Ultrastructure of the larval eye of the scorpionfly Panorpa dubia (mecoptera: Panorpidae) with implications for the evolutionary origin of holometabolous larvae

The evolutionary origin of holometabolous larvae is a long‐standing and controversial issue. The Mecoptera are unique in Holometabola for their larvae possessing a pair of compound eyes instead of stemmata. The ultrastructure of the larval eyes of the scorpionfly Panorpa dubia Chou and Wang, 1981 was investigated using transmission electron microscopy. Each ommatidium possesses a cornea, a tetrapartite eucone crystalline cone, eight retinula cells, two primary pigment cells, and an undetermined number of secondary pigment cells. The rhabdomeres of the eight retinula cells form a centrally‐fused, tiered rhabdom of four distal and four proximal retinula cells. The rhabdomeres of the four distal retinula cells extend distally into a funnel shape around the basal surface of the crystalline cone. Based on the similarity of the larval eyes of Panorpidae to the eyes of the hemimetabolous insects and the difference from the stemmata of the holometabolous larvae, the evolutionary origin of the holometabolous larvae is briefly discussed. Morphol., 2012. © 2012 Wiley Periodicals, Inc.     

Evolutionary history of the scorpionfly Dicerapanorpa magna (Mecoptera,Panorpidae)

Climate changes can have fundamental impacts on the distributional patterns of montane species, and range shifts frequently lead to allopatric divergence followed by the establishment of secondary contact zones. Many European and North American organisms have retreated to southern refugia during glacial periods and colonized northward during postglacial periods, but little is known about the evolutionary response of cold‐adapted insects to Pleistocene climate changes in eastern Asia. The scorpionfly Dicerapanorpa magna (Chou), with cold temperate habitat preference and weak dispersal ability, provides a good model system to explore how climate changes have influenced the distribution and divergence of cold‐adapted insects in eastern Asia. This study reconstructed the demographic dynamics and evolutionary history of D. magna with phylogeographic approaches, and predicted the species’ suitable areas under the Last Glacial Maximum (LGM) and current scenarios with the ecological niche modelling analysis. The mitochondrial cytochrome c oxidase subunit I resolved three phylogenetic lineages in D. magna dating back to Pleistocene, corresponding well with the geographically isolated Qinling, Bashan and Minshan Mountains. The ecological niche modelling recovered the suitable habitats for D. magna were the Qinling and Bashan Mountains under LGM and current conditions. The three lineages of D. magna might be in a process of incipient speciation, and likely derived their current distribution from separate glacial origins, followed by vicariance and divergence.     

Ultrastructure of the larval midgut of Bittacus planus (Mecoptera: Bittacidae) and Neopanorpa longiprocessa (Mecoptera: Panorpidae)

Bittacidae and Panorpidae are the two largest families in Mecoptera. The larvae of Bittacidae are different from those of Panorpidae in external morphology and habits, and have an interesting habit of spraying the body surface with soil through the anus. However, it remains unknown to date whether the larval midguts are different in structure between the two families. Here the ultrastructure of the larval midguts of the hangingfly Bittacus planus Cheng and the scorpionfly Neopanorpa longiprocessa Hua & Chou were compared using light, scanning, and transmission electron microscopy. The midguts of both species are simple tubes of single layered epithelia with digestive and regenerative cells but without diverticula. The basal plasma membrane of epithelial cells exhibits infolding in B. planus, but is closely apposed to its basal lamina in N. longiprocessa. Lymph spaces are present between adjacent epithelial cells in B. planus, but are absent in N. longiprocessa. The regenerative cells are scattered among the digestive cells in B. planus, but are aggregated in N. longiprocessa. The longitudinal muscle bands are compact in B. planus, but are sparse in N. longiprocessa. The compact longitudinal muscle bands are likely associated with their soil-spraying habit in Bittacidae.     

Fine structural description of the compound eye of the Madagascar 'hissing cockroach'Gromphadorhina portentosa (Dictyoptera: Blaberidae)

The compound eyes of the wingless adults of the Madagascar ‘hissing cockroach’Gromphadorhina portentosa Sachum, 1853 were examined by light and electron microscopy. Each eye contains 2 400‐2 500 mostly hexagonal facets. However, irregularities affecting both shape and size of the ommatidia are relatively common, especially towards the margins of the eye. An individual ommatidium of this eucone type of apposition eye contains eight retinula cells, which give rise to a centrally‐fused, tiered rhabdom. The distal end of the latter is funnel‐shaped and accommodates the proximal end of the cone in its midst. Further below, the rhabdom (then formed by the rhabdomeres of four retinula cells) assumes a squarish profile with microvilli aligned in two directions at right‐angle to each other. Cross sections through the proximal regions of the rhabdom display triangular rhabdom outlines and microvilli (belonging to 3‐4 retinula cells different from those involved in the squarish more distal rhabdom) that run in three directions inclined to one another by 120°. Overall the organization of the eye conforms to the orthopteroid pattern and particularly closely resembles that of the American cockroach Periplaneta americana. However, since G. portentosa possesses fewer ommatidia, this could be a consequence of its inability to fly. On the other hand, the large size of the facets and the voluminous rhabdoms suggest considerable absolute sensitivity and an ability to detect the plane of linearly polarized light. Based on the pattern of microvillus orientations in combination with the crepuscular lifestyle G. portentosa leads and the habitat it occurs in, the prediction is made that this insect uses its green receptors for e‐vector discrimination in the environment of down‐welling light that reaches the forest floor.     

Larval morphology of Panorpodes kuandianensis (Insecta,Mecoptera, Panorpodidae) and its evolutionary implications

Larval characters play a significant role in evolutionary and systematic studies of holometabolous insects. However, Panorpodidae, a derived family of Mecoptera, are largely unknown in their immature stages to date. Here, the first instar larva of the short-faced scorpionfly Panorpodes kuandianensis Zhong, Zhang & Hua, 2011 is described and illustrated using light and scanning electron microscopy. The larva of Panorpodes is remarkable for the absence of compound eyes on the head and the presence of seven small unpaired proleg-like processes along the midventral line on abdominal segments II–VIII. The homology of these unpaired appendage-like processes, their ecological adaptation, and the evolutionary implications of some larval characters of Panorpodidae are discussed.     

亲土苔蛾成虫复眼超微结构

【目的】重叠型眼在昆虫复眼演化中起着重要作用。本研究旨在阐明夜出型亲土苔蛾Manulea affineola复眼类型及结构特征,以期填补灯蛾亚科昆虫复眼研究的空白,扩充夜出型昆虫复眼的特征数据,为探讨重叠型眼的变异趋势及复眼演化提供依据。【方法】运用光学和透射电子显微技术观察亲土苔蛾成虫复眼的超微结构。【结果】亲土苔蛾成虫复眼具有一个透明区,由6个次级色素细胞的透明胞质构成。小眼具8个视网膜细胞,其中1个视网膜细胞较短,仅位于小眼基部。在透明区内,7个视网膜细胞聚集成一束,其远端与晶体束末端相接,但并不形成视杆。在透明区下方,这7个视网膜细胞形成一个中心融合的视杆。在复眼背缘区的小眼的视杆具有近似矩形的横截面,而其余小眼的视杆具多分支状截面。【结论】亲土苔蛾成虫复眼属于重叠型眼;复眼背缘区的矩形视杆很可能与昆虫的偏振敏感性有关。     

刘氏蝎蛉卵巢管结构和卵子发生（英文）

卵巢管结构及卵子发生过程在探讨昆虫系统发育关系中有重要意义, 深入研究长翅目昆虫卵巢管结构及卵子发生可为确定其在全变态类昆虫中的系统发育地位提供依据。本文利用光学显微镜和扫描、透射电子显微镜技术研究了刘氏蝎蛉Panorpa liui Hua卵巢管超微结构及卵子发生过程。结果表明：蝎蛉卵巢由12根多滋式卵巢小管组成, 每个卵巢小管分为端丝、生殖区和生长区。根据滋养细胞、卵母细胞及滤泡细胞的变化, 卵子发生过程可分为5个阶段：卵黄发生前早期、卵黄发生前中期、卵黄发生前后期、卵黄发生期及卵壳形成期。在卵黄发生期, 滋养细胞为卵母细胞提供养分后逐渐消亡, 而此时的卵母细胞可通过滤泡之间的细胞间隙从血淋巴中获取营养。在卵壳形成期间, 3种不同类型的滤泡细胞参与形成不同区域的卵壳, 从而形成不同花饰的卵壳表面。据此推测, 与其他目的滋养细胞数目相比, 每个卵室中2次有丝分裂形成3个滋养细胞可能是比较原始的特征, 表明长翅目昆虫可能是全变态类群中近基部的分支。     

长蝎蛉幼期形态附生物学记述

【目的】蝎蛉科(Panorpidae)是长翅目(Mecoptera)最大的科,是重要的生态指示昆虫。然而,由于对环境条件要求苛刻,饲养困难,其幼期研究很不充分。【方法】本研究通过人工饲养成虫获得了长蝎蛉Panorpa macrostyla Hua的卵、幼虫和蛹等全部虫态,运用光学显微镜和扫描电子显微技术观察了其超微形态,并简要记载了其生物学特性。【结果】长蝎蛉每年发生1代,成虫发生于6月末至8月初。卵椭球形,卵壳表面覆盖一层隆起的网状结构。幼虫蠋型,具3对分4节的胸足和8对不分节的腹足;头壳高度骨化,具1对由26个小眼组成的复眼和1对3节的触角,口器咀嚼式;腹部第1-9节背面具有成对的背毛突,第10节仅有1根背毛突,腹部末端具有一个可伸缩的吸盘;呼吸系统为周气门式,具1对前胸气门和8对腹气门。幼虫共4个龄期,以预蛹期在土室内越冬。蛹为强颚离蛹,外形接近成虫,雄蛹腹部末端膨大。【结论】基于幼虫形态特征,长蝎蛉明显区别于新蝎蛉属Neopanorpa、华蝎蛉属Sinopanorpa、双角蝎蛉属Dicerapanorpa以及单角蝎蛉属Cerapanorpa幼虫。然而,长蝎蛉幼虫头部刚毛L2和SO2,腹部末节刚毛D2,SD1和SD2端部均膨大呈棒状,与蝎蛉属Panorpa其他种类区别明显,表明长蝎蛉的属级地位需要进一步研究。     

Molecular phylogeny of the scorpionflies Panorpidae (Insecta: Mecoptera) and chromosomal evolution

Panorpidae is the most species-rich family in Mecoptera with ca. 470 species in the Northern Hemisphere. However, the intergeneric phylogenetic relationships of Panorpidae remain unsatisfactorily resolved to date. Here, we used molecular and cytogenetic approaches to determine the phylogenetic relationships of Panorpidae in the evolutionary scenario of chromosomes, and estimated their divergence times using fossil-calibrated Bayesian analysis. In total, 89 species representing all seven genera of Panorpidae were used to reconstruct the phylogenetic trees using maximum parsimony, maximum likelihood and Bayesian inference based on the nuclear 28S rRNA and mitochondrial cox1 and cox2 genes. The results reveal that Panorpidae is a well-supported monophyletic group that can be categorized into two major clades. Major Clade I comprises Neopanorpa and Leptopanorpa, and Major Clade II consists of all the other genera (Cerapanorpa, Dicerapanorpa, Furcatopanorpa, Panorpa and Sinopanorpa). Neopanorpa and Cerapanorpa are regarded as paraphyletic groups for the first time. BEAST analysis indicates that Panorpidae originated in the Lower Cretaceous approximately 122.5 Ma (96.8–149.3 Ma), and that most diversification occurred from the Selandian (59.8 Ma) to the Middle Pleistocene (0.6 Ma) in the Cenozoic. Cytogenetic data plotted on the cladogram show that the lineage differentiation of Panorpidae is closely related to the chromosomal evolution, especially the reduction of chromosome number. Our study suggests that a taxonomic revision of Panorpidae is urgently needed at the generic level.