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
First Phylogeny of Fulgoridae -- molecular data (Urban & Cryan, accepted MP&E)
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
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 -- 19 equally parsimonious topologies -- robust to different weightings
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 -- 19 equally parsimonious topologies -- robust to different weightings Fulgoridae not monophyletic with respect to Dictyopharidae Dictyopharids Zanna
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 -- 19 equally parsimonious topologies -- robust to different weightings Fulgoridae not monophyletic with respect to Dictyopharidae
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
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: -- 4-7 losses of head process across fulgorid evolution
1 2 5 15 11 14 18 19 6 12 13 16 17 7 10 Pyrops delessertii Anecphora sp. Penthicodes farinosa Penthicodes atomaria 21 20 Birdantis blötei Desudaba danaë 22 23 Eurinopsyche sp. 25 Odontoptera carrenoi 30 31 29 33 32 28 Artacie dufourii 27 34 35 Phrictus quinquepartitus 26 36 Fulgora castresii 37 38 24 Fulgora cearensis Stalubra brunnea 41 43 44 Episcius bolivianus 40 45 Neocynthus crassus Neocynthus sp. 48 42 47 46 49 50 39 51 52 Weyrauchia peruviana Episcius guianensis 55 54 56 57 Calyptoproctus sp. 53 60 Flatolystra distincta 59 Flatolystra verrucosa 61 Enhydria tessellata 58 63 62 64 65 67 66 69 68 3 4 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
Fulgoridae -- molecules + morphology Taxonomic sampling 69 ingroup taxa + 10 outgroups Morphological data -- 110 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
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?
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
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
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
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
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???
Other Lineages? (Moran et al., 2005)
Sulcia in Delphacidae? (Buchner, 1965) loss of Sulcia
New Rentinus species Collected by Simon lab -- Australia