Tracheal ultrastructure of protura (Insecta : Apterygota)

The tracheal systems of Sinentomon and Eosentomon (Apterygota : Protura) were examined in thin sections and compared with the tracheae of collembolan, Allacma. The tracheal system of Protura consists of spiracles and tracheae. The spiracle is a simple, concave cuticular cavity known as an atrium. A globular chamber is present between the atrium and trachea. The atrium of Eosentomon is decorated with ridges and has 2 small openings to tracheal recesses beside the central tracheal opening. The tracheae of Protura are characterized by a high frequency of taenidia and the absence of intima folds and intertaenidial spaces. The taenidia of Sinentomon have a rectangular section and those of Eosentomon are gable-shaped. The results also suggest that the tracheal recess of Eosentomon is a kind of stigmatic gland. The tracheal structure of Protura was compared with that of collembolans, insects, and other arthropods, and discussed in terms of phylogeny.     

Aflagellated spermatozoa of Huhentomon and Acerella (Protura : Apterygota)

The spermatozoa of Huhentomon plicatunguis (Protura : Hesperentomidae) and Acerella tiarnea (Protura : Acerellidae) are aflagellate and of simple structure. An ovoidal nucleus with condensed chromatin is located at the end of the cell; the cytoplasm is devoid of organelles, except for mitochondria, and contains compact material derived from centriolar adjunct-like material. Centriole and mitochondrial cristae degenerate during spermiogenesis. These results show that the evolutionary trends of proturan spermatozoa tend towards aflagellate state.     

从弹尾纲和原尾纲的亲缘关系质疑缺尾纲（=近昆虫纲）的有效性

鉴于弹尾目（跳虫）和原尾目的尾部都没有尾须（cerci）,Brner于1910年就把这两类归并为一个类群缺尾纲（Ellipura）,这一分类阶元长期被许多昆虫学家沿用至今。Kukalová-Peck(1987 )在讨论化石双尾虫（?）附肢的总体结构（ground plan）时,认为跳虫和原尾虫的腹部侧板更原始,附肢无转节,将二者归纳成近昆虫纲（Parainsecta）。但是从形态特征、内部结构、比较精子学、变态类型和胚后发育等的特点以及线粒体DNA和核糖体DNA的测序分析结果,显示弹尾纲与原尾纲之间存在诸多重要差异,不具备较为密切的亲缘关系,我们不支持（弹尾纲+原尾纲）组成缺尾纲或近昆虫纲。据此建议取消缺尾纲（=近昆虫纲）这一分类阶元。     

Protura in native and exotic forests in the North Island of New Zealand

Abstract

This study provides data on population and community ecology of Protura in native forests and Pinus radiata plantations in New Zealand. Abundance, age structure, sex ratios, biodiversity, and relationship with soil chemistry are discussed. Protura were significantly more abundant in pine plantations in comparison to native forests. Among native forests, Protura were most abundant under Southern beech (Nothofagus solandri). The abundance patterns may reflect the association between Protura and fungal communities in the soil. No correlation was found between soil organic matter, nitrogen, phosphorous, pH, cation exchange capacity, and the density of Protura. Protura assemblage composition was significantly related to forest type. New Zealand endemic species were associated with native forests; species with distribution outside New Zealand dominated in pine plantations. The distribution records within New Zealand were expanded for five species. The Protura fauna of New Zealand was increased to 18 species. Berberentulus capensis, Eosentomon australicum and Australentulus tillyardi are new records for New Zealand fauna.     

Systematic and biogeographical study of Protura (Hexapoda) in Russian Far East: new data on high endemism of the group

Proturan collections from Magadan Oblast, Khabarovsk Krai, Primorsky Krai, and Sakhalin Oblast are reported here. Twenty-five species are found of which 13 species are new records for Russian Far East which enrich the knowledge of Protura known for this area. Three new species Baculentulus krabbensis sp. n., Fjellbergella lazovskiensis sp. n. and Yichunentulus alpatovi sp. n. are illustrated and described. The new materials of Imadateiella sharovi (Martynova, 1977) are studied and described in details. Two new combinations, Yichunentulus borealis (Nakamura, 2004), comb. n. and Fjellbergella jilinensis (Wu & Yin, 2007), comb. n. are proposed as a result of morphological examination. Keys to species of the genera Fjellbergella and Yichunentulus are given. An annotated list of all species of Protura from Russian Far East is provided and discussed. Widely distributed species were not recorded in this area. This may be because of the high sensitivity of Protura to anthropogenic impact and low dispersal ability of the group.     

The gut structure of Sinentomon erythanum yin (Protura : Sinentomidae)

The fine structure of the midgut, pyloric region, Malpighian papillae, and hindgut of Sinentomon erythranum (Protura : Sinentomidae) is described. Midgut cells are rich in mineral concretions and are presumably involved in excretory activity; the pyloric chamber, a cavity in the proturan intestine behind the midgut, is formed by cells with microvilli pointing anteriorly; the secretion from 6 Malpighian papillae flows into this cavity. The hindgut consists of 2 regions; the anterior of the 2 has a series of specializations typical of cells engaged in active water reabsorption. Long infoldings of the apical plasma membrane reach deep into the cells. The findings are compared with the gut organization of other genera of Protura examined to date.     

原尾纲重新分群的特征分析(六足总纲)

1996年第20届国际昆虫学大会（意大利佛罗伦萨）上,尹文英根据多年研究的新发现提出了原尾纲Protura分为3目10科的新建议,仅刊登在该次会议的论文摘要中,至今尚未能详细阐述重新分群的重要特征和分析。本文拟就原尾纲不同类群的成虫和幼虫主要特征进行分析和比较,结果证明原尾纲分为3目（Yuan目、华Yuan目和古Yuan目）10科,更符合原尾虫客观发展的自然规律。     

河南伏牛山自然保护区原尾虫群落结构的研究

１９９６－０７对河南伏牛山自然保护区土壤动物进行调查,发现原尾虫９种,隶属于４科６属,其中天目巴元和高绳线毛元为群落优势中。ＥＢＰ－分析表明原尾虫群落多为Ｂ型。群落多样性以落叶阔叶林为最高,针叶林次之,草地最低。     

上海大金山岛原尾虫的物种多样性及生态特征

原尾虫是土壤动物的重要类群之一,然而目前对其物种多样性及生态特征的了解仍然十分有限,岛屿生态系统中原尾虫的研究尤为欠缺。为深入探讨原尾虫的栖息地选择、空间分布、种群密度、繁殖策略及其与土壤真菌的相互作用等生态特征,于2015年秋季在大金山岛竹林、乔木林和灌木林等3种不同的植被中进行了定量采样研究。结果表明,大金山岛共发现8种原尾虫,隶属2科4属,这些物种在上海市陆域植被中均有分布,且均为东洋界种类;大金山岛秋季原尾虫的总密度为竹林(696头/m~2)乔木林(479头/m~2)灌木林(213头/m~2);竹林中原尾虫的Shannon-Wiener多样性指数显著高于灌木林(P0.05),而不同植被类型中原尾虫的Pielou均匀度、Simpson优势度和Margalef丰富度等指数无显著差异;大金山岛的原尾虫在土壤中为隔离分布,具有在适宜的微生境中集中分布的特点;发现原尾虫在野外受到病原真菌感染的直接证据;通过对原尾虫的性比和幼虫比的分析,发现其繁殖可能采取介于k对策和r对策之间的生态策略。     

The phylogenetic positions of three Basal-hexapod groups (protura, diplura, and collembola) based on ribosomal RNA gene sequences

This study combined complete 18S with partial 28S ribosomal RNA gene sequences ( approximately 2,000 nt in total) to investigate the relations of basal hexapods. Ten species of Protura, 12 of Diplura, and 10 of Collembola (representing all subgroups of these three clades) were sequenced, along with 5 true insects and 8 other arthropods, which served as out-groups. Trees were constructed with maximum parsimony, maximum likelihood, Bayesian analysis, and minimum-evolution analysis of LogDet-transformed distances. All methods yielded strong support for a clade of Protura plus Diplura, here named Nonoculata, and for monophyly of the Diplura. Parametric-bootstrapping analysis showed our data to be inconsistent with previous hypotheses (P < 0.01) that joined Protura with Collembola (Ellipura), that said Diplura are sister to true insects or are diphyletic, and that said Collembola are not hexapods. That is, our data are consistent with hexapod monophyly and Collembola grouped weakly with "Protura + Diplura" under most analytical conditions. As a caveat to the above conclusions, the sequences showed nonstationarity of nucleotide frequencies across taxa, so the CG-rich sequences of the diplurans and proturans may have grouped together artifactually; however, the fact that the LogDet method supported this group lessens this possibility. Within the basal hexapod groups, where nucleotide frequencies were stationary, traditional taxonomic subgroups generally were recovered: i.e., within Protura, the Eosentomata and Acerentomata (but Sinentomata was not monophyletic); within Collembola, the Arthropleona, Poduromorpha, and Entomobryomorpha (but Symphypleona was polyphyletic); and in Diplura, the most complete data set (> 2,100 nt) showed monophyly of Campodeoidea and of Japygoidea, and most methods united Projapygoidea with Japygoidea.     

Flagellate spermatozoa of Protura (Insecta,Apterygota) are motile

A new model of sperm axoneme with 16 + 0 doublets is described. The spermatozoon of Acerentulus confinis (Apterygota : Protura) has a short conical acrosome, a long helicoidal nucleus, well-developed centriolar adjunct material, and a long flagellum. Using fixation with a glutaraldehyde-tannic acid mixture, without osmium post-fixation, doublet protofilaments, inner dynein arms, radial spokes, nexin bridges, and Y-links of the sperm axoneme of A. confinis and Acerentomon italicum were clearly observed. Optical observation shows that the proturan flagellate spermatozoa are motile cells. The process involving the transformation of the spermatozoa from a coiled to an elongated swimming form was studied by scanning electron microscope. The findings confirmed that flagellar motility is due to the presence of a single dynein arm on doublets in spite of the unusual axonemal pattern.     

Brain organization in Collembola (springtails)

Arthropoda is comprised of four major taxa: Hexapoda, Crustacea, Myriapoda and Chelicerata. Although this classification is widely accepted, there is still some debate about the internal relationships of these groups. In particular, the phylogenetic position of Collembola remains enigmatic. Some molecular studies place Collembola into a close relationship to Protura and Diplura within the monophyletic Hexapoda, but this placement is not universally accepted, as Collembola is also regarded as either the sister group to Branchiopoda (a crustacean taxon) or to Pancrustacea (crustaceans + hexapods). To contribute to the current debate on the phylogenetic position of Collembola, we examined the brains in three collembolan species: Folsomia candida, Protaphorura armata and Tetrodontophora bielanensis, using antennal backfills, series of semi-thin sections, and immunostaining technique with several antisera, in conjunction with confocal laser scanning microscopy and three-dimensional reconstructions. We identified several neuroanatomical structures in the collembolan brain, including a fan-shaped central body showing a columnar organization, a protocerebral bridge, one pair of antennal lobes with 20-30 spheroidal glomeruli each, and a structure, which we interpret as a simply organized mushroom body. The results of our neuroanatomical study are consistent with the phylogenetic position of Collembola within the Hexapoda and do not contradict the hypothesis of a close relationship of Collembola, Protura and Diplura.     

上海东佘山竹林土壤中原尾虫区系和生态调查

自首次在欧洲发现原尾虫以来,迄今已有七十余年的历史,在此期间虽然五大洲均有原尾虫的发现,但有关生态方面的报道却很少见。偶或在土壤昆虫区系研究中涉及一些,往往非常笼统,大多只有一项“原尾目”,连分科的资料也没有。直到今立源太良(Imadate,1973)才做了出色的报道,Tuxen(1978)也发表了在巴西亚马逊河流域调查的情况。     

低等六足动物系统学研究进展

低等六足动物包括原尾纲、弹尾纲和双尾纲三个类群,是探讨六足动物起源和进化问题的关键类群,近十年来成为节肢动物系统进化研究中的焦点之一。低等六足动物的系统发育地位以及它们之间的关系一直是备受争论的问题。通过介绍三类低等六足动物最新的分类系统,从经典分类学和系统发育两个方面对低等六足动物近十年来的研究进展进行了综述。迄今,对于三类低等六足动物都建立了比较完备的分类体系,原尾纲划分为3目10科,弹尾纲划分为4目30科,双尾纲划分为2亚目3总科10科。系统发育研究中,大多数的系统发育分析结果不支持传统的缺尾类假说,缺尾纲应摒弃不用。分子数据分析的结果普遍支持原尾纲与双尾纲近缘,但仍需要进一步探讨。线粒体基因组、比较胚胎学和比较精子学的研究结果表明,原尾纲可能经历了长期的趋异进化历史。最近的比较精子学研究支持了双尾纲的单系性。总之,三类低等六足动物系统学研究均取得了长足的发展,但仍然存在诸如研究人员匮乏和研究水平不均衡等问题。系统发育研究中,分子系统学研究成为关注的焦点,而基于核基因和线粒体基因的数据分别建立的系统发育假说存在分歧,亟需开发更优的数据分析方法。此外,需加强低等六足动物比较形态学、比较胚胎学、发育生物学等方面的研究,以便将来进行全证据的系统发育研究。     

Is Ellipura monophyletic? A combined analysis of basal hexapod relationships with emphasis on the origin of insects

Hexapoda includes 33 commonly recognized orders, most of them insects. Ongoing controversy concerns the grouping of Protura and Collembola as a taxon Ellipura, the monophyly of Diplura, a single or multiple origins of entognathy, and the monophyly or paraphyly of the silverfish (Lepidotrichidae and Zygentoma s.s.) with respect to other dicondylous insects. Here we analyze relationships among basal hexapod orders via a cladistic analysis of sequence data for five molecular markers and 189 morphological characters in a simultaneous analysis framework using myriapod and crustacean outgroups. Using a sensitivity analysis approach and testing for stability, the most congruent parameters resolve Tricholepidion as sister group to the remaining Dicondylia, whereas most suboptimal parameter sets group Tricholepidion with Zygentoma. Stable hypotheses include the monophyly of Diplura, and a sister group relationship between Diplura and Protura, contradicting the Ellipura hypothesis. Hexapod monophyly is contradicted by an alliance between Collembola, Crustacea and Ectognatha (i.e., exclusive of Diplura and Protura) in molecular and combined analyses.     

Protura of Italy, with a key to species and their distribution

The Italian Protura were studied basing on 5103 specimens from 198 sampling areas, along with bibliographic data from 49 collecting sites. 17 out of the 20 Italian regions are covered. As a result, 40 species have been identified (belonging to 8 genera and 4 families), 6 of which are new records for the Italian fauna.A key to the Italian species is reported, followed by a series of distribution maps and brief remarks for some of them. A preliminary biogeographical overview allowed us to delineate the chorological categories of these species, 10 of which are actually known only in Italy. The comparison with the species richness known for some best studied Central and Eastern European Countries leads us to speculate that widening our research, Italian Protura check-list will be much implemented.     

Testing for misleading effects in the phylogenetic reconstruction of ancient lineages of hexapods: influence of character dependence and character choice in analyses of 28S rRNA sequences

The present analyses employ the almost complete sequence of the 28S rRNA gene to investigate phylogenetic relationships among Pancrustacea, placing special emphasis on the position of basal hexapod lineages. This study utilizes a greater number of characters and taxa of Protura, Collembola and Diplura than previous analyses to focus on conflicts in the reconstruction of the early steps in hexapod evolution. Phylogenetic trees are mainly based on Bayesian approaches, but likewise include analyses with Maximum Likelihood and Maximum Parsimony. Different analyses, including the application of a mixed DNA/RNA substitution model, were performed to narrow possible misleading effects of non-stationarity of nucleotide frequencies, saturation and character independence down to a minimum. This is the first time that a mixed DNA/RNA model is applied to analyse 28S rRNA sequences of basal hexapods. All methods yielded strong support for the monophyly of Collembola, Diplura, Dicondylia and Insecta s.str. , as well as for a cluster composed of Diplura and Protura ('Nonoculata-hypothesis'). However, the last cluster may be an artifact caused by a shared GC bias of the 28S sequences between these orders, in combination with a long branch effect. The instability of the position of the 'Nonoculata' within Pancrustacea further bears out the misleading effect of non-stationarity of nucleotide frequencies. Protura and Diplura either form the sister-group to Collembola (Entognatha) or cluster with branchiopod crustaceans. Overall, the phylogenetic signal of the complete sequences of the 28S rRNA gene favours monophyly of Hexapoda over paraphyly. However, further corroboration from independent data is needed to rule out the competing hypothesis of mutually paraphyletic Crustacea and Hexapoda.     

Possible Involvement of Phage-Like Structures in Antagonism of Cowpea Rhizobia by Rhizobium trifolii

A reduction in the viability of cowpea rhizobia was observed when Rhizobium trifolii IARI and cowpea Rhizobium strain 3824 were inoculated together in soil. The reduction in number of cowpea rhizobia in soil was found to be associated with the reduction in number of nodules per plant and retardation in plant growth. An antimicrobial substance was isolated from R. trifolii which, on electron microscopic investigation, demonstrated the presence of several phage-like structures.     

Cell junctions in the gut of Protura

The main cell junctions in the intestinal tract of a small group of apterygotan insects, Protura, were examined in conventional thin sections, tracer-infiltrated sections and freeze-fracture replicas. The smooth septate junctions in the midgut of collembolan Tomocerus minor were also studied for comparison. Pleated septate junctions are found in foregut, hindgut and Malpighian papillae. They exhibit regular septa crossing the intercellular clefts in thin sections; and the septa with a pronounced zig-zag appearance run parallel to form a honeycomb structure in tracer-impregnated sections. After freeze-fracture undulating rows of intramembranous particles (IMPs) are visible on the P face. Smooth septate juncions are observed in the midgut. The interceullar septa often run in pairs for long tracts and exhibit a wavy course in lanthanum impregnated sections. The IMPs exhibited on the E face are usually separated one from another. Twin arrangement of particle rows is also apparent on the replicas. Gap junctions are frequent in both the midgut and hindgut and possess the conventional characteristics of 'inverted gap junction' with E face connexons. These results provide further evidence relating Protura closely to Collembola as well as to primitive arthropods.