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A Phylogenetic Investigation of<br />

the Planthopper Superfamily<br />

Fulgoroidea (Insecta:<br />

Hemiptera), with Emphasis on<br />

the Family <strong>Fulgoridae</strong><br />

DEFENDED!<br />

Julie Urban<br />

University at Albany &<br />

New York State Museum


First Phylogeny of <strong>Fulgoridae</strong> -molecular<br />

data<br />

(Urban & Cryan, accepted MP&E)


Taxonomic sampling:<br />

<strong>Fulgoridae</strong> -- molecular<br />

Ingroup: 69 taxa in 46 genera<br />

-- representatives of 8 of 8 subfamilies, 10 of 11 tribes<br />

Outgroups:<br />

Dictyopharidae (7)<br />

Cixiidae (2), Delphacidae (1)<br />

Molecular data -- ~ 5000 bp<br />

18S rDNA ~1,900 bp<br />

28S rDNA ~2,200 bp<br />

Histone 3 ~ 350 bp<br />

Wingless ~ 350 bp<br />

COI ~ 500 bp


Bothriocera eborea<br />

Oecleus sp.<br />

Tropidocephala sp.<br />

Zanna sp.1<br />

Zanna sp.2<br />

Zanna sp.3<br />

Raphiophora sp.<br />

Scolops sulcipes<br />

Dictyophara sp.<br />

undet. Dictyopharidae<br />

Rhynchomitra microrhina<br />

Hyalodictyon sp.<br />

Lappida sp.<br />

Xosophara sp.<br />

Neolieftinckana fuscata<br />

Saiva nodata<br />

Pyrops sp.<br />

Pyrops intricata<br />

Penthicodes sp.1<br />

Penthicodes pulchella<br />

Penthicodes sp.2<br />

Kalidasa lanata<br />

Anecphora torrida<br />

undet. <strong>Fulgoridae</strong><br />

Druentia sicca<br />

Rentinus dilatatus<br />

Eurinopsyche obscurata<br />

Birdantis blotei<br />

Desudaba danae<br />

Odontoptera carrenoi<br />

Odontoptera toulgoeti<br />

Stalubra sp.<br />

Cathedra serrata<br />

Fulgora sp.2<br />

Fulgora sp.1<br />

Fulgora laternaria<br />

Phrictus regalis<br />

Phrictus tripartitus<br />

Phrictus quinquepartitus<br />

Enchophora nigromaculata<br />

Enchophora sanguinea var.1<br />

Enchophora sanguinea var.2<br />

Artacie dufouri<br />

Enchophora rosacea<br />

Enchophora stillifera<br />

Amerzanna peruana<br />

Amycle vernalis<br />

Cyrpoptus suavis<br />

Calyptoproctus sp.<br />

Paralystra florenciana<br />

Pterodictya reticularis<br />

Phenax variegata<br />

Episcius boliviana<br />

Episcius guianensis<br />

Weyrauchia fulminea<br />

Echetra modesta<br />

Echetra nebulosa<br />

Aracynthus sanguineus<br />

Neocynthus crassus var. 1<br />

Neocynthus crassus var. 2<br />

Diareusa imitatrix<br />

Diareusa annularis<br />

Aphrodisias cacica<br />

Sinuala tuberculata<br />

Copidocephala guttata<br />

Copidocephala viridiguttata<br />

Flatolystra sp.1<br />

Flatolystra sp.2<br />

Dilobura conspurcata<br />

Enhydria sp.<br />

Lystra lanata<br />

Learcha sponsa<br />

Matacosa miscella<br />

Scaralis semilimpida<br />

Poblicia fuliginosa<br />

Alphina glauca<br />

Scaralis sp.1<br />

Scaralis sp.2<br />

Scaralis sp.3<br />

MP analysis<br />

-- 19 equally parsimonious<br />

topologies<br />

-- robust to different weightings


Bothriocera eborea<br />

Oecleus sp.<br />

Tropidocephala sp.<br />

Zanna sp.1<br />

Zanna sp.2<br />

Zanna sp.3<br />

Raphiophora sp.<br />

Scolops sulcipes<br />

Dictyophara sp.<br />

undet. Dictyopharidae<br />

Rhynchomitra microrhina<br />

Hyalodictyon sp.<br />

Lappida sp.<br />

Xosophara sp.<br />

Neolieftinckana fuscata<br />

Saiva nodata<br />

Pyrops sp.<br />

Pyrops intricata<br />

Penthicodes sp.1<br />

Penthicodes pulchella<br />

Penthicodes sp.2<br />

Kalidasa lanata<br />

Anecphora torrida<br />

undet. <strong>Fulgoridae</strong><br />

Druentia sicca<br />

Rentinus dilatatus<br />

Eurinopsyche obscurata<br />

Birdantis blotei<br />

Desudaba danae<br />

Odontoptera carrenoi<br />

Odontoptera toulgoeti<br />

Stalubra sp.<br />

Cathedra serrata<br />

Fulgora sp.2<br />

Fulgora sp.1<br />

Fulgora laternaria<br />

Phrictus regalis<br />

Phrictus tripartitus<br />

Phrictus quinquepartitus<br />

Enchophora nigromaculata<br />

Enchophora sanguinea var.1<br />

Enchophora sanguinea var.2<br />

Artacie dufouri<br />

Enchophora rosacea<br />

Enchophora stillifera<br />

Amerzanna peruana<br />

Amycle vernalis<br />

Cyrpoptus suavis<br />

Calyptoproctus sp.<br />

Paralystra florenciana<br />

Pterodictya reticularis<br />

Phenax variegata<br />

Episcius boliviana<br />

Episcius guianensis<br />

Weyrauchia fulminea<br />

Echetra modesta<br />

Echetra nebulosa<br />

Aracynthus sanguineus<br />

Neocynthus crassus var. 1<br />

Neocynthus crassus var. 2<br />

Diareusa imitatrix<br />

Diareusa annularis<br />

Aphrodisias cacica<br />

Sinuala tuberculata<br />

Copidocephala guttata<br />

Copidocephala viridiguttata<br />

Flatolystra sp.1<br />

Flatolystra sp.2<br />

Dilobura conspurcata<br />

Enhydria sp.<br />

Lystra lanata<br />

Learcha sponsa<br />

Matacosa miscella<br />

Scaralis semilimpida<br />

Poblicia fuliginosa<br />

Alphina glauca<br />

Scaralis sp.1<br />

Scaralis sp.2<br />

Scaralis sp.3<br />

Cixiidae<br />

Delphacidae<br />

Dictyopharidae<br />

MP analysis<br />

-- 19 equally parsimonious<br />

topologies<br />

-- robust to different weightings<br />

<strong>Fulgoridae</strong> not monophyletic<br />

with respect to Dictyopharidae<br />

Dictyopharids<br />

Zanna


<strong>Fulgoridae</strong><br />

Bothriocera eborea<br />

Oecleus sp.<br />

Tropidocephala sp.<br />

Zanna sp.1<br />

Zanna sp.2<br />

Zanna sp.3<br />

Raphiophora sp.<br />

Scolops sulcipes<br />

Dictyophara sp.<br />

undet. Dictyopharidae<br />

Rhynchomitra microrhina<br />

Hyalodictyon sp.<br />

Lappida sp.<br />

Xosophara sp.<br />

Neolieftinckana fuscata<br />

Saiva nodata<br />

Pyrops sp.<br />

Pyrops intricata<br />

Penthicodes sp.1<br />

Penthicodes pulchella<br />

Penthicodes sp.2<br />

Kalidasa lanata<br />

Anecphora torrida<br />

undet. <strong>Fulgoridae</strong><br />

Druentia sicca<br />

Rentinus dilatatus<br />

Eurinopsyche obscurata<br />

Birdantis blotei<br />

Desudaba danae<br />

Odontoptera carrenoi<br />

Odontoptera toulgoeti<br />

Stalubra sp.<br />

Cathedra serrata<br />

Fulgora sp.2<br />

Fulgora sp.1<br />

Fulgora laternaria<br />

Phrictus regalis<br />

Phrictus tripartitus<br />

Phrictus quinquepartitus<br />

Enchophora nigromaculata<br />

Enchophora sanguinea var.1<br />

Enchophora sanguinea var.2<br />

Artacie dufouri<br />

Enchophora rosacea<br />

Enchophora stillifera<br />

Amerzanna peruana<br />

Amycle vernalis<br />

Cyrpoptus suavis<br />

Calyptoproctus sp.<br />

Paralystra florenciana<br />

Pterodictya reticularis<br />

Phenax variegata<br />

Episcius boliviana<br />

Episcius guianensis<br />

Weyrauchia fulminea<br />

Echetra modesta<br />

Echetra nebulosa<br />

Aracynthus sanguineus<br />

Neocynthus crassus var. 1<br />

Neocynthus crassus var. 2<br />

Diareusa imitatrix<br />

Diareusa annularis<br />

Aphrodisias cacica<br />

Sinuala tuberculata<br />

Copidocephala guttata<br />

Copidocephala viridiguttata<br />

Flatolystra sp.1<br />

Flatolystra sp.2<br />

Dilobura conspurcata<br />

Enhydria sp.<br />

Lystra lanata<br />

Learcha sponsa<br />

Matacosa miscella<br />

Scaralis semilimpida<br />

Poblicia fuliginosa<br />

Alphina glauca<br />

Scaralis sp.1<br />

Scaralis sp.2<br />

Scaralis sp.3<br />

Cixiidae<br />

Delphacidae<br />

Dictyopharidae<br />

MP analysis<br />

-- 19 equally parsimonious<br />

topologies<br />

-- robust to different weightings<br />

<strong>Fulgoridae</strong> not monophyletic<br />

with respect to Dictyopharidae


<strong>Fulgoridae</strong><br />

Bothriocera eborea<br />

Oecleus sp.<br />

Tropidocephala sp.<br />

Zanna sp.1<br />

Zanna sp.2<br />

Zanna sp.3<br />

Raphiophora sp.<br />

Scolops sulcipes<br />

Dictyophara sp.<br />

undet. Dictyopharidae<br />

Rhynchomitra microrhina<br />

Hyalodictyon sp.<br />

Lappida sp.<br />

Xosophara sp.<br />

Neolieftinckana fuscata<br />

Saiva nodata<br />

Pyrops sp.<br />

Pyrops intricata<br />

Penthicodes sp.1<br />

Penthicodes pulchella<br />

Penthicodes sp.2<br />

Kalidasa lanata<br />

Anecphora torrida<br />

undet. <strong>Fulgoridae</strong><br />

Druentia sicca<br />

Rentinus dilatatus<br />

Eurinopsyche obscurata<br />

Birdantis blotei<br />

Desudaba danae<br />

Odontoptera carrenoi<br />

Odontoptera toulgoeti<br />

Stalubra sp.<br />

Cathedra serrata<br />

Fulgora sp.2<br />

Fulgora sp.1<br />

Fulgora laternaria<br />

Phrictus regalis<br />

Phrictus tripartitus<br />

Phrictus quinquepartitus<br />

Enchophora nigromaculata<br />

Enchophora sanguinea var.1<br />

Enchophora sanguinea var.2<br />

Artacie dufouri<br />

Enchophora rosacea<br />

Enchophora stillifera<br />

Amerzanna peruana<br />

Amycle vernalis<br />

Cyrpoptus suavis<br />

Calyptoproctus sp.<br />

Paralystra florenciana<br />

Pterodictya reticularis<br />

Phenax variegata<br />

Episcius boliviana<br />

Episcius guianensis<br />

Weyrauchia fulminea<br />

Echetra modesta<br />

Echetra nebulosa<br />

Aracynthus sanguineus<br />

Neocynthus crassus var. 1<br />

Neocynthus crassus var. 2<br />

Diareusa imitatrix<br />

Diareusa annularis<br />

Aphrodisias cacica<br />

Sinuala tuberculata<br />

Copidocephala guttata<br />

Copidocephala viridiguttata<br />

Flatolystra sp.1<br />

Flatolystra sp.2<br />

Dilobura conspurcata<br />

Enhydria sp.<br />

Lystra lanata<br />

Learcha sponsa<br />

Matacosa miscella<br />

Scaralis semilimpida<br />

Poblicia fuliginosa<br />

Alphina glauca<br />

Scaralis sp.1<br />

Scaralis sp.2<br />

Scaralis sp.3<br />

Cixiidae<br />

Delphacidae<br />

Dictyopharidae<br />

Tests of classification:<br />

-- Reject monophyly of family<br />

-- Reject monophyly of 5 of 7<br />

tested subfamilies<br />

-- Reject monophyly of 5 of 6<br />

tested tribes


NW<br />

Bothriocera eborea<br />

Oecleus sp.<br />

Tropidocephala sp.<br />

Zanna sp.1<br />

Zanna sp.2<br />

Zanna sp.3<br />

Raphiophora sp.<br />

Scolops sulcipes<br />

Dictyophara sp.<br />

undet. Dictyopharidae<br />

Rhynchomitra microrhina<br />

Hyalodictyon sp.<br />

Lappida sp.<br />

Xosophara sp.<br />

Neolieftinckana fuscata<br />

Saiva nodata<br />

Pyrops sp.<br />

Pyrops intricata<br />

Penthicodes sp.1<br />

Penthicodes pulchella<br />

Penthicodes sp.2<br />

Kalidasa lanata<br />

Anecphora torrida<br />

undet. <strong>Fulgoridae</strong><br />

Druentia sicca<br />

Rentinus dilatatus<br />

Eurinopsyche obscurata<br />

Birdantis blotei<br />

Desudaba danae<br />

Odontoptera carrenoi<br />

Odontoptera toulgoeti<br />

Stalubra sp.<br />

Cathedra serrata<br />

Fulgora sp.2<br />

Fulgora sp.1<br />

Fulgora laternaria<br />

Phrictus regalis<br />

Phrictus tripartitus<br />

Phrictus quinquepartitus<br />

Enchophora nigromaculata<br />

Enchophora sanguinea var.1<br />

Enchophora sanguinea var.2<br />

Artacie dufouri<br />

Enchophora rosacea<br />

Enchophora stillifera<br />

Amerzanna peruana<br />

Amycle vernalis<br />

Cyrpoptus suavis<br />

Calyptoproctus sp.<br />

Paralystra florenciana<br />

Pterodictya reticularis<br />

Phenax variegata<br />

Episcius boliviana<br />

Episcius guianensis<br />

Weyrauchia fulminea<br />

Echetra modesta<br />

Echetra nebulosa<br />

Aracynthus sanguineus<br />

Neocynthus crassus var. 1<br />

Neocynthus crassus var. 2<br />

Diareusa imitatrix<br />

Diareusa annularis<br />

Aphrodisias cacica<br />

Sinuala tuberculata<br />

Copidocephala guttata<br />

Copidocephala viridiguttata<br />

Flatolystra sp.1<br />

Flatolystra sp.2<br />

Dilobura conspurcata<br />

Enhydria sp.<br />

Lystra lanata<br />

Learcha sponsa<br />

Matacosa miscella<br />

Scaralis semilimpida<br />

Poblicia fuliginosa<br />

Alphina glauca<br />

Scaralis sp.1<br />

Scaralis sp.2<br />

Scaralis sp.3<br />

Cixiidae<br />

Delphacidae<br />

Dictyopharidae<br />

Nearctic<br />

Nearctic<br />

Biogeography:<br />

-- Old World origin<br />

-- Monophyletic New World<br />

(colonization of Neotropics)<br />

-- Multiple dispersal events<br />

into Nearctic<br />

Mapping Head Process:<br />

-- 4-7 losses of head process<br />

across fulgorid evolution


1<br />

2<br />

5<br />

6<br />

12<br />

13<br />

7<br />

10<br />

Hyalodictyon sp.<br />

Xosophara sp.<br />

Pyrops sp.<br />

Pyrops delessertii<br />

Saiva nodata<br />

Anecphora sp.<br />

Penthicodes farinosa<br />

Penthicodes pulchella<br />

3<br />

4<br />

Zanna sp.3<br />

Zanna sp.1<br />

Zanna sp.2<br />

Zanna<br />

Lappida sp.<br />

Raphiophora sp. LappidaRaphiophoraScolops<br />

8<br />

9<br />

Dictyophara sp.1<br />

Dictyophara sp.2<br />

Scolops sulcipes<br />

Rhynchomitra microrhina<br />

DictyopharaRhynchomitra<br />

Hyalodictyon<br />

sp.1<br />

15<br />

11 14<br />

18<br />

16<br />

17<br />

Penthicodes atomaria<br />

Druentia sicca<br />

undet. <strong>Fulgoridae</strong><br />

21<br />

20<br />

22<br />

23<br />

Kalidasa lanata<br />

Neolieftinckana fuscata<br />

Birdantis blötei<br />

Desudaba danaë<br />

Eurinopsyche sp.<br />

Eurinopsyche obscurata<br />

25<br />

Odontoptera carrenoi<br />

Odontoptera toulgoeti<br />

30<br />

31<br />

Enchophora sanguinea var.1<br />

Enchophora sanguinea var.2<br />

19<br />

29<br />

28<br />

33<br />

32<br />

Enchophora nigromaculata<br />

Enchophora rosacea<br />

Enchophora stillifera<br />

Artacie dufourii<br />

27 34<br />

35<br />

Phrictus tripartitus<br />

Phrictus quinquepartitus<br />

Phrictus regalis<br />

Cathedra serrata<br />

26<br />

24<br />

36<br />

37<br />

Fulgora castresii<br />

Fulgora laternaria<br />

Fulgora cearensis<br />

Stalubra brunnea<br />

Pterodictya reticularis<br />

Phenax variegata<br />

Echetra nebulosa<br />

Episcius bolivianus<br />

Aracynthus sanguineus<br />

Neocynthus crassus<br />

Neocynthus sp.<br />

Diareusa imitatrix<br />

Diareusa annularis<br />

Sinuala tuberculata<br />

Aphrodisias cacica<br />

Copidocephala guttata<br />

Copidocephala viridiguttata<br />

Echetra modesta<br />

Weyrauchia peruviana<br />

Episcius guianensis<br />

Cyrpoptus suavis<br />

Amycle vernalis<br />

Amerzanna peruana<br />

Calyptoproctus sp.<br />

Paralystra florenciana<br />

Flatolystra distincta<br />

Flatolystra verrucosa<br />

Enhydria tessellata<br />

Dilobura conspurcata<br />

Lystra lanata<br />

Learcha sponsa<br />

Poblicia fuliginosa<br />

Scaralis sp.2<br />

Matacosa miscella<br />

Scaralis semilimpida<br />

Alphina glauca<br />

Scaralis sp.1<br />

Scaralis sp.3<br />

38<br />

XosopharaPyrops Saiva<br />

AnecphoraPenthicodesDruentiaUndetermined<br />

Kalidasa<br />

Neolieftinckana BirdantisDesudabaEurinopsyche<br />

Odontoptera<br />

EnchophoraArtacie<br />

Phrictus<br />

Cathedra Fulgora Stalubra<br />

41<br />

43 44<br />

Pterodictya Phenax<br />

40<br />

45<br />

Echetra Episcius AracynthusNeocynthus<br />

39<br />

42<br />

47<br />

46<br />

51<br />

49<br />

52<br />

48<br />

50<br />

DiareusaSinualaAphrodisias Copidocephala<br />

55<br />

54<br />

56<br />

57<br />

53<br />

60<br />

59<br />

61<br />

58<br />

63<br />

62<br />

CyrpoptusAmycle<br />

EchetraWeyrauchiaEpiscius<br />

AmerzannaCalyptoproctus Paralystra<br />

Flatolystra EnhydriaDilobura<br />

64<br />

65<br />

67<br />

66<br />

69<br />

68<br />

Poblicia Scaralis<br />

Lystra Learcha<br />

Matacosa Alphina


<strong>Fulgoridae</strong> -- molecules + morphology<br />

Taxonomic sampling<br />

69 ingroup taxa + 10 outgroups<br />

Morphological data -- 110 characters<br />

Characters and states developed from literature on<br />

<strong>Fulgoridae</strong> and other planthopper families<br />

Molecular data -- ~ 5000 bp<br />

18S rDNA ~1,900 bp<br />

28S rDNA ~2,200 bp<br />

Histone 3 ~ 350bp<br />

Wingless ~ 350bp<br />

COI ~ 500bp


Australia<br />

+ PNG<br />

Africa + Asia<br />

NW<br />

Bothriocera eborea<br />

Oecleus sp.<br />

Tropidocephala sp.<br />

Zanna sp.3<br />

Zanna sp.1<br />

Zanna sp.2<br />

Lappida sp.<br />

Rhaphiophora sp.<br />

Scolops sulcipes<br />

Dictyophara sp.1<br />

Dictyophara sp.2<br />

Rhynchomitra microrhina<br />

Hyalodictyon sp.<br />

Xosophara sp.<br />

Neolieftinckana fuscata<br />

Birdantis blotei<br />

Rentinusdilatatus<br />

Eurinopsyche obscurata<br />

Desudaba danae<br />

Saiva nodata<br />

Pyrops sp.<br />

Pyrops intricata<br />

Kalidasa lanata<br />

Anecphora torrida<br />

Penthicodes farinosa<br />

Penthicodes pulchella<br />

Penthicodes atomaria<br />

Druentia sicca<br />

undet. <strong>Fulgoridae</strong><br />

Stalubra brunnea<br />

Enhydria tessellata<br />

Odontopera carrenoi<br />

Odontoptera toulgoeti<br />

Cathedra serrata<br />

Fulgora cearensis<br />

Fulgora castresii<br />

Fulgora laternaria<br />

Phrictus regalis<br />

Phrictus tripartitus<br />

Phrictusquinquepartitus<br />

Enchophora nigromaculata<br />

Enchophora sanguinea var.1<br />

Enchophora sanguinea var.2<br />

Artacie dufouri<br />

Enchophora rosacea<br />

Enchophora stillifera<br />

Pterodictya reticularis<br />

Phenax variegata<br />

Copidocephala guttata<br />

Copidocephala viridiguttata<br />

Diareusa imitatrix<br />

Diareusa annularis<br />

Aphrodisias cacica<br />

Sinuala tuberculata<br />

Episciusboliviana<br />

Episciusguianensis<br />

Weyrauchia fulminea<br />

Aracynthus sanguineus<br />

Neocynthus crassus<br />

Neocynthus sp.<br />

Echetra modesta<br />

Echetra nebulosa<br />

Cyrpoptus suavis<br />

Amycle vernalis<br />

Amerzanna peruana<br />

Calyptoproctussp.<br />

Paralystra florenciana<br />

Dilobura conspurcata<br />

Flatolystra distincta<br />

Flatolystra verrucosa<br />

Scaralis sp.1<br />

Scaralis sp.3<br />

Scaralis sp.2<br />

Matacosa miscella<br />

Scaralis semilimpida<br />

Alphina glauca<br />

Poblicia fuliginosa<br />

Lystra lanata<br />

Learcha sponsa<br />

Bayesian analysis<br />

Zanna placed outside of<br />

<strong>Fulgoridae</strong><br />

Monophyletic New World<br />

lineage<br />

2 Old World lineages?


Australia<br />

+ PNG<br />

Africa + Asia<br />

NW<br />

Bothriocera eborea<br />

Oecleus sp.<br />

Tropidocephala sp.<br />

Zanna sp.3<br />

Zanna sp.1<br />

Zanna sp.2<br />

Lappida sp.<br />

Rhaphiophora sp.<br />

Scolops sulcipes<br />

Dictyophara sp.1<br />

Dictyophara sp.2<br />

Rhynchomitra microrhina<br />

Hyalodictyon sp.<br />

Xosophara sp.<br />

Neolieftinckana fuscata<br />

Birdantis blotei<br />

Rentinusdilatatus<br />

Eurinopsyche obscurata<br />

Desudaba danae<br />

Saiva nodata<br />

Pyrops sp.<br />

Pyrops intricata<br />

Kalidasa lanata<br />

Anecphora torrida<br />

Penthicodes farinosa<br />

Penthicodes pulchella<br />

Penthicodes atomaria<br />

Druentia sicca<br />

undet. <strong>Fulgoridae</strong><br />

Stalubra brunnea<br />

Enhydria tessellata<br />

Odontopera carrenoi<br />

Odontoptera toulgoeti<br />

Cathedra serrata<br />

Fulgora cearensis<br />

Fulgora castresii<br />

Fulgora laternaria<br />

Phrictus regalis<br />

Phrictus tripartitus<br />

Phrictusquinquepartitus<br />

Enchophora nigromaculata<br />

Enchophora sanguinea var.1<br />

Enchophora sanguinea var.2<br />

Artacie dufouri<br />

Enchophora rosacea<br />

Enchophora stillifera<br />

Pterodictya reticularis<br />

Phenax variegata<br />

Copidocephala guttata<br />

Copidocephala viridiguttata<br />

Diareusa imitatrix<br />

Diareusa annularis<br />

Aphrodisias cacica<br />

Sinuala tuberculata<br />

Episciusboliviana<br />

Episciusguianensis<br />

Weyrauchia fulminea<br />

Aracynthus sanguineus<br />

Neocynthus crassus<br />

Neocynthus sp.<br />

Echetra modesta<br />

Echetra nebulosa<br />

Cyrpoptus suavis<br />

Amycle vernalis<br />

Amerzanna peruana<br />

Calyptoproctussp.<br />

Paralystra florenciana<br />

Dilobura conspurcata<br />

Flatolystra distincta<br />

Flatolystra verrucosa<br />

Scaralis sp.1<br />

Scaralis sp.3<br />

Scaralis sp.2<br />

Matacosa miscella<br />

Scaralis semilimpida<br />

Alphina glauca<br />

Poblicia fuliginosa<br />

Lystra lanata<br />

Learcha sponsa


<strong>Fulgoridae</strong> II (Urban & Cryan, in progress)<br />

Increasing taxonomic sampling<br />

13 genera new to data set<br />

40 species new to data set<br />

Reworking some morphological characters<br />

Objective: Basis for revised classification


Bothriocera eborea<br />

Oecleus sp.<br />

Tropidocephala sp.<br />

Zanna sp.1<br />

Zanna sp.2<br />

Zanna sp.3<br />

Raphiophora sp.<br />

Scolops sulcipes<br />

Dictyophara sp.<br />

undet. Dictyopharidae<br />

Rhynchomitra microrhina<br />

Hyalodictyon sp.<br />

Lappida sp.<br />

Xosophara sp.<br />

Neolieftinckana fuscata<br />

Saiva nodata<br />

Pyrops sp.<br />

Pyrops intricata<br />

Penthicodes sp.1<br />

Penthicodes pulchella<br />

Penthicodes sp.2<br />

Kalidasa lanata<br />

Anecphora torrida<br />

undet. <strong>Fulgoridae</strong><br />

Druentia sicca<br />

Rentinus dilatatus<br />

Eurinopsyche obscurata<br />

Birdantis blotei<br />

Desudaba danae<br />

Odontoptera carrenoi<br />

Odontoptera toulgoeti<br />

Stalubra sp.<br />

Cathedra serrata<br />

Fulgora sp.2<br />

Fulgora sp.1<br />

Fulgora laternaria<br />

Phrictus regalis<br />

Phrictus tripartitus<br />

Phrictus quinquepartitus<br />

Enchophora nigromaculata<br />

Enchophora sanguinea var.1<br />

Enchophora sanguinea var.2<br />

Artacie dufouri<br />

Enchophora rosacea<br />

Enchophora stillifera<br />

Amerzanna peruana<br />

Amycle vernalis<br />

Cyrpoptus suavis<br />

Calyptoproctus sp.<br />

Paralystra florenciana<br />

Pterodictya reticularis<br />

Phenax variegata<br />

Episcius boliviana<br />

Episcius guianensis<br />

Weyrauchia fulminea<br />

Echetra modesta<br />

Echetra nebulosa<br />

Aracynthus sanguineus<br />

Neocynthus crassus var. 1<br />

Neocynthus crassus var. 2<br />

Diareusa imitatrix<br />

Diareusa annularis<br />

Aphrodisias cacica<br />

Sinuala tuberculata<br />

Copidocephala guttata<br />

Copidocephala viridiguttata<br />

Flatolystra sp.1<br />

Flatolystra sp.2<br />

Dilobura conspurcata<br />

Enhydria sp.<br />

Lystra lanata<br />

Learcha sponsa<br />

Matacosa miscella<br />

Scaralis semilimpida<br />

Poblicia fuliginosa<br />

Alphina glauca<br />

Scaralis sp.1<br />

Scaralis sp.2<br />

Scaralis sp.3<br />

Cixiidae<br />

Delphacidae<br />

Dictyopharidae<br />

Increased sampling of Zanna


Bothriocera eborea<br />

Oecleus sp.<br />

Tropidocephala sp.<br />

Zanna sp.1<br />

Zanna sp.2<br />

Zanna sp.3<br />

Raphiophora sp.<br />

Scolops sulcipes<br />

Dictyophara sp.<br />

undet. Dictyopharidae<br />

Rhynchomitra microrhina<br />

Hyalodictyon sp.<br />

Lappida sp.<br />

Xosophara sp.<br />

Neolieftinckana fuscata<br />

Saiva nodata<br />

Pyrops sp.<br />

Pyrops intricata<br />

Penthicodes sp.1<br />

Penthicodes pulchella<br />

Penthicodes sp.2<br />

Kalidasa lanata<br />

Anecphora torrida<br />

undet. <strong>Fulgoridae</strong><br />

Druentia sicca<br />

Rentinus dilatatus<br />

Eurinopsyche obscurata<br />

Birdantis blotei<br />

Desudaba danae<br />

Odontoptera carrenoi<br />

Odontoptera toulgoeti<br />

Stalubra sp.<br />

Cathedra serrata<br />

Fulgora sp.2<br />

Fulgora sp.1<br />

Fulgora laternaria<br />

Phrictus regalis<br />

Phrictus tripartitus<br />

Phrictus quinquepartitus<br />

Enchophora nigromaculata<br />

Enchophora sanguinea var.1<br />

Enchophora sanguinea var.2<br />

Artacie dufouri<br />

Enchophora rosacea<br />

Enchophora stillifera<br />

Amerzanna peruana<br />

Amycle vernalis<br />

Cyrpoptus suavis<br />

Calyptoproctus sp.<br />

Paralystra florenciana<br />

Pterodictya reticularis<br />

Phenax variegata<br />

Episcius boliviana<br />

Episcius guianensis<br />

Weyrauchia fulminea<br />

Echetra modesta<br />

Echetra nebulosa<br />

Aracynthus sanguineus<br />

Neocynthus crassus var. 1<br />

Neocynthus crassus var. 2<br />

Diareusa imitatrix<br />

Diareusa annularis<br />

Aphrodisias cacica<br />

Sinuala tuberculata<br />

Copidocephala guttata<br />

Copidocephala viridiguttata<br />

Flatolystra sp.1<br />

Flatolystra sp.2<br />

Dilobura conspurcata<br />

Enhydria sp.<br />

Lystra lanata<br />

Learcha sponsa<br />

Matacosa miscella<br />

Scaralis semilimpida<br />

Poblicia fuliginosa<br />

Alphina glauca<br />

Scaralis sp.1<br />

Scaralis sp.2<br />

Scaralis sp.3<br />

Cixiidae<br />

Delphacidae<br />

Dictyopharidae<br />

Increased sampling of Zanna<br />

Increased sampling within<br />

Dictyopharidae<br />

Increased Old World<br />

sampling (3 genera)<br />

Increased sampling within<br />

genera


Endosymbionts<br />

Within <strong>Fulgoridae</strong> and Dictyopharidae<br />

Sulcia muelleri<br />

NW<br />

Aphrophora quadrinotata<br />

Clastoptera arizonana<br />

Scolops sulcipes<br />

Scolops sp.<br />

Dictyophara sp.1<br />

Dictyophara sp.2<br />

Rhaphiophora sp.<br />

Zanna sp.<br />

Xosophara sp.<br />

Neolieftinckana fuscata<br />

Rentinusdilatatu s<br />

Desudaba danae<br />

Saiva nodata<br />

Pyrops intricata<br />

Penthicodes sp.2<br />

Anecphora torrida<br />

Druentia sicca<br />

Kalidasa lanata<br />

undet. <strong>Fulgoridae</strong><br />

Poblicia fuliginosa<br />

Poblicia fuliginosa<br />

Lystra lanata<br />

Alphina glauca<br />

Alphina glauca (2)<br />

Scaralis sp.1<br />

Cyrpoptus suavis<br />

Amerzanna peruana<br />

Amycle vernalis<br />

Flatolystra sp.1<br />

Weyrauchia fulminea<br />

Episciusguianensis<br />

Episciusboliviana<br />

Enchophora rosacea<br />

Enchophora stillifera<br />

Odontoptera carrenoi<br />

Odontoptera toulgoeti<br />

Diareusa annularis<br />

Aphrodisias cacica<br />

Diareusa imitatrix<br />

Cathedra serrata<br />

Phrictus tripartitus<br />

Phrictusquinquepartitus<br />

Phrictus regalis<br />

Fulgora laternaria<br />

Enchophora sanguinea<br />

Echetra modesta<br />

Aracynthus sanguineus<br />

Neocynthus sp.<br />

Stalubra brunnea<br />

Enhydria tessellata<br />

? -proteobacteria<br />

? Zanna sp.1<br />

? Xosophara sp.<br />

? Neolieftinckana fuscata<br />

? Druentia sicca<br />

? Echetra nebulosa<br />

? Echetra modesta<br />

? Lystra lanata<br />

? Copidocephala viridiguttata<br />

? Enhydria tessellata<br />

? Cathedra serrata<br />

? Phrictus tripartitus<br />

? Phrictus regalis<br />

? Diareusa imitatrix<br />

? Diareusa annularis<br />

? Cyrpoptus suavis<br />

? Paralystra florenciana<br />

? Flatolystra distincta<br />

? Scaralis sp.2<br />

? Scaralis sp.3<br />

? Aracynthus sanguineus


Other Lineages?<br />

(Moran et al., 2005)


Sulcia in Delphacidae?<br />

loss of<br />

Sulcia<br />

(Buchner, 1965)


New Rentinus species<br />

Collected by Simon lab -- Australia

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