Taxonomy And Phylogeny Of The Leafhopper Tribe Paralimnini(Hemiptera:Cicadellidae:Deltocephalinae) From Chinaand Germany

The tribe Paralimnini of the subfamily Deltocephalinae is one of the larger tribes of leafhoppers comprised 139 genera and 931 species in the world.Leafhoppers of this tribe are common in grassland and feed on the phloem of vascular plants,causing mechanical damage on many cash crops,particularly gramineous plants.Many species are also vectors of plant diseases,causing greater economic loss.Species of the tribe Paralimnini are very similar in external morphology and mainly distinguished by the male genitalia.The relationships among genera is unclear,and there has been a lack of research in many geographic regions.China and Germany are biodiversity-rich countries on the east and west sides of the Old World.Prior to the works on the Paralimnini from China and Germany are scattered among Chinese and German literature.Comprehensive systematic studies on the Chinese and German fauna are needed to provide insights into relationships and to identify traits useful for classification and identification.This thesis describes Chinese and German species of the tribe Paralimnini,with emphasis on the tribal phylogeny of 102 species in 51 genera based on 80 morphological characteristics,the nuclear genes 28 S r DNA and H3,and the mitochondrial 16 S r DNA.The leafhopper tribe Paralimnini from China and Germany is reviewed.The Chinese and German fauna includes 98 species of 46 genera,including 1 new genus Brevipenisus gen.nov.,and 9 new species: Arocephalus maxitettix sp.nov.,Brevipenisus petalinus sp.nov.,Diplocolenus gracilis sp.nov.,Destitutus arctus sp.nov.,Sorhoanus splitus sp.nov.,Jilinga taibaiensis sp.nov.,Futusujinus dietrichi sp.nov.,Futusujinus hastatus sp.nov.and Futusujinus truncatus sp.nov.(these last 4 species were published).Seven genera and ten species are recorded for the first time from China and four new synonyms are proposed.Specimens of available species were photographed and illustrated.We also provide a key to genera and species including 67 external morphological illustrations and 86 male genitalia illustrations.The phylogenetic analysis of 69 representative species from 41 genera of Paralimnini was carried out by using Wincladaver 1.00.08 and TNT ver.1.1.based on 80 morphological characters.One strict consensus tree was obtained and indicated the genus Deltocephalus was the outgroup.The results support Paralimnini as a monophyletic group and Parapotes as the most primitive group.All genera including multiple representative species are monophyletic groups such as Psammotettix and Arocephalus.Among them,Futasujinus,Hengchunia,Takagiella and Yanocephalus all belong to the tribe Paralimnini.In addition,six morphological characteristics from females were selected,such as the ratio of the length of the female forewing to abdomen and the shape of female seventh sternite,that provided support at the important nodes which formed the evolutionary direction of the inner group.Therefore,they played an important role in morphological phylogeny and are important morphological features for defining Paralimnini.The nuclear genes 28 S r DNA and H3 and mitochondrial 16 S r DNA were extracted and247 sequences for 52 genera and 102 species were obtained by using RAx ML7.0.3 and Mr Bayes 3.1.2 software.10 phylogenetic trees using both single and combined genes were constructed.The results show that Arthaldeus is not a monophyletic group with Arthaldeus arenarius being the first to separate out.However,Psammotettix and Sorhoanus are both monophyletic groups.These results indicate that the similarity of nuclear gene sequences is high.And the sequences of mitochondrial genes present more significant differences and have a greater influence on the phylogenetic tree of joint genes than do the sequences of nuclear genes.This study provides a molecular basis for the current classification system of the tribe Paralimnini.We also reconstruct the phylogeny of the tribe Paralimnini with comprehensive generic representation(45 taxa in 26 genera)based on molecular(28S,H3 and 16S)and morphological data(80 morphological characters)using Mr Bayes.The combined phylogenetic results indicate that the molecular data had a greater impact on the phylogenetic tree topology and the morphological data increases the overall support rate for the phylogenetic tree.Based on these results,Deltocephalini and Paralimnini are sister groups and,except for Arthaldeus,the genera of Paralimnini are monophyletic clades which are well supported in all analyses.Cosmotettix and Calamotettix,Paralimnus and Paramesus,Diplocolenus and Turrutus,Errastunus and Adarrus,Yanocephalus and Futasujinus,and Henschia and Mocuellus are each sister groups.These results show the relationship among genera and species for 41 representative genera of Paralimnini and provide more morphological and molecular data as a basis for the classification of tribe Paralimnini.