A Phylogenetic Investigation of the Planthopper Superfamily Fulgoroidea (Insecta: Hemiptera), with Emphasis on the Family Fulgoridae DEFENDED!

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1 A Phylogenetic Investigation of the Planthopper Superfamily Fulgoroidea (Insecta: Hemiptera), with Emphasis on the Family Fulgoridae DEFENDED! Julie Urban University at Albany & New York State Museum

2 First Phylogeny of Fulgoridae -- molecular data (Urban & Cryan, accepted MP&E)

3 Taxonomic sampling: Fulgoridae -- molecular Ingroup: 69 taxa in 46 genera -- representatives of 8 of 8 subfamilies, 10 of 11 tribes Outgroups: Dictyopharidae (7) Cixiidae (2), Delphacidae (1) Molecular data -- ~ 5000 bp 18S rdna 28S rdna Histone 3 Wingless COI ~1,900 bp ~2,200 bp ~ 350 bp ~ 350 bp ~ 500 bp

4 Bothriocera eborea Oecleus sp. Tropidocephala sp. Raphiophora sp. Dictyophara sp. undet. Dictyopharidae Penthicodes sp.1 Penthicodes sp.2 Rentinus dilatatus Birdantis blotei Odontoptera carrenoi Stalubra sp. Fulgora sp.2 Fulgora sp.1 Phrictus quinquepartitus Artacie dufouri Calyptoproctus sp. Episcius boliviana Episcius guianensis Neocynthus crassus var. 1 Neocynthus crassus var. 2 Flatolystra sp.1 Flatolystra sp.2 Enhydria sp. MP analysis equally parsimonious topologies -- robust to different weightings

5 Bothriocera eborea Oecleus sp. Tropidocephala sp. Raphiophora sp. Dictyophara sp. undet. Dictyopharidae Penthicodes sp.1 Penthicodes sp.2 Rentinus dilatatus Birdantis blotei Odontoptera carrenoi Stalubra sp. Fulgora sp.2 Fulgora sp.1 Phrictus quinquepartitus Artacie dufouri Calyptoproctus sp. Episcius boliviana Episcius guianensis Neocynthus crassus var. 1 Neocynthus crassus var. 2 Flatolystra sp.1 Flatolystra sp.2 Enhydria sp. Cixiidae Delphacidae Dictyopharidae MP analysis equally parsimonious topologies -- robust to different weightings Fulgoridae not monophyletic with respect to Dictyopharidae Dictyopharids Zanna

6 Fulgoridae Bothriocera eborea Oecleus sp. Tropidocephala sp. Raphiophora sp. Dictyophara sp. undet. Dictyopharidae Penthicodes sp.1 Penthicodes sp.2 Rentinus dilatatus Birdantis blotei Odontoptera carrenoi Stalubra sp. Fulgora sp.2 Fulgora sp.1 Phrictus quinquepartitus Artacie dufouri Calyptoproctus sp. Episcius boliviana Episcius guianensis Neocynthus crassus var. 1 Neocynthus crassus var. 2 Flatolystra sp.1 Flatolystra sp.2 Enhydria sp. Cixiidae Delphacidae Dictyopharidae MP analysis equally parsimonious topologies -- robust to different weightings Fulgoridae not monophyletic with respect to Dictyopharidae

7 Fulgoridae Bothriocera eborea Oecleus sp. Tropidocephala sp. Raphiophora sp. Dictyophara sp. undet. Dictyopharidae Penthicodes sp.1 Penthicodes sp.2 Rentinus dilatatus Birdantis blotei Odontoptera carrenoi Stalubra sp. Fulgora sp.2 Fulgora sp.1 Phrictus quinquepartitus Artacie dufouri Calyptoproctus sp. Episcius boliviana Episcius guianensis Neocynthus crassus var. 1 Neocynthus crassus var. 2 Flatolystra sp.1 Flatolystra sp.2 Enhydria sp. Cixiidae Delphacidae Dictyopharidae Tests of classification: -- Reject monophyly of family -- Reject monophyly of 5 of 7 tested subfamilies -- Reject monophyly of 5 of 6 tested tribes

8 NW Bothriocera eborea Oecleus sp. Tropidocephala sp. Raphiophora sp. Dictyophara sp. undet. Dictyopharidae Penthicodes sp.1 Penthicodes sp.2 Rentinus dilatatus Birdantis blotei Odontoptera carrenoi Stalubra sp. Fulgora sp.2 Fulgora sp.1 Phrictus quinquepartitus Artacie dufouri Calyptoproctus sp. Episcius boliviana Episcius guianensis Neocynthus crassus var. 1 Neocynthus crassus var. 2 Flatolystra sp.1 Flatolystra sp.2 Enhydria sp. Cixiidae Delphacidae Dictyopharidae Nearctic Nearctic Biogeography: -- Old World origin -- Monophyletic New World (colonization of Neotropics) -- Multiple dispersal events into Nearctic Mapping Head Process: losses of head process across fulgorid evolution

9 Pyrops delessertii Anecphora sp. Penthicodes farinosa Penthicodes atomaria Birdantis blötei Desudaba danaë Eurinopsyche sp. 25 Odontoptera carrenoi Artacie dufourii Phrictus quinquepartitus Fulgora castresii Fulgora cearensis Stalubra brunnea Episcius bolivianus Neocynthus crassus Neocynthus sp Weyrauchia peruviana Episcius guianensis Calyptoproctus sp Flatolystra distincta 59 Flatolystra verrucosa 61 Enhydria tessellata Zanna Raphiophora sp. LappidaRaphiophoraScolops 8 Dictyophara sp.1 9 Dictyophara sp.2 DictyopharaRhynchomitra Hyalodictyon sp.1 XosopharaPyrops Saiva AnecphoraPenthicodesDruentiaUndetermined Kalidasa Neolieftinckana BirdantisDesudabaEurinopsyche Cathedra Odontoptera EnchophoraArtacie Fulgora Phrictus Stalubra Pterodictya Phenax Echetra Episcius AracynthusNeocynthus DiareusaSinualaAphrodisias Copidocephala EchetraWeyrauchiaEpiscius CyrpoptusAmycle AmerzannaCalyptoproctus Paralystra Poblicia Flatolystra EnhydriaDilobura Lystra Learcha Scaralis Matacosa Alphina

10 Fulgoridae -- molecules + morphology Taxonomic sampling 69 ingroup taxa + 10 outgroups Morphological data characters Characters and states developed from literature on Fulgoridae and other planthopper families Molecular data -- ~ 5000 bp 18S rdna 28S rdna Histone 3 Wingless COI ~1,900 bp ~2,200 bp ~ 350bp ~ 350bp ~ 500bp

11 Australia + PNG Africa + Asia NW Bothriocera eborea Oecleus sp. Tropidocephala sp. Rhaphiophora sp. Dictyophara sp.1 Dictyophara sp.2 Birdantis blotei Rentinusdilatatus Penthicodes farinosa Penthicodes atomaria Stalubra brunnea Enhydria tessellata Odontopera carrenoi Fulgora cearensis Fulgora castresii Phrictusquinquepartitus Artacie dufouri Episciusboliviana Episciusguianensis Neocynthus crassus Neocynthus sp. Calyptoproctussp. Flatolystra distincta Flatolystra verrucosa Bayesian analysis Zanna placed outside of Fulgoridae Monophyletic New World lineage 2 Old World lineages?

12 Australia + PNG Africa + Asia NW Bothriocera eborea Oecleus sp. Tropidocephala sp. Rhaphiophora sp. Dictyophara sp.1 Dictyophara sp.2 Birdantis blotei Rentinusdilatatus Penthicodes farinosa Penthicodes atomaria Stalubra brunnea Enhydria tessellata Odontopera carrenoi Fulgora cearensis Fulgora castresii Phrictusquinquepartitus Artacie dufouri Episciusboliviana Episciusguianensis Neocynthus crassus Neocynthus sp. Calyptoproctussp. Flatolystra distincta Flatolystra verrucosa

13 Fulgoridae II (Urban & Cryan, in progress) Increasing taxonomic sampling 13 genera new to data set 40 species new to data set Reworking some morphological characters Objective: Basis for revised classification

14 Bothriocera eborea Oecleus sp. Tropidocephala sp. Raphiophora sp. Dictyophara sp. undet. Dictyopharidae Penthicodes sp.1 Penthicodes sp.2 Rentinus dilatatus Birdantis blotei Odontoptera carrenoi Stalubra sp. Fulgora sp.2 Fulgora sp.1 Phrictus quinquepartitus Artacie dufouri Calyptoproctus sp. Episcius boliviana Episcius guianensis Neocynthus crassus var. 1 Neocynthus crassus var. 2 Flatolystra sp.1 Flatolystra sp.2 Enhydria sp. Cixiidae Delphacidae Dictyopharidae Increased sampling of Zanna

15 Bothriocera eborea Oecleus sp. Tropidocephala sp. Raphiophora sp. Dictyophara sp. undet. Dictyopharidae Penthicodes sp.1 Penthicodes sp.2 Rentinus dilatatus Birdantis blotei Odontoptera carrenoi Stalubra sp. Fulgora sp.2 Fulgora sp.1 Phrictus quinquepartitus Artacie dufouri Calyptoproctus sp. Episcius boliviana Episcius guianensis Neocynthus crassus var. 1 Neocynthus crassus var. 2 Flatolystra sp.1 Flatolystra sp.2 Enhydria sp. Cixiidae Delphacidae Dictyopharidae Increased sampling of Zanna Increased sampling within Dictyopharidae Increased Old World sampling (3 genera) Increased sampling within genera

16 Endosymbionts Within Fulgoridae and Dictyopharidae Sulcia muelleri NW Aphrophora quadrinotata Clastoptera arizonana Scolops sp. Dictyophara sp.1 Dictyophara sp.2 Rhaphiophora sp. Zanna sp. Rentinusdilatatu s Penthicodes sp.2 (2) Flatolystra sp.1 Episciusguianensis Episciusboliviana Odontoptera carrenoi Phrictusquinquepartitus Enchophora sanguinea Neocynthus sp. Stalubra brunnea Enhydria tessellata? -proteobacteria????????? Enhydria tessellata???????? Flatolystra distincta???

17 Other Lineages? (Moran et al., 2005)

18 Sulcia in Delphacidae? (Buchner, 1965) loss of Sulcia

19 New Rentinus species Collected by Simon lab -- Australia