A Phylogenetic Investigation of Hemiptera), with Emphasis...

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







Sinuala tuberculata















Phrictus regalis


Artacie dufouri




Penthicodes sp.1








Fulgoridae not monophyletic with respect to Dictyopharidae

Zanna

Dictyopharids



Fulgoridae not monophyletic with respect to Dictyopharidae

CixiidaeDelphacidae

Dictyopharidae

Fulgoridae





Copidocephala viridiguttata


Eurinopsyche obscurata

Neolieftinckana fuscataXosophara sp.



Echetra modesta

Aphrodisias cacica


Copidocephala guttataSinuala tuberculata














Phrictus regalis


Artacie dufouri


Rentinus dilatatus

Birdantis bloteiDesudaba danae


Penthicodes sp.1



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





Copidocephala viridiguttata


Eurinopsyche obscurata

Neolieftinckana fuscataXosophara sp.



Echetra modesta

Aphrodisias cacica


Copidocephala guttataSinuala tuberculata














Phrictus regalis


Artacie dufouri


Rentinus dilatatus

Birdantis bloteiDesudaba danae


Penthicodes sp.1



Saiva nodata

CixiidaeDelphacidae

Dictyopharidae

Echetra modesta

Aphrodisias cacica







Sinuala tuberculata















Phrictus regalis


Artacie dufouri




Penthicodes sp.1






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


Cyrpoptus suavisAmycle vernalis

Calyptoproctus sp.Paralystra florenciana

Amerzanna peruana

Flatolystra distinctaFlatolystra verrucosa

Enhydria tessellataDilobura conspurcata



Alphina glaucaScaralis sp.1

Scaralis sp.3

Scaralis sp.2Poblicia fuliginosa



Sinuala tuberculataAphrodisias cacica

Neocynthus crassusNeocynthus sp.

Episcius bolivianusAracynthus sanguineus

Echetra nebulosa

Weyrauchia peruvianaEpiscius guianensis

Echetra modesta

Pterodictya reticularisPhenax variegata


Phrictus regalis


Artacie dufourii



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.3

Rhynchomitra microrhinaHyalodictyon sp.

Dictyophara sp.1Dictyophara sp.2


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


Odontopera carrenoiOdontoptera toulgoeti

Copidocephala guttataCopidocephala viridiguttataDiareusa imitatrixDiareusa annularisAphrodisias cacicaSinuala tuberculata

EpisciusguianensisWeyrauchia fulminea

Episciusboliviana


Aracynthus sanguineus

Echetra modestaEchetra nebulosaCyrpoptus suavisAmycle vernalis

Calyptoproctussp.Paralystra florenciana

Amerzanna peruana


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




Artacie dufouri


Zanna sp.3

Neolieftinckana fuscataBirdantis bloteiRentinusdilatatusEurinopsyche obscurataDesudaba danae

Stalubra brunneaEnhydria tessellata


Saiva nodata


Penthicodes farinosaAnecphora torrida

Druentia siccaundet. Fulgoridae

Kalidasa lanata

Xosophara sp.


Scolops sulcipesRhaphiophora sp.


Lappida sp.

Bothriocera eboreaOecleus sp.Tropidocephala sp.

Africa + Asia

NW

Australia+ PNG


Odontopera carrenoiOdontoptera toulgoeti

Copidocephala guttataCopidocephala viridiguttataDiareusa imitatrixDiareusa annularisAphrodisias cacicaSinuala tuberculata

EpisciusguianensisWeyrauchia fulminea

Episciusboliviana


Aracynthus sanguineus

Echetra modestaEchetra nebulosaCyrpoptus suavisAmycle vernalis

Calyptoproctussp.Paralystra florenciana

Amerzanna peruana


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




Artacie dufouri


Zanna sp.3

Neolieftinckana fuscataBirdantis bloteiRentinusdilatatusEurinopsyche obscurataDesudaba danae

Stalubra brunneaEnhydria tessellata


Saiva nodata


Penthicodes farinosaAnecphora torrida

Druentia siccaundet. Fulgoridae

Kalidasa lanata

Xosophara sp.


Scolops sulcipesRhaphiophora 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







Sinuala tuberculata















Phrictus regalis


Artacie dufouri




Penthicodes sp.1






Increased sampling of Zanna

CixiidaeDelphacidae

Dictyopharidae

Echetra modesta

Aphrodisias cacica







Sinuala tuberculata















Phrictus regalis


Artacie dufouri




Penthicodes sp.1






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


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

A Phylogenetic Investigation of Hemiptera), with Emphasis...

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