Molecular phylogeny and divergence times of Hormaphidinae (Hemiptera: Aphididae) indicate Late Cretaceous tribal diversification

The aphid subfamily Hormaphidinae is a good candidate for the study of the evolution of insect – plant relationships. Most hormaphidine species depend on woody primary host plants and woody or herbaceous secondary host plants, and represent high host specificity, especially to their primary hosts. No detailed molecular phylogeny of Hormaphidinae has been reported, and the taxonomic positions of some taxa in this group remain unclear. To reconstruct major phylogenetic relationships and to understand the evolution of host association patterns for major lineages, we present the first detailed molecular phylogeny of Hormaphidinae, as inferred from nuclear and mitochondrial DNA sequences, using maximum parsimony, maximum likelihood, and Bayesian methods. The monophyly of Hormaphidinae and its three traditional tribes was supported, and a sister relationship between Hormaphidini and Nipponaphidini was suggested. Most inner relationships within tribes were also supported, and some novel relationships were revealed. Two subtribes of Cerataphidini are proposed. Divergence times estimated using a Bayesian approach indicate that tribal diversifications occurred during the Late Cretaceous and were coincident with the appearance of their primary host plants. The current pattern of secondary host association for the three tribes may have evolved in different time ranges. © 2012 The Linnean Society of London, Zoological Journal of the Linnean Society, 2012, 165 , 73–87.     

Development of setal patterns in embryonic and adult aphids of the Hormaphidinae （Hemiptera： Aphididae）

The elementary traits and diversity of embryonic setal patterns of 31 species of 22 genera, in three tribes of Hormaphidinae are studied. The embryonic setal patterns to every species are provided. Based on the parsimony principle and the developmental principle, these patterns are analyzed in each tribe following the terminology of Richards （1965）. It is suggested that the setal patterns of abdominal tergites are the most important characters in embryonic and adult setal patterns. In conclusion, neoteny is a phylogenetic trend in the Hormaphidinae. Nipponaphidini and Hormaphidini have more typical neoteny in each genus and more diversity in different types of adult viviparous females than Cerataphidini, by showing more frequent and various protopatterns or mesopatterns.     

Social parasitism and behavioral interactions between two gall-forming social aphids

We investigated ecological and behavioral aspects of the interactions between two social aphids, Pseudoregma bambucicola and Astegopteryx bambucifoliae (Hormaphidinae, Cerataphidini), both of which produce second-instar, sterile soldiers in galls formed on Styrax suberifolius, in Taiwan. By censusing their galls, either species was found to invade galls of the other species. Twenty-eight (58%) out of 48 A. bambucifoliae galls contained P. bambucicola, while four (6%) of 69 P. bambucicola galls contained A. bambucifoliae. Furthermore, P. bambucicola behaved like social parasites in galls of A. bambucifoliae. Colonies of P. bambucicola produced much fewer or even no soldiers compared with those in natal galls. Our experiments also revealed that individual aphids of P. bambucicola more successfully intruded into galls of A. bambucifoliae than into conspecific galls, and that guarding soldiers of P. bambucicola effectively prevented aphids from invading their galls and permitted only conspecific soldiers to join their colonies. These differences in behavior provide good explanations for the differences in the frequency of invaded galls found between the two species. Received 4 September 2007; revised 7 January 2008; accepted 17 January 2008.     

拟爪绵蚜新属及三新种记述 (同翅目:蚜总科:瘿绵蚜科)

该文记述了瘿绵蚜科一新属拟爪绵蚜属ＳｉｃｉｕｎｇｕｉｓＺｈａｎｇｅｔＱｉａｏｇｅｎ．ｎｏｖ．和这个属的二个新种,即十绵蚜Ｓ．ｄｅｃｉｍａＺｈａｎｇｅｔＱｉａｏ,ｓｐ．ｎｏｖ．,九绵蚜Ｓ．ｎｏｖｅｎａＺｈａｎｇｅｔＨｕ,ｓｐ．ｎｏｖ．,另外还记述了爪绵蚜属ＡｐｈｉｄｏｕｎｇｕｉｓＴａｋａｈａｓｈｉ,１９６３一个新种,苹果根爪绵蚜Ａ．ｐｏｍｉｒａｄｉｃｏｌａＺｈａｎｇｅｔＨｕ,ｓｐ．ｎｏｖ．,模式标本存放在中国科学院动物研究所．拟爪绵蚜新属ＳｉｃｉｕｎｇｕｉｓＺｈａｎｇｅｔＱｉａｏｇｅｎ．ｎｏｖ．模式种：十绵蚜ＳｉｃｉｕｎｇｕｉｓｄｅｃｉｍａＺｈａｎｇｅｔＱｉａｏ,ｓｐ．ｎｏｖ．本新属与爪绵蚜属ＡｐｈｉｄｏｕｎｇｕｉｓＴａｋａｈａｓｈｉ,１９６３和绵蚜属ＥｒｉｏｓｏｍａＬｅａｃｈ,１８１８有较近的亲缘关系,不同在于：有翅孤雌蚜腹管位于腹部背片Ⅵ,隆起,周围有毛环绕（前者：位于背片Ⅵ,不隆起,无毛环绕;后者：位于背片Ⅴ,隆起,有毛环绕）;触角稍长,至少为体长０．３５,具有线形次生感觉圈;后者：稍长,至少为体长０．４０,具有环形次生感觉圈）。十绵蚜ＳｉｃｉｕｎｇｕｉｓｄｅｃｉｍａＺｈａｎｇｅｔＱｉａｏ,ｓｐ．ｎｏｖ     

Formation of long‐lasting galls by overwintered nymphs in the Japanese aphid Quadrartus yoshinomiyai (Hemiptera: Aphididae: Hormaphidinae)

This paper examines the life history of a generation of galls created by the aphid Quadrartus yoshinomiyai (Hormaphidinae: Nipponaphidini) on its primary host plant, Distylium racemosum. First‐instar fundatrix nymphs of Q. yoshinomiyai initiated galls on stems of developing shoots in early April and incipient enclosed galls were found from later the same month. The galls lasted for up to 14 months, during which they grew to maturity, opened in early or mid‐April of the following year and dried up by the end of June. First‐instar fundatrix nymphs were found on winter buds, indicating that they hatched from eggs in autumn and overwintered as nymphs. These results suggest that Q. yoshinomiyai has a three‐year life cycle.     

NINE NEW SPECIES OF PEMPHIGINAE (HOMOPTERA:PEMPHIGI-DAE) FROM CHINA*

Abstract The present paper describes 9 new species of Pemphiginae from China, vizr Mimeuria graminiradicis Zhang sp. nov., Pachypappa, aigkros Zhang sp, nov., P. tortuosae Zhang sp. nov., Pachypapella diquiplla Zhang sp. nov., Pemphigus turritus Zhang sp. nov., P. wuduenris Zhang sp. nov., Prociphilus aurus Zhang et Qiao sp. nov., P. emeiensis Zhang, sp. nov. and P. trinus Zhang sp. nov. All type specimens are deposited in the Institute of Zoology, Academia Sinica.     

TWO NEW GENERA,FIVE NEW SPECIES AND ONE NEW SUBSPECIES OF FORDINAE (HOMOPTERA: PEMPHIGIDAE) FROM CHINA*

Abstract The present paper describes 2 new genera, viz_:Dimelaphis Zhang, gen. nov and Siciforda Zhang, gen. nov, 5 new species viz: Chaetogeoica ulmidrupa Zhang sp. nov, D. obtusifalcata Zhang, sp. nov., Forda auralenta Zhang et Qiao, sp. nov., F. multicoma Zhang, sp. nov. and S. sexiarticulata Zhang, sp.nov. and 1 new subspecies Paracletus cimiciformis zhanhuanus Zhang, ssp. nov. from China. All type specimens are deposited in the Institute of Zoology, Academia Sinica.     

Molecular phylogeny of Lymantriinae (Lepidoptera,Noctuoidea, Erebidae) inferred from eight gene regions

To understand the evolutionary history of Lymantriinae and test the present higher‐level classification, we performed the first broad‐scale molecular phylogenetic analysis of the subfamily, based on 154 exemplars representing all recognized tribes and drawn from all major biogeographical regions. We used two mitochondrial genes (cytochrome c oxidase subunit I and 16S ribosomal RNA) and six nuclear genes (elongation factor‐1α, carbamoylphosphate synthase domain protein, ribosomal protein S5, cytosolic malate dehydrogenase, glyceraldehyde‐3‐phosphate dehydrogenase and wingless). Data matrices (in total 5424 bp) were analysed by parsimony and model‐based evolutionary methods (maximum likelihood and Bayesian inference). Based on the results of the analyses, we present a new phylogenetic classification for Lymantriinae composed of seven well‐supported tribes, two of which are proposed here as new: Arctornithini, Leucomini, Lymantriini, Orgyiini, Nygmiini, Daplasini trib. nov. and Locharnini trib. nov. We discuss the internal structure of each of these tribes and address some of the more complex problems with the genus‐level classification, particularly within Orgyiini and Nygmiini.     

Morphological classification of loaches (Nemacheilinae)

Comprehensive comparative morphological analysis of loaches from the subfamily Nemacheilinae is conducted. Forty significant phylogenetic characters are suggested, and phylogenetic relationships are reconstructed. The subfamily Nemacheilinae is subdivided into five tribes (Vaillantellini, Lefuini nov., Yunnanilini nov., Triplophysini nov., and Nemacheilini), and the relationships between these tribes can be expressed by the following formula: Vaillantellini (Lefuini (Yunnanilini (Triplophysini + Nemacheilini))). The classification of highland Asian loaches (Triplophysini) is specified, and the revised diagnoses of the genera from this tribe are given. Heterogeneity of the genus Triplophysa is demonstrated. This genus can be divided into several subgenera, and three new subgenera (Labiatophysa subgen. nov., Indotriplophysa subgen. nov., and Tarimichthys subgen. nov.) are described. A new species Hedinichthys grummorum sp. n is described from the Turpan Depression (northwest China).     

中国福建省瘿绵蚜科和大蚜科一新纪录属和三新种（同翅目：蚜总科）

作者在整理福建省蚜虫标本时,发现瘿绵蚜科中国１新纪录属和３新种。模式标本均保存在中国科学院动物研究所昆虫标本馆。     

Molecular phylogeny of Pemphiginae (Hemiptera: Aphididae) inferred from nuclear gene EF-1alpha sequences

Three traditional tribes of Fordini, Pemphigini and Eriosomatini comprise Pemphiginae, and there are two subtribes in Fordini and Pemphigini, respectively. Most of the species in this subfamily live heteroecious holocyclic lives with distinct primary host specificity. The three tribes of Pemphigini (except Prociphilina), Eriosomatini and Fordini use three families of plants, Salicaceae (Populus), Ulmaceae (Ulums) and Anacardiaceae (Pistacia and Rhus), as primary hosts, respectively, and form galls on them. Therefore, the Pemphigids are well known as gall makers, and their galls can be divided into true galls and pseudo-galls in type. We performed the first molecular phylogenetic study of Pemphiginae based on molecular data (EF-1alpha sequences). Results show that Pemphiginae is probably not a monophylum, but the monophyly of Fordini is supported robustly. The monophyly of Pemphigini is not supported, and two subtribes in it, Pemphigina and Prociphilina, are suggested to be raised to tribal level, equal with Fordini and Eriosomatini. The molecular phylogenetic analysis does not show definite relationships among the four tribes of Pemphiginae, as in the previous phylogenetic study based on morphology. It seems that the four tribes radiated at nearly the same time and then evolved independently. Based on this, we can speculate that galls originated independently four times in the four tribes, and there is no evidence to support that true galls are preceded by pseudo-galls, as in the case of thrips and willow sawflies.     

Dermaphis coccidiformis sp. nov. (Hemiptera), an aphid species with asymmetrically sclerotized apterae and “winter alates”

Dermaphis coccidiformis sp. nov. (Hormaphidinae: Nipponaphidini) is described from Japan. Apterous adults of the species were found between winter buds (or between a winter bud and a leaf petiole) of the evergreen oaks Quercus glauca, Q. myrsinifolia and Q. salicina. Their morphology is peculiar in that their tergites are heavily sclerotized only in the part that seems to have been exposed to sunlight. The new species is also peculiar in that nymphs to be alates (sexuparae) were found on the upper surfaces of leaves of the host oak only during winter, from December to March or early April, before the bud break of the oak. Our molecular phylogenetic analysis indicated that the new species is closely related to Dermaphis spp., therefore it was placed in the genus. The analysis incidentally indicated that “Dinipponaphis” autumna, a monoecious species forming galls on Distylium racemosum, was included in the clade of the genus Dermaphis, and therefore it was transferred to this genus.     

New fossil Ochteridae (Hemiptera: Heteroptera: Ochteroidea) from the Upper Mesozoic of north‐eastern China,with phylogeny of the family

Specimens of the velvety shore bugs (Hemiptera: Ochtheridae) occur in the Early Cretaceous Yixian Formation in north‐eastern China. We describe two new genera and three new species –Pristinochterus ovatus Yao, Zhang & Ren sp. nov., Floricaudus multilocellus Yao, Ren & Shih gen.n. et sp.n. and Angulochterus quatrimaculatus Yao, Zhang & Ren gen.n. et sp.n. – from this deposit. A cladistic analysis based on a combination of fossil and extant taxa, and their morphological characters, clarifies the phylogenetic status of the new fossils and allows the reconstruction of relationships within the family Ochteridae. Two main clades within Ochteridae are recognized from the cladistic analysis: Pristinochterus Yao Cai & Ren and Floricaudus Yao, Ren & Shih gen.n. form a monophyletic lineage; and Angulochterus Yao, Zhang & Ren gen.n. emerges as a sister group to all extant velvety shore bugs. A key to all fossil and extant genera of Ochteridae is provided.     

Phylogeny of the Braconinae (Hymenoptera: Braconidae): A new tribal order!

Generic relationships within the parasitoid wasp subfamily Braconinae are assessed based on a molecular phylogenetic analysis of four gene fragments: mitochondrial cytochrome c oxidase subunit I, 16S rDNA, nuclear 28S D2-D3 rDNA and elongation factor 1-alpha. Our results support the recognition of Aphrastobraconini, Braconini and Coeloidini plus three new tribes: Compsobraconini tribus nov ., Tropobraconini tribus nov . and Virgulibraconini tribus nov . The first of these new tribes is restricted to the New World; the second includes the Old World genera Tropobracon Cameron, Trispinaria Quicke and Grangerbracon Samartsev and Belokobylskij and possibly others, whereas the third comprises the Australian genus Virgulibracon Quicke, plus several other described and undescribed Australian genera. Consistent placement of Amyosoma Viereck with members of the Virgulibraconini tribus nov . is discussed, whereas Amyosoma is left currently unplaced. A preliminary key to tribes is presented, and the characters used to differentiate between Aphrastobraconini and Braconini are revised. Megacoeloides Quicke was never recovered with Coeloides, the type genus of Coeloidini, so it is treated as Braconinae incertae sedis. By combining molecular and morphological traits, nearly all valid genera are assigned to tribes, and the possible relationships of the remainder are discussed. Alienoclypeus Shenefelt, 1978 is synonymised with Atanycolus Förster, 1862 (Atanycolus insolitus (Shenefelt) comb. nov .). Several new genera have been revealed and will be described elsewhere.     

Systematic revision of Aluntiini Emeljanov, 1979 (Hemiptera: Fulgoromorpha: Dictyopharidae: Dictyopharinae): reclassification,phylogenetic analysis,and biogeography

The dictyopharid planthopper tribe Aluntiini s.l. is revised and reclassified into two tribes: Aluntiini s.s. and Arjunini Song & Szwedo trib. nov. The tribe Aluntiini s.s. includes five genera: Aluntia Stål, 1866; D endrophora Melichar, 1903 stat. rev. ; Dictyomorpha Melichar, 1912; Indodictyophara Liang & Song, 2012; and Madagascaritia Song & Liang gen. nov. The new tribe Arjunini comprises two genera – Arjuna Muir, 1934 and Pippax Emeljanov, 2008 – both moved from Aluntiini s.l. Four new species – A luntia longicephalica Song & Szwedo sp. nov. , Madagascaritia angusta Song & Liang sp. nov. , Arjuna maai Song & Wang sp. nov. , and Arjuna muiri Song & Wang sp. nov. – are described. A morphologically based phylogenetic analysis is undertaken for Aluntiini, Arjunini, and the representatives of Dictyopharini, Hastini, Orthopagini, and the fossil Worskaitini within Dictyopharinae, all distributed in the Old World. A matrix of 129 characters of the habitus, coloration, head, thorax, and male and female genitalia of the adults was used for the cladistic analysis. The phylogenetic results show that Aluntiini s.l. as placed in Dictyopharidae is well supported, but it is distinctly paraphyletic and should be separated into two unambiguous tribes. A palaeotropical distribution pattern displayed by Aluntiini is suggested. The origin and diversification of Aluntiini are discussed preliminarily. © 2015 The Linnean Society of London     

中国斑蚜科一新属一新种一新亚种记述(同翅目:蚜总科)

本文记述了中国斑蚜科一新属,茴斑蚜属Ｆｏｅｎｉａｐｈｉｓ Ｚｈａｎｇ ｅｔ Ｑｉａｏ,ｇｅｎ．ｎｏｖ．,一新种,长圈茴斑蚜Ｆ．ｏｂｌｏｎｇｉｓｅｎｓｏｒｉａ Ｑｉａｏ ｅｔ Ｚｈａｎｇ,ｓｐ．ｎｏｖ．和一新亚种聂长斑蚜连城亚种Ｔｉｎｏｃａｌｌｉｓ ｎｅｖｓｋｙｉ ｌｉａｎｃｈｅｎｇｅｎｓｉｓ Ｑｉａｏ ｅｔ Ｚｈａｎｇ,ｓｓｐ．ｎｏｖ．。所有模式标本均存放在中国科学院动物研究所昆虫标本馆内。     

Diversity of bacteria associated with Hormaphidinae aphids (Hemiptera: Aphididae)

Bacteria are ubiquitous inhabitants of animals.Hormaphidinae is a particular aphid group exhibiting very diverse life history traits.However,the microbiota in this group is poorly known.In the present study,using high-throughput sequencing of bacterial 16S ribosomal RNA gene amplicons,we surveyed the bacterial flora in hormaphidine aphids and explored whether the aphid tribe,host plant and geographical distribution are associated with the distribution of secondary symbionts.The most dominant bacteria detected in hormaphidine species are heritable symbionts.As expected,the primary endosymbiont Buchnera aphidicola is the most abundant symbiont across all species and has cospeciated with its host aphids.Six secondary symbionts were detected in Hormaphidinae.Arsenophonus is widespread in Hormaphidinae species,suggesting the possibility of ancient acquisition of this symbiont.Ordination analyses and statistical tests show that the symbiont composition does not seem to relate to any of the aphid tribes,host plants or geographical distributions,which indicate that horizontal transfers might occur for these symbionts in Hormaphidinae.Correlation analysis exhibits negative interference between Buchnera and coexisting secondary symbionts,while the interactions between different secondary symbionts are complicated.These findings display a comprehensive picture of the microbiota in Hormaphidinae and may be helpful in understanding the symbiont diversity within a group of aphids.     

Revision and phylogenetic assessment of the rove beetle genus Pseudohesperus Hayashi,with broad reference to the subtribe Philonthina (Coleoptera: Staphylinidae: Staphylinini)

This paper studies the phylogeny of the rove beetle subtribe Philonthina, to test its hypothetical monophyly and to unravel the evolutionary relationships of the subtribe and its included genus‐level taxa, with emphasis on the genus Pseudohesperus and its close‐allied relatives. The phylogenetic analyses are based on 105 adult morphological characters and 66 terminal taxa, i.e., all six members of Pseudohesperus, 51 species to represent 29 other genera of the subtribe Philonthina, seven species to represent the other six subtribes of Staphylinini, one species of the tribes Arrowinini, and one of the Platyprosopini. According to the phylogenetic results obtained, the genus Erichsonius should move out from the hitherto‐defined subtribe Philonthina and thus the monophyly of this taxon is challenged. The phylogenetic tree suggests that the genera Hesperus and Belonuchus might not be monophyletic, but the monophyly of Pseudohesperus and the sister relationship between it and Bisnius are well supported. The species‐level phylogenetic relationships of the genus Pseudohesperus reveal a clear pattern of species diversification that can be correlated well with the species' zoogeographical patterns. The paper also revises the taxonomy of Pseudohesperus and describes five new species from China: Pseudohesperus luteus Li & Zhou sp. nov. , Pseudohesperus pedatiformis Li & Zhou sp. nov. , Pseudohesperus tripartitus Li & Zhou sp. nov. , Pseudohesperus sparsipunctatus Li & Zhou sp. nov. , and Bisnius lubricus Li & Zhou sp. nov. An identification key to the species of Pseudohesperus is provided and their geographical distributions are mapped. © 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 163 , 679–722.     

A new genus of Entiminae from North Africa,supported by a phylogenetic analysis (Coleoptera: Curculionidae: Entiminae)

The genus Holcophloeus gen. nov. is here proposed to include Trachyphloeus cruciatus Seidlitz, 1868, and two new species native to North Africa, based on a phylogenetic analysis and an evaluation of the diagnostic characters. The taxonomic position of Holcophloeus in relation to the tribes Trachyphloeini Lacordaire, 1863, and Holcorhinini Desbrochers, 1898, is discussed, and the new genus is attributed to the Holcorhinini. Holcophloeus laurae sp. nov. from south‐eastern Morocco and Holcophloeus weilli sp. nov. from northern Libya are described and illustrated and a key to the species of the new genus is given. The lectotype of Trachyphloeus cruciatus Seidlitz, 1868, is designated. The genus Massimiellus Borovec, 2009, is transferred from Trachyphloeini to Holcorhinini. © 2013 The Linnean Society of London