Phylogenetic And Historical Biogeography Of Partial Genera Of The Tribe Blaptini From The Qinghai-Xizang Plateau

The tribe Blaptini is a species-rich group in the subfamily Tenebrioninae(Coleoptera,Tenebrionidae),whose species are mostly distributed in the Central-Eastern Asia.It includes approximately 741 species/subspecies that belong to 27 genera of 6 subtribes,and 269 species/subspecies of 20 genera have been recorded from China.Researchers have paid much attention to the classification of this tribe and constructed diverse classification systems,e.g.23 genera of 5 subtribes by Medvedev(2001)and 27 genera of 6 subtribes by Ren(2016).However,these classification systems are mainly based on comparative morphological characteristics.The works to molecular phylogenetic studies are extended presently,and the first molecular phylogenetic tree based on three mitochondrial molecular markers(COI,Cytb,and 16 S rDNA)and one nuclear molecular marker(28S rDNA-D2)representing 12 genera of the tribe Blaptini are reconstructed.Phylogenetics,molecular dating,and ancestral area reconstruction are used to investigate the endemic genus Gnaptorina from the Qinghai-Xizang Plateau.The results of phylogenetic and molecular species delimitation analyses clarify the taxonomy and systematics of the genus.The following results have been preliminarily obtained:1.The first molecular phylogenetic tree of the genus Gnaptorina was constructed based on four DNA markers.Three distinct molecular species delimitation approaches,Automatic Barcode Gap Discovery(ABGD),Generalized Mixed Yule Coalescent(GMYC),and Poisson Tree Processes(PTP),are implemented to reassess the status of 28 morphological species of the genus Gnaptorina.The results are consistent with morphological classification,and confirmed the classification status of most extant morphological species and four putative new species.Based on the results,a new taxonomic arrangement for Gnaptorina is proposed.A new subgenus Austroptorina Bai,Li & Ren subgen.n.is defined.Consequently,the new generic synonymy is established: the subgenus Boreoptorina Medvedev,2009,syn.n.is a junior synonym of the subgenus Hesperoptorina Medvedev,2009;and the subgenus Gnaptorina is rendered monophyletic thanks to the inclusion of G.dongdashanensis Shi,2013.Besides,G.minxiana Medvedev,2008,formerly treated as a subspecies of G.potanini Reitter,1889,is elevated to species rank.2.Dating and historical biogeography analyses based on 58 individuals of the genus GnaptorinaThe results of dating and historical biogeographic analyses based on DNA and the distributions of 28 extant species(58 individuals)of the genus Gnaptorina suggested an early Eocene origin of the genus with the South-Eastern regions of the Qinghai-Xizang Plateau as most likely areas of origin.The uplifting events of the Qinghai-Xizang Plateau blocked gene flow among species or populations,and likely influenced the current diversity patterns of the genus Gnaptorina.Deep valleys and the multitude of high mountain ranges likely prohibited a rapid spread of Gnaptorina from one valley to another.The scenario derived from the analyses suggested a long evolutionary history of the genus Gnaptorina at the Qinghai-Xizang Plateau.The most likely scenario is that the genus Gnaptorina originated in the South-Eastern regions of the Qinghai-Xizang Plateau regions and then dispersed progressively eastward and westwards.The uplifting of the Qinghai-Xizang Plateau led to the further formation of many geographical isolated habitats,and promoted the generation of many new species.Variance was likely a major evolutionary driver for the genus at that time.The scenario derived from our analyses suggested a long evolutionary history of Gnaptorina in the Qinghai-Xizang Plateau region,with three distinct lineages progressively colonizing distinct areas during the uplifting of the plateau.3.Part genera of the tribe Blaptini are rendered monophyletic and explained reasonably the molecular phylogenetic relationship from Blaptini.(1)In this study,the first molecular phylogenetic tree of the tribe Blaptini was constructed based on four DNA markers.The tree revealed the phylogenetic relationships within the tribe Blaptini.Also,the results show that the genera Asidoblaps,Blaptogonia,Belousovia,and Agnaptoria are closer living relatives,and the genus Itagonia is the closest existing relatives to the genus Gnaptorina.Most genera were recovered with strong support as monophyletic,such as Prosodes,Asidoblaps,Blaptogonia,Belousovia and Agnaptoria.(2)Three species of the genus Itagonia,one species of the genus Pseudognaptorina and species of the genus Gnaptorina were formed a paraphyletic taxon,but all species were classified to a monophyletic group in the phylogenetic tree with strong support.One species of the genus Dila and two species of the genus Coelocnemode were paraphyletic,but three species forming a monophyletic group in the phylogenetic tree with strong support.(3)Three species from the genus Itagonia(Itagonia mera Medvedev,1998,Itagonia zayica Shi & Ren,2007,and Itagonia baxoica Liu & Ren,2009)and one species from the genus Pseudognaptorina(P.obtuse Shi,Ren & Merkl,2007)were transferred to the genus Gnaptorina based on molecular and morphological evidences through rechecking the type specimens and the determined specimens.Also,two Pseudognaptorina species,Pseudognaptorina exsertogena Shi,Ren & Merkl,2007 and Pseudognaptorina flata Liu & Ren,2007,were transferred to the genus Gnaptorina according to the morphological characteristics.Dila bomina Ren & Li,2001 was transferred to the genus Coelocnemodes based on molecular and morphological evidences.Three reintegrated genera(Gnaptorina,Itagonia,and Coelocnemodes)were recovered with molecular data strongly supporting to be monophyletic.(4)Dating analyses of the tribe Blaptini.Dating analyses based on 154 individuals of the tribe Blaptini suggested the late Cretaceous Period origin.The tribe Blaptini was rapidly radiation evolution in the Oligocene and Miocene with the rapidly uplifting of the Qinghai-Xizang Plateau.The evolutionary of the tribe Blaptini may have been caused by the collision of two plates in the Oligocene.The continuous uplifting of the Qinghai-Xizang Plateau at the end of the Eocene to the Indian plate fault ended,which led to dramatic changes in geological structure.The continuous orogeny created isolated speciation events,which also contributed to the rapid radiation evolution of endemic genera and species during this period.5.Discovery of a new subgenus,seven new species,and six new combinations:A new subgenus: Gnaptorina(Austroptorina)Bai,Li & Ren subgen.n.Seven new species: Blaptogonia zhentanga Bai,Li & Ren,2018(type locality: Xizang),Gnaptorina(Austroptorina)polita Bai,Li & Ren sp.nov.(type locality: Xizang),Gnaptorina(Hesperoptorina)ampliptera Bai,Li & Ren sp.nov.(type locality: Sichuan),Gnaptorina(Gnaptorina)acutangula Bai,Li & Ren,2019(type locality: Sichuan),Gnaptorina(Gnaptorina)artipenis Bai,Li & Ren sp.nov.(type locality: Sichuan),Gnaptorina(Gnaptorina)crassitibia Bai,Li & Ren,2019(type locality: Sichuan),Gnaptorina(Gnaptorina)crenata Bai,Li & Ren sp.nov.(type locality: Sichuan).Seven new combinations: Gnaptorina(Gnaptorina)obtuse(Shi,Ren & Merkl,2005)comb.nov.,Gnaptorina(Gnaptorina)exsertogena(Shi,Ren & Merkl,2005)comb.nov.,Gnaptorina(Gnaptorina)flata(Liu & Ren Merkl,2009)comb.nov.,Gnaptorina(Gnaptorina)baxoica(Liu & Ren,2009)comb.nov.,Gnaptorina(Gnaptorina)mera(Medevedev,1998)comb.nov.,Gnaptorina(Austroptorina)zayica(Shi & Ren,2007)comb.nov.,and Coelocnemodes bomina(Ren & Li,2001)comb.nov.