterior density of the age estimates . gray boxes mark the oldest fossils for a given lineage , those with dashed lines were used as calibration age priors ( see materials and methods ) and those with solid black lines were not used as age calibrations . line drawings of ray - finned fish species are based on photographs of specimens housed at the peabody museum of natural history , yale university , new haven , ct . ( b ) frequency of calibrations shared between this study and those using whole mtdna genomes ( blue ) and the rag1 nuclear gene ( orange ) binned by the age of the fossil calibration in millions of years ( ma ) .
despite the apparent gap in the fossil record for early crown–group teleosts , we find that most major teleost lineages originated in a period spanning the late mesozoic into the early cenozoic ( figs . 1 and 2 a ) , which corresponds to patterns apparent in the fossil record ( 39 ) . we identify this interval as the “second age of fishes . ” the devonian age of fishes is characterized by the presence of all major vertebrate lineages referred to as “fishes , ” both living and extinct [ e . g . , ostracoderms , placoderms , acanthodians , chondrichthyans , and so forth ( 40 ) ] . although this period in time appears to mark the origin of crown actinopterygii ( figs . 1 and 2 a ) , it does not correspond to the divergence of the major lineages that comprise the bulk of living actinopterygian biodiversity . instead , the second age of fishes represents the interval in geologic time where these species - rich lineages ( e . g . , otophysians and acanthomorphs ) originated and eventually flourished , becoming the dominant vertebrate component of marine and freshwater habitats .
ray - finned fishes include half of the entire species richness of vertebrates ( 2 , 3 ) , but had ranked last , by a wide margin , in the degree of phylogenetic resolution offered by available dna sequence and genomic resources ( 1 ) . our phylogeny , based on a multilocus dataset , provides robust resolution and strong support across all major lineages of ray - finned fishes and teleosts . additionally , our divergence time estimates reconcile inferences from paleontology with those obtained from other studies that used molecular methods , providing a molecular time scale that is more consistent with ages implied by the fossil record . this comprehensive molecular perspective on the evolutionary diversification of one - half of all vertebrate species provides dna sequence data and calibration information from which to integrate resolution of clades at lower taxonomic levels ( e . g . , families ) and estimate ages of actinopterygian lineages that lack a fossil record .
standard phenol - chloroform extraction protocol or qiagen dneasy blood and tissue kits were used to isolate dna from tissue biopsies sampled from 232 ray - finned fish species ( table s2 ) . previously published pcr primers were used to amplify and sequence an exon from each of nine nuclear genes [ glyt , myh6 , plagl2 , ptr , rag1 , sh3px3 , sreb2 , tbr1 , and zic1 ( 22 , 41 ) ] . the genes were aligned by eye using the inferred amino acid sequences . no frame mutations or dna substitutions that resulted in stop codons were observed in the aligned dna sequences . the combined nine - gene dataset contained 7 , 587 base pairs .
twenty - seven data partitions were designated that corresponded to the three separate codon positions for each of the nine genes . a gtr + g substitution model was used in a portioned maximum - likelihood analysis using the computer program raxml 7 . 2 . 6 ( 42 ) run with the –d option . support for nodes in the raxml tree was assessed with a thorough bootstrap analysis ( option –f i ) with 1 , 000 replicates .
a species tree was inferred using gene tree parsimony implemented in the computer program igtp ( 43 ) . individual gene trees estimated using raxml were used as input files . several rooting strategies were used . the individual gene trees were rooted using erpetoichthys calabaricus or polypterus ornatipinnis , except in three cases when these species were not sampled for a specific gene . in these cases the individual gene trees were rooted using scaphirhynchus platorynchus , amia calva , or atractosteus spatula . a heuristic search using randomized hill climbing was performed to find the species tree that minimized the reconciliation cost for deep coalescence . this search was bootstrapped by performing it 100 times and bootstrap proportions for the resulting species trees were calculated using sumtrees in the dendropy package ( 44 ) .
for each fossil calibration prior , we identify the calibrated node in the ray - fin fish phylogeny , list the taxa that represent the first occurrence of the lineage in the fossil record , describe the character states that justify the phylogenetic placement of the fossil taxon , provide information on the stratigraphy of the rock formations bearing the fossil , give the absolute age estimate for the fossil , outline the prior age setting in the beast relaxed - clock analysis , and provide any additional notes on the calibration ( si text ) . each calibration is numbered and the phylogenetic placement of the calibration is highlighted in fig . s2 .
we thank t . - y . cheng and k . - t . shao of the biodiversity research museum , academia sinica ; j . friel of the cornell university museum of vertebrates ; p . a . hastings and h . j . walker of the scripps institution of oceanography ; k . p . maslenikov and t . w . pietsch of the burke museum of natural history and culture , university of washington ; and a . c . bentley and e . o . wiley of the biodiversity institute of the university of kansas for generous gifts of tissue specimens . j . s . albert , j . w . armbruster , l . bernatchez , t . m . berra , c . p . burridge , c . d . hulsey , s . lavoué , j . g . lundberg , m . miya , n . merret , p . j . unmack , k . watanabe , j . m . waters provided additional specimens ; c . m . bossu , r . c . harrington , p . r . hollingsworth , c . d . hulsey , b . p . keck , and the staff of the carribean research management of biodiversity biological research station in curaçao provided assistance in sampling expeditions ; gregory watkins - colwell assisted with museum collections . k . l . ilves provided insight on taxon sampling . this research was supported by the peabody museum of natural history and national science foundation grants deb - 0444842 , deb - 0716155 , deb - 0717009 , deb - 0732642 , ant - 0839007 , deb - 1060869 , deb - 1061806 , and deb - 1061981 ( to w . l . s . , p . c . w . , and t . j . n . ) ; and natural environment research council grant nerc ne / i005536 / 1 ( to m . f . ) .
author contributions : t . j . n . , r . i . e . , a . d . , j . a . m . , m . p . d . , p . c . w . , m . f . , and w . l . s . designed research ; t . j . n . , r . i . e . , a . d . , k . l . k . , p . c . w . , m . f . , and w . l . s . performed research ; t . j . n . , r . i . e . , a . d . , m . f . , and w . l . s . analyzed data ; and t . j . n . , r . i . e . , a . d . , j . a . m . , p . c . w . , m . f . , and w . l . s . wrote the paper .
data deposition : the sequence reported in this paper has been deposited in the genbank database ( accession no . jx190073 – jx191369 ) .
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researchers used field data from 2012 to 2015 to study mortality and allele frequency changes in the sea star pisaster ochraceus during a mass mortality event in northcentral california , and found that surviving adult and juvenile sea stars experienced 81 % mortality and allele shifts , according to the authors .
a survey of more than 4 , 600 american adults conducted in 1995 - 1996 and in 2011 - 2014 suggests that among individuals of low socioeconomic status , negative affect increased significantly between the two survey waves , whereas life satisfaction and psychological well - being decreased .
a study of cognitive ability in norwegian males born from 1962 to 1991 suggests that environmental factors rather than changing genetic composition of families likely account for most of the change in norwegian population iq .
the mathematical tools behind recent gerrymandering cases have brought a modicum of precision into the political arena—but this rigor hasn’t always been enough to spur policy changes .
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urltoken
betancur - r . r , broughton re , wiley eo , carpenter k , lópez ja , li c , holcroft ni , arcila d , sanciangco m , cureton ii jc , zhang f , buser t , campbell ma , ballesteros ja , roa - varon a , willis s , borden wc , rowley t , reneau pc , hough dj , lu g , grande t , arratia g , ortí g . the tree of life and a new classification of bony fishes . plos currents tree of life . 2013 apr 18 . edition 1 . doi : 10 . 1371 / currents . tol . 53ba26640df0ccaee75bb165c8c26288 .
department of biological sciences , the george washington university , washington , dc , usa .
oklahoma biological survey and department of biology , university of oklahoma , norman , ok , usa .
biodiversity institute and department of ecology & evolutionary biology , university of kansas , lawrence , ks , usa , biologist at the university of kansas .
department of biological sciences , old dominion university , norfolk , virginia , u . s . a . , international union for conservation of nature .
fishes , university of alaska fairbanks , fairbanks , alaska , united states of america , university of alaska museum .
key laboratory of exploration and utilization of aquatic genetic resources , shanghai ocean university , ministry of education , shanghai 201306 , china .
science division , johnson county community college , overland park , kansas , usa .
department of biological sciences , old dominion university , norfolk , virginia , usa .
department of biology and oklahoma biological survey , university of oklahoma , norman , oklahoma , usa .
department of biology and oklahoma biologcial survey , university of oklahoma , norman , ok , usa , ph . d . student .
department of biology and wildlife , university of alaska fairbanks , fairbanks , ak , usa .
department of biological sciences , the george washington university , washington , dc , united states .
department of biological sciences , george washington university , washington , dc , usa , research assistant .
school of biological sciences , university of nebraska - lincoln , lincoln , ne , usa .
biology , university of nebraska - omaha , omaha , nebraska , united states .
department of biological sciences , florida a & m university , tallahassee , fl , usa .
systematics and ecology and biodiversity institute , university of kansas , lawrence , kansas , usa , courtesy research professor and associated researcher .
this work was supported by nsf awards deb - 0732988 ( to reb ) , deb - 0732838 , deb - 1019308 ( to go and cl ) , deb - 0732819 ( eow ) , deb 0732589 ( tg ) , deb - 0732894 ( kc ) , deb 0963767 ( jal ) , and deb - 0732969 ( gl ) ; gwu selective excellence in diversity of life program ( to rbr ) ; leading academic discipline project of shanghai municipal education commission , project number s30701 ( cl ) . the biodiversity institute , university of kansas , provided financial support for the collection used in this study .
our view of the phylogeny and classification of bony fishes is rapidly changing under the influence of molecular phylogenetic studies based on larger and more taxonomically comprehensive datasets . classification schemes displayed in widely used text books on fish biodiversity ( e . g . , 2 , 3 ) have been based on loosely formulated syntheses ( supertrees ) and community consensus views of largely disconnected studies . the phylogenetic structure underpinning such classifications has many areas that are notably unresolved and poorly known , providing weak or no justification for many groups that , although formally recognized , are implicitly known to be polyphyletic ( e . g . percoids , perciforms , scorpaeniforms ) . a comprehensive phylogenetic tree for all major groups of fishes has been elusive because explicit analyses including representatives across their diversity have never been accomplished . detailed morphological cladistic investigations of fish relationships have typically focused on lower taxonomic scales and few attempts to synthesize morphology at higher taxonomic levels proved to be challenging and met limited success ( e . g . , 4 ) . a recent effort to systematically collect morphological synapomorphies from published records for all currently recognized groups resulted in the first teleost classification based on monophyletic groups 5 . this effort , however , did not produce a global phylogenetic hypothesis . similarly , molecular analyses have been limited and many times conflicting in terms of genetic coverage and taxonomic sampling .
this study is the main product of the euteleost tree of life project ( etol ) . a total of 21 molecular markers with a genome - wide distribution were examined , the majority of which were developed by etol using a genomic screen pipeline 25 . this pipeline compared the danio rerio and takifugu rubripes genomes to identify single - copy genes with long exons ( > 800 bp ) and divergence levels suggesting they evolve at rates appropriate for phylogenetic resolution among distantly related taxa . exons markers were sequenced from 11 nuclear genes previously published by our group ( kiaa1239 , ficd , myh6 , panx2 , plagl2 , ptchd4 ( = ptr ) , ripk4 , sidkey , snx33 ( = sh3px3 ) , tbr1b ( = tbr1 ) , and zic1 ) and three additional markers , including one intron ( hoxc6a ) and two exons ( svep1 , and vcpip ) , were newly developed for this study using the same approach . sequence data from seven additional markers , including etol markers ( enc1 , gtdc2 ( = glyt ) , and gpr85 ( = sreb2 ) ) or markers developed by others ( 16s mtdna , rag1 , rag2 , and rh ) , were generated for our previous studies ( e . g . , 25 , 26 , 27 , 28 , 66 , 96 ) or obtained from ncbi , ensembl , or other genomic databases .
fish diversity is represented in the phylogenetic data matrix by a sample of 1410 bony fish species ( of ca . 31000 51 ) plus four tetrapod species and two chondrichthyan outgroups ( total 1416 terminals ) . the taxonomic sampling of bony fishes consists of 1093 genera ( of ca . 4300 ) , 369 families ( of 502 ; see below ) , and all traditionally recognized orders ( e . g . , 5 ) . our taxonomic sampling emphasizes representation of percomorph groups , with 1037 ( of > 15000 ) species in 201 families . all scientific names were checked against the catalog of fishes 51 . a complete list of material examined is given in table s1 .
* 1st and 2nd are primers for the first and nested / seminested ( optional ) rounds of pcr , respectively .
contigs were assembled from forward and reverse sequences using codoncode aligner v3 . 5 . 4 ( codoncode corporation ) , sequencher v4 ( gene codes corporation ) , or geneious pro v4 . 5 ( biomatters ltd . ) . exon markers were aligned individually based on their underlying reading frame in translatorx 52 using the mafft aligner 53 . the hoxc6a and 16s sequences were aligned with mafft v6 . 9 53 using 1000 iterations and the genafpair algorithm . because nested pcr is highly prone to cross - contamination , we vetted the data by visually inspecting individual gene trees estimated with the geneious tree builder algorithm in geneious . to qualitatively assess gene - tree congruence , the final gene alignments were analyzed under maximum likelihood ( ml ) in raxml using ten independent runs for each ; exon alignments were partitioned by codon position . alternative approaches to analyze combined data based on species - tree methods that account for gene - tree heterogeneity due to lineage sorting ( e . g . , 54 , 55 , 56 , 57 ) could not be applied to this dataset due to high proportion of missing data ( see results ) .
individual genes were concatenated using sequencematrix v1 . 7 . 8 58 or geneious . two datasets were assembled and analyzed separately , one including all 1416 taxa with sequence data from three genes or more ( 3 + dataset ) and a subset including 1020 taxa with sequence data from seven genes or more ( 7 + dataset ) . analyses of the 3 + dataset were performed under maximum likelihood ( ml ) using two partitioning schemes , a simple one determined arbitrarily with 5 data partitions ( 3 codon positions across all exons plus 16s and hoxc6a ) , and a more complex scheme with 24 partitions ( a combination of codon positions and individual genes plus 16s and hoxc6a ) indicated by partitionfinder 59 . to make the partitionfinder analysis scalable , a representative subset of 201 taxa was run under the bayesian information criterion 59 . the 7 + dataset was analyzed with the 24 - partition scheme only . analyses for both datasets and partition schemes were conducted in raxml using 30 independent replicates under the gtrgamma model . nodal support was assessed using the rapid bootstrapping algorithm of raxml with 1000 replicates estimated under the gtrcat model 60 , and the collection of sample trees was used to draw the bibartition frequencies on the optimal tree . all raxml analyses were conducted in the cipres portal v3 . 1 .
for comparison purposes , the 3 + dataset was also analyzed under implied - weighted parsimony 61 . the optimal tree search and bootstrap trees were set to run independently . gaps were treated as missing characters and all parsimony uninformative characters were ignored . a relatively mild value of k ( 20 ) was chosen arbitrarily due to computational limitations to explore sensitivity of the nodes to other weighting functions . tree searches were performed in tnt 1 . 1 63 using a driven - search strategy combining the following tree - search algorithms : ratchet , drift , sectorial searches and tree fusion . the exhaustiveness of the search parameters was self - adjusted every 2 hits of the current best score . to maximize tree - space exploration , the final searches implemented tree - bisection - reconnection ( tbr ) . a strict consensus of nine equally optimal trees ( length 407187 steps ; fit 7309 . 19 ) was computed . bootstrap search strategies were relaxed to ten random addition sequences and tbr , saving only one tree per replicate ( 1000 replicates ) ; bootstrap bipartition frequencies were drawn on the consensus tree .
the complete tree with 1416 taxa was time - calibrated under penalized likelihood ( pl 67 ) with treepl 69 . the pl model , which assumes rate autocorrelation , has been shown to perform poorly in simulation studies resulting in high stochastic error of divergence time estimates 70 . to ameliorate this problem , mean highest posterior density estimates of clade ages obtained with the subset in beast were imposed as fixed secondary calibrations for the pl analysis , rather than using primary calibrations with minimum and maximum age constrains . a total of 126 secondary calibrations were used for this analysis , including the ages obtained for all major groups in the tree as well as the nodes near which primary calibrations were defined . the rate smoothing parameter was set to 10 based on the cross - validation procedure and the χ 2 test in treepl ( four smoothing values between 1 and 1000 were compared ) .
based on the topology obtained ( figs . 1 - 10 , s1 ) we propose a new classification for ordinal and subordinal groups of bony fishes and subsequently discuss some of the most significant findings .
fig . 1 : main phylogenetic hypothesis of bony fish groups collapsed to depict higher - level clades .
the phylogenetic tree was estimated in raxml using the 3 + dataset ( 1416 taxa ) and 24 partitions with divergence times estimated under pl using 126 fixed secondary calibrations from the beast analysis ( see fig . 11 ) . terminal clades are either orders or supraordinal taxa with multiple orders included . values in parentheses indicate number of families examined . see also figs . 3 - 10 for relationship details on selected percomorph clades . the complete phylogeny with bootstrap support values and names for supraordinal taxa is in fig . s1 ) .
fish illustrations were obtained from fishes of the world ( nelson [ 2 ] ) and are reproduced with permission of john wiley & sons , inc . .
fig . 2 : sensitivity analyses for selected clades obtained in this study ( shown in figs . 1 , 3 - 10 ) and for selected alternative hypotheses .
for each case , we assess support from individual gene trees ( indicating whether the group was obtained ) or from the concatenated data sets ( indicating whether the group was obtained and showing boostrap support ) . for some gene trees , monophyletic groups ignore a few rogue taxa falling outside . n / a : insufficient taxonomic sampling to test hypothesis . 1 excluding gymnotiformes ( e . g . , saitoh et al . 77 ) ; 2 stomiatii ; 3 stiassny and moore 75 ; 4 nelson 2 and wiley and johnson 5 ; 5 shan and gras 47 ; 6 patterson and rosen 78 ; 7 arratia 79 ; 8 fink and fink 80 ; 9 nelson 81 ; 10 olney et al . 82 ; 11 johnson and patterson 4 ; 12 miya et al . 8 ; 13 johnson 83 ; 14 kaufman and liem 84 ; 15 gill 85 ; 16 acanthuriformes sensu tyler et al . 86 ( i . e . , acanthuriformes sensu stricto 87 plus scatophagidae and siganidae ) ; 17 jordan 88
the nomenclatural arrangement presented in appendix 2 builds on the existing classification by wiley and johnson 5 and intends to preserve names and taxonomic composition of groups whenever possible . however , adjustments are made to recognize new well - supported molecular clades , many of which also have been obtained by previous molecular studies ( several examples discussed below ) . order - level or supraordinal taxa are erected ( new ) or resurrected on the basis of well - supported clades only ( > 90 % bootstrap values ) . current taxon names supported by previous molecular or morphological studies are retained if congruent with our results , even if bootstrap support is low ( e . g . , osteoglossocephalai sensu arratia 79 with only 38 % bootstrap ) . in some cases , ordinal or subordinal taxa that were not monophyletic in our analysis are also validated , as long as the incongruence is not supported by strong bootstrap values . examples include the suborder blennioidei ( not monophyletic here but monophyletic in wainwright et al . 31 ) and the order pleuronectiformes ( not monophyletic here but monophyletic in betancur - r . et al . 28 ) .
our results ( appendix 2 ) invite comparison to the recent classification of wiley and johnson 5 based on morphological evidence gleamed from many investigators . of 123 clades recognized by them , 70 ( 56 . 9 % ) are congruent with bootstrap values > 95 % obtained in this study . five of these 70 clades are included in our sample by only one family and thus their monophyly is not critically tested . another six clades ( 4 . 9 % ) are congruent but are supported by lower bootstrap values ; seven additional clades ( 5 . 7 % ) are monotypic . forty clades ( 32 . 5 % ) are incongruent , with some being grossly polyphyletic in our tree . notable examples are protacanthopterygii , smegmamorpharia , and labriformes . others are incongruent based on exclusion of subclades and are rendered monophyletic in our classification by the addition or removal of smaller clades . examples include stomiatii ( inclusion of osmeriformes sensu stricto ) , otomorpha ( inclusion of alepocephaliformes ) , neoteleostei ( removal of stomiatiformes ) , and lampridiformes ( removal of stylephorus ) .
there is considerable consensus between morphology and the interrelationships of major clades . for example , the major cohorts of living teleosts and their interrelationships are congruent with the listing convention employed by wiley and johnson 5 ; this is also true within many of the major clades ( e . g . relationships within elopomorpha ) . but there is also incongruence . for example , relationships among early - branching acanthomorph groups differ considerably from previous morphological hypotheses ( e . g . , johnson and patterson 4 ) with lampridiforms , percopsiforms , zeiforms and gadiforms branching off basally relative to polymixiiforms . more explicit tests of new and alternative phylogenetic hypotheses based on multiple analyses of our dataset are presented in fig . 2 .
the following sections highlight some of the salient features of this global phylogeny and classification of bony fishes , especially in reference to well - established relationships and newly found clades among the euteleosts . we do not attempt to provide a complete account of all taxonomic issues , but to give some perspective and contrast to discuss the evidence supporting novel and established taxa .
our analyses support several recent hypotheses based on molecular data that contradict the consensus based on morphology 2 , 5 relative to the composition of “protacanthopterygians . ” although our results fall short of resolving with confidence circumscription and relationships among taxa in this group ( hence protacathopterygii is a sedis mutabilis taxon in our proposed classification ) , some relationships are well supported and consistent with previous studies ( fig . 1 ) . first , is the hypothesis that alepocephalid fishes ( slickheads ) have affinities within otomorpha , instead of argentiformes , as proposed by johnson and patterson 4 . this result was first proposed on the basis of mitogenomic data 10 , 41 , 100 , 101 and recently corroborated with a subset of the nuclear markers used in this study 29 . second , is the sister group relationship of osmeriformes and stomiatiformes ( = stomiiformes ) , first proposed by lópez et al . 21 based on mtdna and rag1 sequence data . finally , the position of lepidogalaxias at the base of the euteleosts rendering galaxiidae non - monophyletic also was proposed previously 102 , 29 and supported by our data ( see also fig . 2 ) .
a corollary of the increased resolution of percomorph relationships is the demise of the smegmamorpharia sensu johnson and patterson 4 ( see also wiley and johnson 5 ; fig . 2 ) . elements included in this supraordinal taxon are now scattered throughout the molecular phylogeny , placed within many of the newly found clades with high bootstrap support . for example , the pygmy sunfishes ( elassoma ) are back with the other sunfishes ( centrarchids ) , as suggested by earlier classifications and recently confirmed by molecules 30 . centrarchids plus elassomatids are placed here in the resurrected order centrarchiformes ( within percomorpharia , fig . 9 ) . mugiliforms ( mullets ) and atherinomorphs ( silversides , needlefishes , halfbeaks , guppies and allies ) are placed within ovalentariae ( fig . 8 ) . the swamp eels and spiny eels ( order synbranchiformes , suborders synbranchoidei and mastacembeloidei ) are placed with confidence in anabantomorphariae ( fig . 5 ) , together with armored sticklebacks ( indostomidae ) , one of the 11 families previously included in the order gasterosteiformes . the polyphyly of gasterosteiformes ( another large clade assigned to smegmamorpha ) was first pointed out by mitogenomic evidence 12 . our results place the sticklebacks , tubesnouts and sand eels ( previously assigned to gasterosteoidei ) in our newly defined perciformes ( suborder cottioidei ; fig . 10 ) and the rest of the families previously assigned to the suborder syngnathoidei were relocated to our newly defined order syngnathiformes within the scombrimorpharia ( fig . 4 , see below ) .
phylogenetic resolution within five newly discovered clades , however , will require additional study . relationships within syngnathiformes , scombriformes , carangimorphariae , ovalentariae , and percomorpharia may be challenging to recover given the rapid radiation and diversification of these clades .
based on a phylogeny estimated with four mitochondrial markers , thacker 33 resurrected the order gobiiformes , to accommodate three suborders : gobioidei ( gobies and sleepers ) , kurtidoidei ( nurseryfish ) , and apogonoidei ( including apogonids and pempherids ) . previous molecular studies have shown affinities between gobioids , apogonids , kurtids and , to some extent , pempherids and dactylopterids 8 , 11 , 16 . there is also morphological evidence supporting a close relationship between gobids and apogonids 108 , 109 as well as between kurtids and apogonids 110 . our results provide partial support for the gobiiformes sensu thacker 33 but we treat it here as a supraordinal group ( gobiomorpharia ) . a major difference is that our hypothesis segregates the family pempheridae ( sweepers ) to its own order ( pempheriformes , together with glaucosomatidae ) , within percomorpharia ( figs . 1 , 3 , 9 ) .
fig . 3 : detailed relationships among orders and families of gobiomorpharia ( see also fig . 1 ) .
fish illustrations were obtained from fishes of the world ( nelson [ 2 ] ) and are reproduced with permission of john wiley & sons , inc .
scombrimorpharia : sea horses and tunas are close relatives ( figs . 1 , 4 and 5 )
one of the most unanticipated new percomorph clades is the scombrimorpharia , grouping such disparate fishes as seahorses and tunas . this clade includes the newly circumscribed orders syngnathiformes ( fig . 4 ) and scombriformes ( fig . 5 ) . not surprisingly , a close relationship among taxa contained within this group , including syngnathids , mullids , callionymids , dactylopterids , scombrids , stromateids , an others , has never been proposed on morphological grounds . the syngnathiformes , as defined here ( fig . 4 ) , comprises mostly tropical marine reef - dwellers , traditionally placed in three distinct percomorph orders , including gasterosteiformes ( syngnathids ) , “perciformes” ( mullids and callionymids ) and “scorpaeniformes” ( dactylopterids ) . recent molecular studies have emphasized the non - monophyly of scorpaeniformes 74 . we have noted above the dissolution of gasterosteiformes 12 and , as discussed below , we provide a restricted definition for perciformes that includes many scorpaeniform taxa ( fig . 10 ) .
fig . 4 : detailed relationships among families of syngnathiformes ( see also fig . 1 ) .
our new order scombriformes ( fig . 5 ) includes most of the families previously grouped in the perciform suborder scombroidei 2 or the order scombriformes 5 , except for the barracudas ( sphyraenidae ) and the billfishes and swordfishes ( here placed in their own order , istiophoriformes ) . sphyraenidae and istiophoriformes are now firmly placed within carangimorphariae ( fig . 7 ) together with disparate taxa such as remoras ( echeneidae ) , archer fishes ( toxotidae ) , jacks ( carangidae ) , flatfishes ( pleuronectiformes ) , and others ( see below ) . because billfishes and tunas are not closely related as previously suggested by anatomical studies 83 ( fig . 2 ) , the new hypothesis implies that endothermy has evolved at least twice independently in teleosts 111 , 112 . this new circumscription of scombriformes also comprises families belonging to multiple orders in previous classifications , such as stromateiformes ( centrolophidae , nomeidae , ariommatidae , stromateidae ) , trachiniformes ( chiasmodontidae ) , icosteiformes ( icosteidae ) , and perciformes ( bramidae , pomatomidae , and caristiidae ) . despite the disparate morphology among members of scombriformes , most are offshore fishes that inhabit pelagic and / or deep - sea waters .
fig . 5 : detailed relationships among families of scombriformes ( see also fig . 1 ) .
another major percomorph group proposed here is the series carangimorpharia , including three subseries : anabantomorphariae , carangimorphariae , and ovalentariae ( fig . 1 ) . species in anabantomorphariae include representatives placed in three separate orders by wiley and johnson 5 : synbranchiformes ( swamp eels ) , gasterosteiformes ( indostomus , the armored stickleback ) , and anabantiformes ( gouramis ) ( fig . 6 ) . while the first two orders belonged to the smegmamorpharia 4 , 5 , the anabantiformes were placed as incertae sedis in percomorphacea 5 . the monophyly of anabantomorphariae has also been supported on the basis of mitogenomics 8 , 11 , 12 and nuclear markers 28 . a remarkable condition shared by members of this novel grouping is their mostly freshwater origin and restriction to africa and south east asia ( although some members in the family synbranchidae occur in mexico , and central and south america ) . most are able to occupy marginal , stagnant waters due to their capacity to tolerate anoxia and to obtain oxygen directly from the air . anabantiforms have a suprabranchial organ and synbranchids have suprabranchial pouches with respiratory function .
fig . 6 : detailed relationships among orders and families of anabantomorphariae ( see also fig . 1 ) .
a close affinity between other seemingly disparate groups , including barracudas , swordfishes , jacks , flatfishes , and others , has been well established by recent molecular studies 10 , 16 , 19 , 24 , 27 , 28 , 112 ( fig . 7 ) . this higher - level group has been referred to as ‘‘clade l’’ sensu chen et al . 19 or carangimorpha by li et al . 24 ( see also 27 , 28 ) . in looking for possible anatomical synapomorphies uniting flatfishes , billfishes , and carangids , little et al . 112 found that most taxa share a relatively low number of vertebrae , have multiple dorsal pterygiophores inserting before the second neural spine , and lack supraneurals , among others . however , according to friedman 113 , some of these characters are symplesiomorphies while others are absent in the remaining carangimorph groups . it thus seems paradoxical that despite the apparent lack of morphological synapomorphies for carangimorphs there is a strong molecular signal supporting their monophyly , whereas the opposite is true for pleuronectiforms 28 . for additional insights and discussion on carangimorphariae we refer the reader to recent studies 24 , 27 , 28 , 112 , 113 .
fig . 7 : detailed relationships among orders and families of carangimorphariae ( see also fig . 1 ) .
values in parentheses indicate number of genera examined ( see also betancur - r . et al . 28 ) .
ovalentariae is one of the most spectacular percomorph radiations , including more than 5000 species in some 44 families , grouping seemingly distinct groups such as cichlids , mullets , blennies , and atherinomorphs ( atheriniforms , beloniforms , and cyprinodontiforms ) . this clade was first found on the basis of mitogenomic evidence 8 , 12 and later confirmed with nuclear sequence data 23 , 24 , 26 , 31 . our results suggest that this group can be divided into four subgroups ( superorders ) , two of which already existed ( atherninomorphae and mugilomorphae ) and two that are new : ( i ) cichlomorphae ( cichlidae plus pholidichthyidae ) and ( ii ) blennimorphae ( blennioids plus clingfishes , jawfishes and basslets ) . many families in ovalentariae , however , remain incertae sedis ( e . g . , embiotocidae and pseudochromidae ) . two different studies have coined a name for this group ; first stiassnyiformes by li et al . 24 and , more recently , ovalentaria by wainwright et al . 31 for their characteristic demersal , adhesive eggs with chorionic filaments ( lost secondarily in some groups ) . an interesting implication of this phylogenetic hypothesis is that the pharyngeal jaw apparatus ( pharyngognathy ) , present in many members of this clade ( e . g . , cichlidae , pomacentridae , hemiramphidae ) , has evolved multiple times in percomorphs 31 . we refer the reader to wainwright et al . 31 for additional discussion on ovalentariae .
fig . 8 : detailed relationships among orders and families of ovalentariae ( see also fig . 1 ) .
values in parentheses indicate number of genera examined ( see also wainwright et al . 31 ) . many clades lacking taxonomic annotations on nodes are incertae sedis taxa ( for details , see classification ) .
percomorpharia is by far the largest percomorph clade , including 11 orders with some of the most prominent ones such as perciformes , labriformes , lophiiformes , and tetraodontiformes . at least 151 families ( 105 examined ) belong in percomorpharia , including three of the top ten most diverse families of fishes ( i . e . , labridae , serranidae , and scorpaenidae ) 2 . more than one third ( 514 ) of the species in our bony fish phylogeny are placed in this clade . previous molecular studies obtained monophyletic groups with a combination of taxa here assigned to percomorpharia , but with far more limited sampling ( e . g . , 8 , 11 , 16 , 74 ) . although most family - level and ordinal groups within percomorpharia receive high bootstrap support , interrelationships among them are largely unresolved ( hence , the new bush at the top ; fig . 9 ) . several of these groups are newly proposed or resurrected orders under new circumscription ( e . g . , uranoscopiformes , ephippiformes , pempheriformes ) . our new arrangement removes anglerfishes ( lophiiformes ) from paracanthomorphacea , as was suggested by previous classifications 78 , and places them close to tetraodontiforms , caproids , acanthuriforms , chaetodontids , pomacanthids , ephippids and others ( see also 87 , 114 , 115 ) . the largest group within percomorpharia is the order perciformes .
fig . 9 : detailed relationships among orders and families of percomorpharia ( the new bush at the top ; see also fig . 1 ) .
values in parentheses indicate number of genera examined in each terminal family or number of families and genera , respectively , in each terminal order . see also fig . 10 for expanded relationships on perciform groups . many clades lacking taxonomic annotations on nodes are incertae sedis taxa ( for details , see classification ) .
for the first time , a monophyletic definition of perciformes can be recovered from phylogenetic analysis of a comprehensive taxon sampling . the new circumscription of perciformes reduces significantly the number of included taxa , while retaining remarkable diversity that can be organized into several suborders and infraorders . nelson’s classification 2 included 160 families in perciformes , making it the largest order of all vertebrates . our definition indicates unambiguous membership for 38 families and uncertain membership for an additional 42 that were not examined in our study but that have been assigned to either “perciformes” ( 10 ) , “scorpaeniformes” ( 14 ) , cottiformes ( 8 ) , or trachiniformes ( 1 ) in previous classifications 2 , 5 . hence , the maximum possible number of families in the newly defined perciformes is reduced to 71 . this number is closer to the 90 families proposed by wiley and johnson 5 for their perciformes , but with a very different composition .
for a long time , perciformes has been regarded as a “taxonomic waste basket” 2 , 5 with ‘‘percoids’’ scattered throughout percomorpha and no clear phylogenetic distinction among percoidei , perciformes , and percomorpha 74 . earlier molecular studies lacked sufficient sampling to resolve phylogenetic questions among “percoids , ” but close relationships among groupers ( serranidae ) , perches ( percidae ) , sticklebacks ( gasterosteidae ) , searobins ( triglidae ) , icefishes ( notothenioidei ) , sculpins ( cottoidei ) , eelpouts ( zoarcoidei ) and scorpionfishes ( scorpaenoidei ) have been obtained in one form or another , and in different combinations , by several authors 16 , 19 , 20 , 23 , 24 , 29 , 74 , 116 . all of these taxa are included in our definition of perciformes ( fig . 10 ) .
within perciformes , we tentatively propose suborders ( notothenioidei , scorpaenoidei , trigloidei , cottoidei ) for clades with high support that also represent some well - established groups , but two incertae sedis ( percophidae and platycephalidae ) , and several unexamined families remain unclassified . additional taxon sampling and more data are needed to resolve interrelationships among these taxa . four suborders / infraorders were recognized as separate orders by wiley and johnson 5 : percoidei , scorpaenoidei , cottioidei , and gasterosteales ( an infraorder of cottioidei ) .