Taxonomy,Phylogeny And Biotope Characteristics Of Major Groups Of The Oriental And Palaearctic Issidae (Hemiptera:Fulgoroidea)

This research is on the taxonomy,phylogeny and biotope characteristics of major groups of the Oriental and Palaearctic Issidae(Hemiptera: Fulgoroidea).In the taxonomic section,Issidae specimens of three tribes,90 genera,242 species(32genera and 92 species from Northwest A&F University(NWAFU)and 67 genera and 151 species from the Muséum National d'Histoire Naturelle(MNHN))were examined,of which seven new genera and 16 new species of Issidae are described and two new combinations are proposed.Photos of the external morphology along with male and female genitalia of these species are illustrated.Some species of Issidae were revised.New taxa and new combinations are as follows(the new names of the taxa mentioned below will only be valid after publication and registration in Zoobank):New genera: Longieusarima gen.nov.,Brevsarima gen.nov.,Kodaianellissus gen.nov.,Homotetricodes gen.nov.,Neocoruncanius gen.nov.,Tetricissus gen.nov.,Nongelastyrella gen.nov.New species: Macroduruma simipertinax sp.nov.,Longieusarima lunulia sp.nov.,Brevsarima brunnea sp.nov.,Kodaianellissus intorqueus sp.nov.,Homotetricodes helvimacula sp.nov.,Neocoruncanius tenuvirgus sp.nov.,Eusarima fuscvincla sp.nov.,Eusarima granula sp.nov.,Eusarima camptoocula sp.nov.,Eusarima helvifacia sp.nov.,Eusarima rectiocula sp.nov.,Eusarima sancta sp.nov.,Euxaldar lenis sp.nov.,Nongelastyrella sufflava sp.nov.,Tetrica popula sp.nov.,Neodurium chenissum sp.nov.New combination: Bruneastrum quinquemaculatus(Che,Zhang & Wang,2007)comb.nov.(= Gergithus quinquemaculatus Che,Zhang & Wang,2007),Tetricissus philo(Fennah,1978)comb.nov.(= Tetrica philo Fennah,1978).Taxa published:Tetricodissus Wang,Bourgoin & Zhang,2015;Tetricodissus pandlineus Wang,Bourgoin & Zhang,2015;Connelicita Wang & Bourgoin,2015;Connelicita backyensis Stroi?ski & Bourgoin,2015;Connelicita haiphongensis Wang & Zhang,2015;Connelicita lungchowensis(Chou & Lu,1977).The type specimens of new species are deposited in the Entomological Museum,Northwest A&F University,Yangling,Shaanxi,China(NWAFU)or Muséum National d'Histoire Naturelle,Paris,France(MNHN).In the phylogeny work,a molecular phylogeny of the planthopper family Issidae(Hemiptera,Fulgoroidea)of 50 genera 79 species with whole length of 4702 bp is constructed,based on four genes: nuclear genes 18 S and D3–D7 regions of 28 S,mitochondrial genes COXI and Cytb.For sampling,this study not only took into account the major lineages of Issidae s.s.to test the relationships inside the family,but also included the major lineages(Caliscelidae,Acanaloniidae,Tonginae,Trienopinae,Bladininae)which were recently excluded from the Issidae.This analysis was conducted using both Maximum likelihood and Bayesian inference analyses.In our results,all previously-excluded taxa are confirmed as not being part of the Issidae family.However for the first time,all analyses depict several fully supported groups,which challenge the recent proposed "modern classification" of the family.Based on this molecular phylogenetic study,a new classification of the family Issidae is proposed which now divides the Issidae into 3 subfamilies and 7 tribes,among them:1)Neotropical Thioniini is re-established as an independent sister lineage to all other Issidae;2)Palaearctic Issidae are weakly supported(UFB=62)as a monophyletic lineage Issinae including two tribes: Issini and Hysteropterini(both strongly supported with UFB=100);3)All Oriental Issidae cluster together to form a strong monophyletic subfamily group(UFB=99)Hemisphaeriinae which includes four tribes: Kodaianellini(UFB=99),Sarimini(UFB=96),Parahiraciini(UFB=100),and Hemisphaeriini(UFB=100),the latter including two sub-tribes: Mongolianina and Hemisphaeriina.4)The Neotropical lineage including the genus Picumna is regarded as incertae sedis within the Issidae.The classification system of Issidae proposed by this study:Family Issidae Spinola,1839-Thioniinae Melichar,1906Thioniini Melichar,1906-Issinae Spinola,1839Issini Spinola,1839Hysteropterini Melichar,1906-Hemisphaeriinae Melichar,1906Kodaianellini Wang,Zhang & Bourgoin,2016Sarimini Wang,Zhang & Bourgoin,2016Parahiraciini Cheng & Yang,1991Hemisphaeriini Melichar,1906Mongolianina Wang,Zhang & Bourgoin,2016Hemisphaeriina Melichar,1906Biotope characteristics of the major groups were investigated based on the main monophyletic clades obtained in the molecular phylogeny tree,using 3556 geographical distribution data and nine bioclimatic variables related to temperature and humidity,to model the niche of the Oriental and Palaearctic Issidae.The main results are as follows:1)From a biogeographical view,the Palaearctic Issini are mainly distributed in the northwestern of Europe.However,the Palaearctic Hysteropterini has a unique distribution,its biogeography showing a horizontal distribution from the west to east,mainly inland.The Oriental Issidae are mainly located in the south of China.2)Palaearctic Issidae prefer a lower temperature and cooler climate than do Oriental Issidae.The Palaearctic Issidae usually live in areas with an annual mean temperature ranging from 1°C to 13°C;however,most of the Oriental Issidae favor a warm habitat in a area with annual mean temperature ranging from 15°C to 23°C.3)Palaearctic Issini and all tribes of Oriental Issidae prefer a humid environment with abundant rainwater(annual precipitation of about 700 mm~2500 mm);only the Palaearctic Hysteropterini live in semi-humid regions(annual precipitation of about 200 mm~500 mm).4)Under more extreme conditions,Issidae are very sensitive to excessive high temperature and this is one of the main factors limiting the range of the different main groups of Issidae.In addition,Issidae are also susceptible to drought,the precipitation of the driest quarter also acting as one of the main factors.5)There is no evidence that large-scale climatic factors caused family level evolutionary diversification of the Oriental Issidae(four tribes),but climatic factors had a very important role in the diversity patterns of the Palaearctic Issidae(Issini and Hysteropterini).