Guyiling jianboni gen. et sp.n., an antlion‐like lacewing,illuminating homologies and transformations in Neuroptera wing venation

A new genus and species of antlion‐like fossil lacewing, Guyiling jianboni gen. et sp.n. (Insecta: Neuroptera) are described based on a single specimen from the Yixian Formation (Liaoning Province, China; Early Cretaceous). The new taxon exhibits derived traits such as distally dilated antennae and well‐developed anterior Banksian line (known in Myrmeleontidae), but also genuine plesiomorphies (at the level of Myrmeleontiformia), such as the divergence of a distinct CuA1 stem from MP2 + CuA1 (forewing), and a basal origin of MA (diverging from RP + MA; both wing pairs). This combination is unique among the ‘Palaeoleontidae’, a group better considered as a paraphyletic assemblage of various stem‐Myrmeleontiformia. The wing morphology of the new species is considered in the light of a survey of wing venation topological homologies (and implied transformations) among several Neuroptera families. The survey includes a revision of the holotype of Leptolingia jurassica Ren (Grammolingiidae; Jiulongshan Formation, Daohugou locality, Inner Mongolia Autonomous Region, China; middle Jurassic). The forewing morphology of Guyiling jianboni gen. et sp.n. demonstrates that the fusion of MP2 with CuA, and the differentiation of CuA into two distinct main stems (namely CuA1 and CuA2) are traits shared with Myrmeleontidae and Ascalaphidae (and possibly Nemopteridae). However, the survey also demonstrates that a fusion of MP2 with CuA occurred repeatedly among Neuropterida, although by means of various modalities (namely translocation vs regular fusion). The ‘pectinate fusion’ of CuA(1) with MP2 [i.e. CuA1 has no distinct stem and emits successive branches from MP2 + CuA(1) partim] is a further step in this fusion, and occurred repeatedly as well (at least in the hind wings of Osmylidae and Nymphidae, and both fore‐ and hindwings of a sub‐group of Myrmeleontiformia including Myrmeleontidae and Ascalaphidae, and possibly Nemopteridae). It is anticipated that the current contribution will constitute useful background information for further studies, focusing on particular transformation cases, ideally including a documentation of intraspecific variation.     

A new Upper Carboniferous stem‐orthopteran (Insecta) from Ningxia (China)

Abstract During the Upper Carboniferous, orthopteran insects (grasshoppers, katydids, and crickets) were represented by numerous species distantly related to crown‐orthopterans, such as lobeattid and cnemidolestodean insects. The panorthopterans, including total‐orthopterans and their closest relatives, are represented by comparatively rarer species in localities of this period. Here we describe Heterologus duyiwuer sp. nov., an infrequent panorthopteran from the Late Carboniferous locality of Xiaheyan Village (Zhongwei City, Ningxia Hui Autonomous Region, China). The only available specimen is composed of an isolated forewing exhibiting a combination of character states previously unknown, in particular the lack of posterior radius (RP) / anterior Media (MA) connection, late branchings of the media (M) and anterior cubitus (CuA), and a branched posterior branch of the posterior cubitus (CuPb). Based on its unusual branching pattern, the composite stem resulting from the fusion of CuA and CuPaα (second anterior branch of CuP) is assumed to be composed of a branched CuA and a simple CuPaα.     

Evidence for Carboniferous origin of the order Mantodea (Insecta: Dictyoptera) gained from forewing morphology

Homologies of the forewing venation pattern of the order Mantodea (Insecta: Dictyoptera) consistent with the accepted insect wing venation groundplan are proposed. A comparative morphological analysis was carried out based on a broad taxonomic sample of extant taxa. Besides macromorphological aspects, focus is given to the pattern of the tracheal system as a basis for establishing primary homologies. All extant praying mantids exhibit a composite stem composed of the posterior radius (RP) and the media (M) and most praying mantids exhibit a fusion of the anterior branch of RP + M with the anterior radius (RA). The wing venation of the species ?Mesoptilus dolloi, previously assigned to the polyphyletic fossil assemblage ‘Protorthoptera’, is re‐interpreted in the light of the new homology statement. Our interpretation suggests that it is a putative stem‐Mantodea, as are some other ‘protorthopterous’ taxa. This hypothesis implies that the total‐group Mantodea arose as soon as the Late Carboniferous, i.e. about 175 million years earlier than previously estimated. This analysis contributes to the view that most of the Late Carboniferous ‘Protorthoptera’ are stem‐representatives of the major polyneopteran clades (e.g. cockroaches, grasshoppers and crickets, rock‐crawlers), suggesting a survivorship of several main Pterygota lineages at the end‐Permian extinction event higher than previously expected. © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 156 , 79–113.     

中国锯天牛族后翅翅脉研究(鞘翅目,天牛科,锯天牛亚科,锯天牛族)(英文)

对中国锯天牛族的后翅基部关节和后翅翅脉特征进行了研究,发现利用Kukalová-Peck和Lawrence (2004)的后翅命名系统能够很好地对中国锯天牛族后翅翅脉进行命名。但是在中国锯天牛族中,后中脉( MP)和前肘脉(CuA)在后缘并不合并;当前臀脉( AA3)和前肘脉(CuA3 +4)与后肘脉(CuP)相遇时,前臀脉(AA3)消失,前肘脉(CuA3 +4)和后肘脉(CuP)合并,因此楔室(W)仅由肘脉(Cu)的分支脉围成。尽管基部翅关节在研究的各属和各种之间没有表现出差异,但是后翅翅脉在土天牛属Dorysthenes和锯天牛属Prionus不同种类之间差异明显,这些特征包括径室的长宽比例和各边的长度关系、r3存在与否及其长度、后径脉的长度、楔室的长宽比例、以及后中脉(MP3 +4)和前肘脉(CuA3 +4)端部是否分叉等。因此,后翅翅脉特征在土天牛属Dorysthenes和锯天牛属Prionus分种时可能具有分类学意义。     

How Gerarus lost its head: stem-group Orthoptera and Paraneoptera revisited

Abstract.  The Upper Carboniferous insect Gerarus from Mazon Creek was reinvestigated to assess the affinities of the Geraridae. Specimens were examined using an environmental scanning electron microscope. The evidence, including an inflated clypeus and characters of the wing venation, for placing the family in Paraneoptera (hemipteroids) is equivocal; Gerarus is assigned to the Archaeorthoptera in the stem-group Orthoptera. The presence of leg exites in Gerarus and other Mazon Creek specimens could not be confirmed. Definitive evidence for the origin of insect wings remains to be discovered.     

Lodetiella magnifica nov. gen. and nov. sp. (Insecta: Palaeodictyoptera; Permian), an extreme situation in wing morphology of palaeopterous insects

Based on published data and review of Mecynoptera splendida Handlirsch, 1904, a wing venation ground plan applicable to most taxa of the Palaeozoic superorder Palaeodictyopteroidea is proposed, where each major vein system has a simple anterior sector, and, primitively, a dichotomously branched posterior sector. Lodetiella magnifica nov. gen. and sp. is described, based on an isolated hind wing from the Salagou Formation (Middle to Late Permian; Hérault, France). The venation of Lodetiella magnifica conforms to the predicted ground plan, but differs by its simple posterior cubitus sector. This character is diagnostic of the family Calvertiellidae (Palaeodictyoptera), whose diagnosis is updated. Wing movement in flight is inferred from the peculiar wing vein organization of Lodetiella magnifica. Some ecological preferences might explain the relative rarity of calvertiellids in the fossil record.     

High variability in tegminal venation of primitive cercopoids (Hemiptera: Cicadomorpha), as implied by the new discovery of fossils from the Middle Jurassic of China

The Procercopidae is the most ancient family of the Cercopoidea and is known to be low in diversity. Herein, a new genus and species of Procercopidae, Titanocercopis borealis Chen, Zhang & Wang, is erected from the Middle Jurassic of Daohugou of Inner Mongolia, China. The venation patterns show considerable intra‐specific or even intra‐individual variation in specimens involved in this study, suggesting that the flight ability of these primitive cercopoids might be not very strong. The new genus differs from other known genera by the following combination of morphological characters: body and tegmen large; tegmen with clavus long and narrow; apical margin rounded; vein M bifurcating basal of or almost at the same level of vein CuA bifurcation; stem ScP+R short, nearly straight, and branching into veins ScP+RA and RP near basal 1/3 wing length.     

中国西北地区下侏罗统三工河组原沫蝉科(昆虫纲:半翅目)一新属种*

本文根据一枚产自中国新疆维吾尔自治区下侏罗统三工河组的昆虫化石标本建立1新属新种——秉氏巨沫蝉(Magnicercopis pingi Zhang, Chen et Zhang, gen. et sp. nov.), 归入已灭绝的半翅目原沫蝉科。本新属的衍征包括: MP脉3分支, 存在横脉im, MP脉分支处于CuA脉分支处的基侧。本文对新属种与中国侏罗纪的原沫蝉科类群, 尤其是Longucercopis xinjiangensis Zhang, Chen et Zhang, 2022进行了详细的翅脉对比: M. pingi盖翅长约为L. xinjiangensis的两倍、MP脉3分支(后者2分支)、MP脉首次分支处位于CuA脉分支处基侧(后者位于CuA脉分支处端侧)、M. pingi具横脉im (后者未见im脉)。这一发现不仅提高了对原沫蝉科多样性的认识, 也丰富了该科的翅脉信息。     

现存蜉蝣翅基纵脉走向及愈合模式(昆虫纲：蜉蝣目)

有翅昆虫翅基纵脉的走向及愈合模式在系统发育重建中占有重要地位。然而,现存蜉蝣翅基纵脉的走向及愈合状况在大部分种类变化极大,无法推测其原始状况,只在极少数种类保留有部分可见残迹。中国拟短丝蜉Siphluriscus chinensis的翅基保留有独立的亚前缘脉弓、部分中脉M和肘脉Cu主干以及前中脉MA及径分脉Rs的走向痕迹。据此并结合红斑蜉Ephemera rufomaculata 和大网脉蜉Chromarcys magnifica翅基的相关特征,本文提出了蜉蝣目主要纵脉基部走向及愈合的基本模式,其要点有：中脉主干在基部与径脉主干独自发出后先接近或愈合后又分离、它们各自分成两支后的前中脉及径分脉又先愈合再分离、肘脉始终独立。这种中脉与径脉先接近或愈合后分离的模式非常接近新翅类的情况而与蜻蜓很不相同(在蜻蜓,中脉与肘脉在基部愈合) 。亚前缘脉弓的作用相信是加强了因翅基骨板发达而相互远离的纵脉间的连结作用。这个假说也可以来解释蜻蜓复杂脉相的形成原因。     

Revision of the enigmatic Upper Carboniferous insect Campyloptera eatoni Brongniart, 1893 (Insecta: Odonatoptera)

Campyloptera eatoni Brongniart, 1893, the type species of the type genus of the Upper Carboniferous family Campylopteridae Handlirsch, is redescribed. It is not a Megasecoptera as previously supposed, but an Odonatoptera with a specialized wing venation. Although it has a more basal position than the Meganeuridae because of the absence of any nodal or subnodal structure, it has acquired a simple vein MA and a widening area between MP and CuA, convergently with the highly derived Discoidalia clade that includes the modern Odonata. A new diagnosis is given for Campylopteridae and its type genus, Campyloptera Brongniart. Campylopterodea Rohdendorf, 1962 falls as a new junior synonym under Odonatoptera Martynov, 1932.     

Carboniferous Onychophora from Montceau‐les‐Mines,France, and onychophoran terrestrialization

The geological age of the onychophoran crown‐group, and when the group came onto land, have been sources of debate. Although stem‐group Onychophora have been identified from as early as the Cambrian, the sparse record of terrestrial taxa from before the Cretaceous is subject to contradictory interpretations. A Late Carboniferous species from the Mazon Creek biota of the USA, Helenodora inopinata, originally interpreted as a crown‐group onychophoran, has recently been allied to early Cambrian stem‐group taxa. Here we describe a fossil species from the Late Carboniferous Montceau‐les‐Mines Lagerstätte, France, informally referred to as an onychophoran for more than 30 years. The onychophoran affinities of Antennipatus montceauensis gen. nov., sp. nov. are indicated by the form of the trunk plicae and the shape and spacing of their papillae, details of antennal annuli, and the presence of putative slime papillae. The poor preservation of several key systematic characters for extant Onychophora, however, prohibits the precise placement of the Carboniferous fossil in the stem or crown of the two extant families, or the onychophoran stem‐group as a whole. Nevertheless, A. montceauensis is the most compelling candidate to date for a terrestrial Paleozoic onychophoran.     

Dpp/BMP transport mechanism is required for wing venation in the sawfly Athalia rosae

The pattern of wing venation varies considerably among different groups of insects and has been used as a means of species-specific identification. However, little is known about how wing venation is established and diversified among insects. The decapentaplegic (Dpp)/bone morphogenetic protein (BMP) signaling pathway plays a critical role in wing vein formation during the pupal stages in Drosophila melanogaster. A key mechanism is BMP transport from the longitudinal veins (LVs) to the posterior crossvein (PCV) by the BMP-binding proteins, short gastrulation (Sog) and twisted gastrulation2/crossveinless (Tsg2/Cv). To investigate whether the BMP transport mechanism is utilized to specify insect wing vein patterns in other than Drosophila, we used the sawfly Athalia rosae as a model, which has distinct venation patterns in the fore- and hindwings. Here, we show that Ar-dpp is ubiquitously expressed in both the fore- and hindwings, but is required for localized BMP signaling that reflects distinct wing vein patterns between the fore- and hindwings. By isolating Ar-tsg/cv in the sawfly, we found that Ar-Tsg/Cv is also required for BMP signaling in wing vein formation and retains the ability to transport Dpp. These data suggest that the BMP transport system is widely used to redistribute Dpp to specify wing venation and may be a basal mechanism underlying diversified wing vein patterns among insects.     

On the phylogenetic position of the palaeopteran Syntonopteroidea (Insecta: Ephemeroptera), with a new species from the Upper Carboniferous of England

A new syntonopterid, Anglolithoneura magnifica gen. et sp. n., is described from a siderite concretion (nodule) from the Late Carboniferous (Langsettian) of Lancashire County (UK). The new genus is diagnosed on hind wing venation and compared with other syntonopterid genera. The new species is the first syntonopterid formally described from the Late Carboniferous of Europe. The systematic positions of other potential Syntonopteroidea (Miracopteron mirabile, Bojophlebia prokopi, and specimens described in 1985 by J. Kukalová-Peck from Obora in the Czech Republic) are reconsidered. Wing venation synapomorphies are proposed for the Syntonopteroidea (sensu novo), and for a potential clade ((Ephemeroptera+Syntonopteroidea)+Odonatoptera) separated from the Palaeodictyopterida. The close relations of the new species with Lithoneura lameerei Carpenter, 1938 from Mazon Creek (Illinois, USA) provide additional support for a Euramerican connection during the Late Carboniferous.     

Fluctuating asymmetry in the honey bee,Apis mellifera: effects of ploidy and hybridization

We present a new measure of morphological asymmetry that avoids most of the statistical problems inherent in character-by-character analysis of size or shape. The method is an application of Procrustes analysis, which computes best-fitting super-positions of configurations of landmarks to the left and right sides of a single specimen. The Procrustes method combines subtle deviations in all aspects of the landmark configuration into one net asymmetry score. Directional asymmetry is separated from fluctuating asymmetry in a simple partition of a net sum-of-squares, and geometrical details of either component can be inspected by traditional methods of multivariate statistical analysis of landmarks. We demonstrate this method in a comparison of wing venation asymmetry in male (haploid) and female (diploid) honey bees (Apis mellifera). In addition we investigate the effects of ploidy and inter-subspecies hybridization on asymmetry and wing venation abnormalities, using the subspecies A. m. mellifera, A. m. carnica, and the hybrid strain “Nigra”. Results suggest that while the haploid males showed a higher frequency of wing venation abnormalities and greater total asymmetry than the diploid females, most of the asymmetry difference between males and females was in the form of directional, not fluctuating, asymmetry. Hybrid females had a higher frequency of wing venation abnormalities than females of either subspecies, but there were no significant differences in the mean level of asymmetry among females of A. m. mellifera, A. m. carnica and hybrid Nigra. Hybrid males had higher absolute frequency of wing venation abnormalities and asymmetry than males of either subspecies. However the mean frequency of venation abnormalities did not differ significantly between Nigra and A. m. carnica males, and mean asymmetries were not significantly different between Nigra and A. m. mellifera males. We discuss the relationship which is assumed to exist between developmental stability and fluctuating asymmetry in light of our result.     

Insights into the molecular mechanisms underlying diversified wing venation among insects

Insect wings are great resources for studying morphological diversities in nature as well as in fossil records. Among them, variation in wing venation is one of the most characteristic features of insect species. Venation is therefore, undeniably a key factor of species-specific functional traits of the wings; however, the mechanism underlying wing vein formation among insects largely remains unexplored. Our knowledge of the genetic basis of wing development is solely restricted to Drosophila melanogaster. A critical step in wing vein development in Drosophila is the activation of the decapentaplegic (Dpp)/bone morphogenetic protein (BMP) signalling pathway during pupal stages. A key mechanism is the directional transport of Dpp from the longitudinal veins into the posterior crossvein by BMP-binding proteins, resulting in redistribution of Dpp that reflects wing vein patterns. Recent works on the sawfly Athalia rosae, of the order Hymenoptera, also suggested that the Dpp transport system is required to specify fore- and hindwing vein patterns. Given that Dpp redistribution via transport is likely to be a key mechanism for establishing wing vein patterns, this raises the interesting possibility that distinct wing vein patterns are generated, based on where Dpp is transported. Experimental evidence in Drosophila suggests that the direction of Dpp transport is regulated by prepatterned positional information. These observations lead to the postulation that Dpp generates diversified insect wing vein patterns through species-specific positional information of its directional transport. Extension of these observations in some winged insects will provide further insights into the mechanisms underlying diversified wing venation among insects.     

中国谷蛾一新属新种(鳞翅目:谷蛾科)

本文记述地谷蛾亚科Meessiinae的1新属:合腺地谷蛾属Unilepidotricha gen.nov.,及新种细弯合腺地谷蛾Unilepidotricha gracilicurva sp.nov.(图1~6)。新属与Xeringinia属近似且新种与X.altilis(Meyrick)很相似,但雄性腹部背面第3与第4节之间有特殊的狭长鳞片斑,颚形突和阳茎基环发达;雌性交配囊有2个囊突等可以区别。模式标本采自云南大理州巍山县巍宝山自然保护区,保存在南开大学生命科学院昆虫标本室。     

Records of bizarre Jurassic brachycerans in the Daohugou biota,China (Diptera,Brachycera, Archisargidae and Rhagionemestriidae)

Abstract: Four new Brachycera fossils were collected from the Daohugou biota, China. Among these, two impressions demonstrating peculiar wing venation can be designated as two new species of a new genus (Mostovskisargus portentosus gen. et sp. nov. and M. signatus sp. nov.) referred to a new subfamily Mostovskisarginae (subfam. nov.) within Archisargidae. The third impression belongs to a new species (Calosargus (Pterosargus) sinicus sp. nov.) referred to the subgenus Pterosargus Mostovski, 1997 of Calosargus Mostovski, 1997 within Archisarginae, Archisargidae. It is only the second member of the subgenus Pterosargus worldwide. The fourth impression is a nearly complete fly, Jurassinemestrinus orientalis gen. et sp. nov. referable to Rhagionemestriidae. It reveals new morphological data about rhagionemestriids and extends the geographical distribution of this group beyond Europe and Central Asia during the Mesozoic. A correlation for the archisargid composition of taxa in Daohugou and Karabastau formations is discussed. The age of the fly‐bearing strata is briefly reassessed and can be more accurately limited to the Callovian–Oxfordian based on both biostratigraphical correlation and radiometric dating.     

叶甲亚科后翅比较形态学研究

通过对叶甲亚科昆虫20属20种后翅的比较形态学研究,主要对叶甲亚科翅脉名称、翅脉变化类型进行了探讨,给出了叶甲亚科翅脉模式图、20种翅脉的形态特征图和翅脉变化类型图,并对其进化意义进行了初步的总结,认为中国叶甲亚科中最原始的后翅类型应为：长宽比例大,Cu1a与Cu1b之间具cv横脉或Cu1b与Pcu之间具cp横脉以及Cu1b与Pcu端部分开,不融合为一根。     

中国隐斑谷蛾属一新种(鳞翅目:谷蛾科)

隐斑谷蛾属Crypsithyris全世界已知38种,其中5种分布于中国。本文记述了采自湖南、贵州和江西的1新种:锯隐斑谷蛾Crypsithyris serrata Xiao et Li,sp.nov.(Figs.1～3)。新种与分布日本的Crypsithyris japonica Petersen et Gaedike和Crypsithyris saigusai Gaedike相似,主要区别如下:新种前翅暗褐色,具1大的椭圆形透明斑以及沿后缘具1赭白色纵带;抱器瓣近矩形,末端斜直;抱器背基突大,W形;阳茎约与囊形突等长,两侧具数个锯齿。后2种前翅乳白色,具1模糊的透明斑,在透明斑外侧有1暗斑;抱器瓣基半部宽于端半部,末端尖或钝圆;抱器背基突窄并相连成桥;阳茎明显长于囊形突且两侧无锯齿。模式标本保存在南开大学生命科学学院昆虫标本室。     

Systematic position of Triplosoba,hitherto the oldest mayfly,from Upper Carboniferous of Commentry in Central France (Insecta: Palaeodictyopterida)

A revision of the holotype of Triplosoba pulchella (Brongniart, 1883), the ‘oldest mayfly’, provides evidence for new attribution to Palaeodictyopterida sensu Bechly. New character states derived from the venation pattern, namely the basal connection of the median anterior with the radius and radius posterior in the forewing, the median anterior remote from the radius posterior and basally fused with the median posterior in the hindwing, and a strong costal margin formed by two connected or fused veins with a series of small spines (costa and subcosta anterior), support an ordinal placement of Triplosoba into Palaeodictyopterida (inclusive of Diphanopterodea sensu Bechly) versus Ephemeroptera as supposed by previous authors. It shares with Diaphanopterodea a potential synapomorphy in the presence of long curved connections between the three stems of the radius, media, and cubitus in the basal part of the forewing, a character not present in other groups of Palaeodictyopterida.