A Phylogenetic Investigation of the Planthopper Superfamily
Fulgoroidea (Insecta: Hemiptera), with Emphasis on
the Family Fulgoridae
Julie UrbanUniversity at Albany &
New York State Museum
DEFENDED!
First Phylogeny of Fulgoridae --molecular data (Urban & Cryan, accepted MP&E)
Fulgoridae -- molecular
Molecular data -- ~ 5000 bp
18S rDNA ~1,900 bp
28S rDNA ~2,200 bp
Histone 3 ~ 350 bp
Taxonomic sampling:
Ingroup: 69 taxa in 46 genera-- representatives of 8 of 8 subfamilies, 10 of 11 tribes
Outgroups:
Wingless ~ 350 bp
Dictyopharidae (7)
Cixiidae (2), Delphacidae (1)
COI ~ 500 bp
Echetra modesta
Aphrodisias cacica
Dictyophara sp.undet. Dictyopharidae
Scolops sulcipesRaphiophora sp.
Rhynchomitra microrhinaHyalodictyon sp.Lappida sp.
Diareusa imitatrixDiareusa annularis
Neocynthus crassus var. 1Neocynthus crassus var. 2
Copidocephala guttataCopidocephala viridiguttata
Sinuala tuberculata
Episcius bolivianaEpiscius guianensisWeyrauchia fulminea
Aracynthus sanguineusEchetra nebulosa
Cyrpoptus suavisAmycle vernalisAmerzanna peruana
Calyptoproctus sp.Paralystra florencianaPterodictya reticularisPhenax variegata
Matacosa miscellaScaralis semilimpida
Lystra lanataLearcha sponsa
Poblicia fuliginosaAlphina glaucaScaralis sp.1Scaralis sp.2Scaralis sp.3
Flatolystra sp.1Flatolystra sp.2
Enhydria sp.Dilobura conspurcata
Fulgora sp.1Fulgora laternaria
Fulgora sp.2Cathedra serrataStalubra sp.
Enchophora sanguinea var.1Enchophora sanguinea var.2
Enchophora nigromaculata
Phrictus tripartitusPhrictus quinquepartitus
Phrictus regalis
Enchophora rosaceaEnchophora stillifera
Artacie dufouri
Odontoptera carrenoiOdontoptera toulgoeti
Rentinus dilatatusEurinopsyche obscurataBirdantis bloteiDesudaba danae
Penthicodes pulchellaPenthicodes sp.2
Penthicodes sp.1
Druentia siccaundet. FulgoridaeAnecphora torridaKalidasa lanata
Pyrops sp.Pyrops intricata
Saiva nodataNeolieftinckana fuscataXosophara sp.
Zanna sp.2Zanna sp.3
Zanna sp.1Tropidocephala sp.Oecleus sp.Bothriocera eborea
MP analysis -- 19 equally parsimonious
topologies-- robust to different weightings
CixiidaeDelphacidae
Dictyopharidae
Echetra modesta
Aphrodisias cacica
Dictyophara sp.undet. Dictyopharidae
Scolops sulcipesRaphiophora sp.
Rhynchomitra microrhinaHyalodictyon sp.Lappida sp.
Diareusa imitatrixDiareusa annularis
Neocynthus crassus var. 1Neocynthus crassus var. 2
Copidocephala guttataCopidocephala viridiguttata
Sinuala tuberculata
Episcius bolivianaEpiscius guianensisWeyrauchia fulminea
Aracynthus sanguineusEchetra nebulosa
Cyrpoptus suavisAmycle vernalisAmerzanna peruana
Calyptoproctus sp.Paralystra florencianaPterodictya reticularisPhenax variegata
Matacosa miscellaScaralis semilimpida
Lystra lanataLearcha sponsa
Poblicia fuliginosaAlphina glaucaScaralis sp.1Scaralis sp.2Scaralis sp.3
Flatolystra sp.1Flatolystra sp.2
Enhydria sp.Dilobura conspurcata
Fulgora sp.1Fulgora laternaria
Fulgora sp.2Cathedra serrataStalubra sp.
Enchophora sanguinea var.1Enchophora sanguinea var.2
Enchophora nigromaculata
Phrictus tripartitusPhrictus quinquepartitus
Phrictus regalis
Enchophora rosaceaEnchophora stillifera
Artacie dufouri
Odontoptera carrenoiOdontoptera toulgoeti
Rentinus dilatatusEurinopsyche obscurataBirdantis bloteiDesudaba danae
Penthicodes pulchellaPenthicodes sp.2
Penthicodes sp.1
Druentia siccaundet. FulgoridaeAnecphora torridaKalidasa lanata
Pyrops sp.Pyrops intricata
Saiva nodataNeolieftinckana fuscataXosophara sp.
Zanna sp.2Zanna sp.3
Zanna sp.1Tropidocephala sp.Oecleus sp.Bothriocera eborea
MP analysis -- 19 equally parsimonious
topologies-- robust to different weightings
Fulgoridae not monophyletic with respect to Dictyopharidae
Zanna
Dictyopharids
MP analysis -- 19 equally parsimonious
topologies-- robust to different weightings
Fulgoridae not monophyletic with respect to Dictyopharidae
CixiidaeDelphacidae
Dictyopharidae
Fulgoridae
Dictyophara sp.undet. Dictyopharidae
Scolops sulcipesRaphiophora sp.
Rhynchomitra microrhinaHyalodictyon sp.Lappida sp.
Neocynthus crassus var. 1Neocynthus crassus var. 2
Copidocephala viridiguttata
Enchophora nigromaculata
Eurinopsyche obscurata
Neolieftinckana fuscataXosophara sp.
Zanna sp.2Zanna sp.3
Zanna sp.1Tropidocephala sp.Oecleus sp.Bothriocera eborea
Echetra modesta
Aphrodisias cacica
Diareusa imitatrixDiareusa annularis
Copidocephala guttataSinuala tuberculata
Episcius bolivianaEpiscius guianensisWeyrauchia fulminea
Aracynthus sanguineusEchetra nebulosa
Cyrpoptus suavisAmycle vernalisAmerzanna peruana
Calyptoproctus sp.Paralystra florencianaPterodictya reticularisPhenax variegata
Matacosa miscellaScaralis semilimpida
Lystra lanataLearcha sponsa
Poblicia fuliginosaAlphina glaucaScaralis sp.1Scaralis sp.2Scaralis sp.3
Flatolystra sp.1Flatolystra sp.2
Enhydria sp.Dilobura conspurcata
Fulgora sp.1Fulgora laternaria
Fulgora sp.2Cathedra serrataStalubra sp.
Enchophora sanguinea var.1Enchophora sanguinea var.2
Phrictus tripartitusPhrictus quinquepartitus
Phrictus regalis
Enchophora rosaceaEnchophora stillifera
Artacie dufouri
Odontoptera carrenoiOdontoptera toulgoeti
Rentinus dilatatus
Birdantis bloteiDesudaba danae
Penthicodes pulchellaPenthicodes sp.2
Penthicodes sp.1
Druentia siccaundet. FulgoridaeAnecphora torridaKalidasa lanata
Pyrops sp.Pyrops intricata
Saiva nodata
Tests of classification:
-- Reject monophyly of family-- Reject monophyly of 5 of 7 tested subfamilies-- Reject monophyly of 5 of 6 tested tribes
CixiidaeDelphacidae
Dictyopharidae
Fulgoridae
Dictyophara sp.undet. Dictyopharidae
Scolops sulcipesRaphiophora sp.
Rhynchomitra microrhinaHyalodictyon sp.Lappida sp.
Neocynthus crassus var. 1Neocynthus crassus var. 2
Copidocephala viridiguttata
Enchophora nigromaculata
Eurinopsyche obscurata
Neolieftinckana fuscataXosophara sp.
Zanna sp.2Zanna sp.3
Zanna sp.1Tropidocephala sp.Oecleus sp.Bothriocera eborea
Echetra modesta
Aphrodisias cacica
Diareusa imitatrixDiareusa annularis
Copidocephala guttataSinuala tuberculata
Episcius bolivianaEpiscius guianensisWeyrauchia fulminea
Aracynthus sanguineusEchetra nebulosa
Cyrpoptus suavisAmycle vernalisAmerzanna peruana
Calyptoproctus sp.Paralystra florencianaPterodictya reticularisPhenax variegata
Matacosa miscellaScaralis semilimpida
Lystra lanataLearcha sponsa
Poblicia fuliginosaAlphina glaucaScaralis sp.1Scaralis sp.2Scaralis sp.3
Flatolystra sp.1Flatolystra sp.2
Enhydria sp.Dilobura conspurcata
Fulgora sp.1Fulgora laternaria
Fulgora sp.2Cathedra serrataStalubra sp.
Enchophora sanguinea var.1Enchophora sanguinea var.2
Phrictus tripartitusPhrictus quinquepartitus
Phrictus regalis
Enchophora rosaceaEnchophora stillifera
Artacie dufouri
Odontoptera carrenoiOdontoptera toulgoeti
Rentinus dilatatus
Birdantis bloteiDesudaba danae
Penthicodes pulchellaPenthicodes sp.2
Penthicodes sp.1
Druentia siccaundet. FulgoridaeAnecphora torridaKalidasa lanata
Pyrops sp.Pyrops intricata
Saiva nodata
CixiidaeDelphacidae
Dictyopharidae
Echetra modesta
Aphrodisias cacica
Dictyophara sp.undet. Dictyopharidae
Scolops sulcipesRaphiophora sp.
Rhynchomitra microrhinaHyalodictyon sp.Lappida sp.
Diareusa imitatrixDiareusa annularis
Neocynthus crassus var. 1Neocynthus crassus var. 2
Copidocephala guttataCopidocephala viridiguttata
Sinuala tuberculata
Episcius bolivianaEpiscius guianensisWeyrauchia fulminea
Aracynthus sanguineusEchetra nebulosa
Cyrpoptus suavisAmycle vernalisAmerzanna peruana
Calyptoproctus sp.Paralystra florencianaPterodictya reticularisPhenax variegata
Matacosa miscellaScaralis semilimpida
Lystra lanataLearcha sponsa
Poblicia fuliginosaAlphina glaucaScaralis sp.1Scaralis sp.2Scaralis sp.3
Flatolystra sp.1Flatolystra sp.2
Enhydria sp.Dilobura conspurcata
Fulgora sp.1Fulgora laternaria
Fulgora sp.2Cathedra serrataStalubra sp.
Enchophora sanguinea var.1Enchophora sanguinea var.2
Enchophora nigromaculata
Phrictus tripartitusPhrictus quinquepartitus
Phrictus regalis
Enchophora rosaceaEnchophora stillifera
Artacie dufouri
Odontoptera carrenoiOdontoptera toulgoeti
Rentinus dilatatusEurinopsyche obscurataBirdantis bloteiDesudaba danae
Penthicodes pulchellaPenthicodes sp.2
Penthicodes sp.1
Druentia siccaundet. FulgoridaeAnecphora torridaKalidasa lanata
Pyrops sp.Pyrops intricata
Saiva nodataNeolieftinckana fuscataXosophara sp.
Zanna sp.2Zanna sp.3
Zanna sp.1Tropidocephala sp.Oecleus sp.Bothriocera eborea
NW
Biogeography:
-- Old World origin-- Monophyletic New World (colonization of Neotropics)-- Multiple dispersal events into Nearctic
Nearctic
Nearctic
Mapping Head Process:
-- 4-7 losses of head process across fulgorid evolution
Pyrops sp.Pyrops delessertii
Saiva nodata
Odontoptera carrenoiOdontoptera toulgoeti
Cyrpoptus suavisAmycle vernalis
Calyptoproctus sp.Paralystra florenciana
Amerzanna peruana
Flatolystra distinctaFlatolystra verrucosa
Enhydria tessellataDilobura conspurcata
Lystra lanataLearcha sponsa
Matacosa miscellaScaralis semilimpida
Alphina glaucaScaralis sp.1
Scaralis sp.3
Scaralis sp.2Poblicia fuliginosa
Copidocephala guttataCopidocephala viridiguttata
Diareusa imitatrixDiareusa annularis
Sinuala tuberculataAphrodisias cacica
Neocynthus crassusNeocynthus sp.
Episcius bolivianusAracynthus sanguineus
Echetra nebulosa
Weyrauchia peruvianaEpiscius guianensis
Echetra modesta
Pterodictya reticularisPhenax variegata
Phrictus tripartitusPhrictus quinquepartitus
Phrictus regalis
Enchophora rosaceaEnchophora stillifera
Artacie dufourii
Enchophora sanguinea var.1Enchophora sanguinea var.2
Enchophora nigromaculata
Fulgora castresiiFulgora laternariaFulgora cearensis
Cathedra serrata
Stalubra brunnea
Neolieftinckana fuscataBirdantis blötei
Eurinopsyche sp.Eurinopsyche obscurata
Desudaba danaë
Penthicodes pulchellaPenthicodes atomaria
Penthicodes farinosaAnecphora sp.
Druentia siccaundet. FulgoridaeKalidasa lanata
Xosophara sp.
Zanna sp.1Zanna sp.2
Zanna sp.3
Rhynchomitra microrhinaHyalodictyon sp.
Dictyophara sp.1Dictyophara sp.2
Scolops sulcipesRaphiophora sp.
Lappida sp.
1
2
5
3
6
78
10
12 13
1516
17
18
2120
22
1411
9
23
4
19
25
24
27
28
3435
2930
3332
31
26 3637
38
39
40
41
42
43 44
45
4647
49
48
50
5152
53
54
57
55
56
58
5961
60
6362
6465
6669
6768
Zanna
LappidaRaphiophoraScolops
Dictyopharasp.1
RhynchomitraHyalodictyon
XosopharaPyrops Saiva
AnecphoraPenthicodesDruentia KalidasaUndetermined
NeolieftinckanaBirdantisDesudabaEurinopsyche
Odontoptera
Enchophora Artacie
Phrictus
StalubraCathedra Fulgora
Pterodictya Phenax
Echetra Episcius AracynthusNeocynthus
DiareusaSinualaAphrodisiasCopidocephala
EchetraWeyrauchiaEpiscius
CyrpoptusAmycle Amerzanna ParalystraCalyptoproctus
Flatolystra DiloburaEnhydria
Lystra Learcha
MatacosaScaralisPoblicia Alphina
Fulgoridae -- molecules + morphology
Morphological data -- 110 characters
Molecular data -- ~ 5000 bp18S rDNA ~1,900 bp
28S rDNA ~2,200 bp
Histone 3 ~ 350bp
Wingless ~ 350bp
COI ~ 500bp
Taxonomic sampling
Characters and states developed from literature on Fulgoridae and other planthopper families
69 ingroup taxa + 10 outgroups
Bayesian analysis
Zanna placed outside of Fulgoridae
Monophyletic New World lineage
2 Old World lineages?
NW
Australia+ PNG
Pterodictya reticularisPhenax variegata
Odontopera carrenoiOdontoptera toulgoeti
Copidocephala guttataCopidocephala viridiguttataDiareusa imitatrixDiareusa annularisAphrodisias cacicaSinuala tuberculata
EpisciusguianensisWeyrauchia fulminea
Episciusboliviana
Neocynthus crassusNeocynthus sp.
Aracynthus sanguineus
Echetra modestaEchetra nebulosaCyrpoptus suavisAmycle vernalis
Calyptoproctussp.Paralystra florenciana
Amerzanna peruana
Flatolystra distinctaFlatolystra verrucosa
Dilobura conspurcata
Scaralis sp.1Scaralis sp.3
Matacosa miscellaScaralis semilimpidaAlphina glauca
Scaralis sp.2
Poblicia fuliginosaLystra lanataLearcha sponsa
Fulgora castresiiFulgora laternaria
Fulgora cearensisCathedra serrata
Phrictus tripartitusPhrictusquinquepartitus
Phrictus regalis
Enchophora sanguinea var.1Enchophora sanguinea var.2
Enchophora nigromaculata
Enchophora rosaceaEnchophora stillifera
Artacie dufouri
Zanna sp.1Zanna sp.2
Zanna sp.3
Neolieftinckana fuscataBirdantis bloteiRentinusdilatatusEurinopsyche obscurataDesudaba danae
Stalubra brunneaEnhydria tessellata
Pyrops sp.Pyrops intricata
Saiva nodata
Penthicodes pulchellaPenthicodes atomaria
Penthicodes farinosaAnecphora torrida
Druentia siccaundet. Fulgoridae
Kalidasa lanata
Xosophara sp.
Dictyophara sp.1Dictyophara sp.2
Scolops sulcipesRhaphiophora sp.
Rhynchomitra microrhinaHyalodictyon sp.
Lappida sp.
Bothriocera eboreaOecleus sp.Tropidocephala sp.
Africa + Asia
NW
Australia+ PNG
Pterodictya reticularisPhenax variegata
Odontopera carrenoiOdontoptera toulgoeti
Copidocephala guttataCopidocephala viridiguttataDiareusa imitatrixDiareusa annularisAphrodisias cacicaSinuala tuberculata
EpisciusguianensisWeyrauchia fulminea
Episciusboliviana
Neocynthus crassusNeocynthus sp.
Aracynthus sanguineus
Echetra modestaEchetra nebulosaCyrpoptus suavisAmycle vernalis
Calyptoproctussp.Paralystra florenciana
Amerzanna peruana
Flatolystra distinctaFlatolystra verrucosa
Dilobura conspurcata
Scaralis sp.1Scaralis sp.3
Matacosa miscellaScaralis semilimpidaAlphina glauca
Scaralis sp.2
Poblicia fuliginosaLystra lanataLearcha sponsa
Fulgora castresiiFulgora laternaria
Fulgora cearensisCathedra serrata
Phrictus tripartitusPhrictusquinquepartitus
Phrictus regalis
Enchophora sanguinea var.1Enchophora sanguinea var.2
Enchophora nigromaculata
Enchophora rosaceaEnchophora stillifera
Artacie dufouri
Zanna sp.1Zanna sp.2
Zanna sp.3
Neolieftinckana fuscataBirdantis bloteiRentinusdilatatusEurinopsyche obscurataDesudaba danae
Stalubra brunneaEnhydria tessellata
Pyrops sp.Pyrops intricata
Saiva nodata
Penthicodes pulchellaPenthicodes atomaria
Penthicodes farinosaAnecphora torrida
Druentia siccaundet. Fulgoridae
Kalidasa lanata
Xosophara sp.
Dictyophara sp.1Dictyophara sp.2
Scolops sulcipesRhaphiophora sp.
Rhynchomitra microrhinaHyalodictyon sp.
Lappida sp.
Bothriocera eboreaOecleus sp.Tropidocephala sp.
Africa + Asia
Fulgoridae II (Urban & Cryan, in progress)
Reworking some morphological characters
Objective: Basis for revised classification
Increasing taxonomic sampling13 genera new to data set
40 species new to data set
CixiidaeDelphacidae
Dictyopharidae
Echetra modesta
Aphrodisias cacica
Dictyophara sp.undet. Dictyopharidae
Scolops sulcipesRaphiophora sp.
Rhynchomitra microrhinaHyalodictyon sp.Lappida sp.
Diareusa imitatrixDiareusa annularis
Neocynthus crassus var. 1Neocynthus crassus var. 2
Copidocephala guttataCopidocephala viridiguttata
Sinuala tuberculata
Episcius bolivianaEpiscius guianensisWeyrauchia fulminea
Aracynthus sanguineusEchetra nebulosa
Cyrpoptus suavisAmycle vernalisAmerzanna peruana
Calyptoproctus sp.Paralystra florencianaPterodictya reticularisPhenax variegata
Matacosa miscellaScaralis semilimpida
Lystra lanataLearcha sponsa
Poblicia fuliginosaAlphina glaucaScaralis sp.1Scaralis sp.2Scaralis sp.3
Flatolystra sp.1Flatolystra sp.2
Enhydria sp.Dilobura conspurcata
Fulgora sp.1Fulgora laternaria
Fulgora sp.2Cathedra serrataStalubra sp.
Enchophora sanguinea var.1Enchophora sanguinea var.2
Enchophora nigromaculata
Phrictus tripartitusPhrictus quinquepartitus
Phrictus regalis
Enchophora rosaceaEnchophora stillifera
Artacie dufouri
Odontoptera carrenoiOdontoptera toulgoeti
Rentinus dilatatusEurinopsyche obscurataBirdantis bloteiDesudaba danae
Penthicodes pulchellaPenthicodes sp.2
Penthicodes sp.1
Druentia siccaundet. FulgoridaeAnecphora torridaKalidasa lanata
Pyrops sp.Pyrops intricata
Saiva nodataNeolieftinckana fuscataXosophara sp.
Zanna sp.2Zanna sp.3
Zanna sp.1Tropidocephala sp.Oecleus sp.Bothriocera eborea
Increased sampling of Zanna
CixiidaeDelphacidae
Dictyopharidae
Echetra modesta
Aphrodisias cacica
Dictyophara sp.undet. Dictyopharidae
Scolops sulcipesRaphiophora sp.
Rhynchomitra microrhinaHyalodictyon sp.Lappida sp.
Diareusa imitatrixDiareusa annularis
Neocynthus crassus var. 1Neocynthus crassus var. 2
Copidocephala guttataCopidocephala viridiguttata
Sinuala tuberculata
Episcius bolivianaEpiscius guianensisWeyrauchia fulminea
Aracynthus sanguineusEchetra nebulosa
Cyrpoptus suavisAmycle vernalisAmerzanna peruana
Calyptoproctus sp.Paralystra florencianaPterodictya reticularisPhenax variegata
Matacosa miscellaScaralis semilimpida
Lystra lanataLearcha sponsa
Poblicia fuliginosaAlphina glaucaScaralis sp.1Scaralis sp.2Scaralis sp.3
Flatolystra sp.1Flatolystra sp.2
Enhydria sp.Dilobura conspurcata
Fulgora sp.1Fulgora laternaria
Fulgora sp.2Cathedra serrataStalubra sp.
Enchophora sanguinea var.1Enchophora sanguinea var.2
Enchophora nigromaculata
Phrictus tripartitusPhrictus quinquepartitus
Phrictus regalis
Enchophora rosaceaEnchophora stillifera
Artacie dufouri
Odontoptera carrenoiOdontoptera toulgoeti
Rentinus dilatatusEurinopsyche obscurataBirdantis bloteiDesudaba danae
Penthicodes pulchellaPenthicodes sp.2
Penthicodes sp.1
Druentia siccaundet. FulgoridaeAnecphora torridaKalidasa lanata
Pyrops sp.Pyrops intricata
Saiva nodataNeolieftinckana fuscataXosophara sp.
Zanna sp.2Zanna sp.3
Zanna sp.1Tropidocephala sp.Oecleus sp.Bothriocera eborea
Increased sampling of Zanna
Increased sampling within Dictyopharidae
Increased Old World sampling (3 genera)
Increased sampling within genera
Within Fulgoridae and Dictyopharidae
Endosymbionts
Sulcia muelleriAphrophora quadrinotataClastoptera arizonanaScolops sulcipesScolops sp. Dictyophara sp.1Dictyophara sp.2Rhaphiophora sp.
Rentinusdilatatu sDesudaba danae
Neolieftinckana fuscata
Saiva nodataPyrops intricataPenthicodes sp.2
Druentia siccaAnecphora torrida
Kalidasa lanataundet. Fulgoridae
Xosophara sp.
Poblicia fuliginosaPoblicia fuliginosaLystra lanataAlphina glaucaAlphina glauca (2)Scaralis sp.1
Odontoptera carrenoiOdontoptera toulgoetiDiareusa annularisAphrodisias cacicaDiareusa imitatrixCathedra serrata
Enchophora sanguineaFulgora laternaria
Phrictus tripartitusPhrictusquinquepartitusPhrictus regalis
Enchophora rosaceaEnchophora stillifera
Enhydria tessellata
Cyrpoptus suavisAmerzanna peruanaAmycle vernalisFlatolystra sp.1
Echetra modesta
Zanna sp.
Weyrauchia fulmineaEpisciusguianensisEpisciusboliviana
Aracynthus sanguineusNeocynthus sp.Stalubra brunnea
NW
? -proteobacteria
? Zanna sp.1? Xosophara sp.? Neolieftinckana fuscata? Druentia sicca? Echetra nebulosa
? Lystra lanata? Copidocephala viridiguttata
? Cathedra serrata? Phrictus tripartitus? Phrictus regalis? Diareusa imitatrix? Diareusa annularis? Cyrpoptus suavis? Paralystra florenciana
? Scaralis sp.3? Aracynthus sanguineus
? Echetra modesta
? Enhydria tessellata
? Scaralis sp.2? Flatolystra distincta
Other Lineages?(Moran et al., 2005)
Sulcia in Delphacidae?
loss ofSulcia
(Buchner, 1965)
New Rentinus speciesCollected by Simon lab -- Australia