Phylogenetic position of the Strepsiptera: Review of molecular and morphological evidence

Molecular evidence of the monophyly of the Halteria (Strepsiptera + Diptera) is reviewed. The majority of morphological characters, which have classically been used to establish a Strepsiptera + Coleoptera sister group, are rejected, because they are based on erroneous interpretations of strepsipteran morphology. The scorings of 31 morphological characters, which directly relate to the phylogenetic position of Strepsiptera, are provided, and their distribution and optimization on the molecular + morphological tree is discussed. Of these, 13 characters specifically support the placement of Strepsiptera within the Mecopterida; seven of which are based on the optimization of inapplicable or missing data, and six of which are based on states that can be scored for Strepsiptera. Only a single character (posteromotorism) suggests a sister group relationship with the Coleoptera. The morphological and molecular data are largely congruent, and suggest that the Strepsiptera are sister group to the Diptera.     

The Strepsiptera problem: phylogeny of the holometabolous insect orders inferred from 18S and 28S ribosomal DNA sequences and morphology

Phylogenetic relationships among the holometabolous insect orders were inferred from cladistic analysis of nucleotide sequences of 18S ribosomal DNA (rDNA) (85 exemplars) and 28S rDNA (52 exemplars) and morphological characters. Exemplar outgroup taxa were Collembola (1 sequence), Archaeognatha (1), Ephemerida (1), Odonata (2), Plecoptera (2), Blattodea (1), Mantodea (1), Dermaptera (1), Orthoptera (1), Phasmatodea (1), Embioptera (1), Psocoptera (1), Phthiraptera (1), Hemiptera (4), and Thysanoptera (1). Exemplar ingroup taxa were Coleoptera: Archostemata (1), Adephaga (2), and Polyphaga (7); Megaloptera (1); Raphidioptera (1); Neuroptera (sensu stricto = Planipennia): Mantispoidea (2), Hemerobioidea (2), and Myrmeleontoidea (2); Hymenoptera: Symphyta (4) and Apocrita (19); Trichoptera: Hydropsychoidea (1) and Limnephiloidea (2); Lepidoptera: Ditrysia (3); Siphonaptera: Pulicoidea (1) and Ceratophylloidea (2); Mecoptera: Meropeidae (1), Boreidae (1), Panorpidae (1), and Bittacidae (2); Diptera: Nematocera (1), Brachycera (2), and Cyclorrhapha (1); and Strepsiptera: Corioxenidae (1), Myrmecolacidae (1), Elenchidae (1), and Stylopidae (3). We analyzed approximately 1 kilobase of 18S rDNA, starting 398 nucleotides downstream of the 5' end, and approximately 400 bp of 28S rDNA in expansion segment D3. Multiple alignment of the 18S and 28S sequences resulted in 1,116 nucleotide positions with 24 insert regions and 398 positions with 14 insert regions, respectively. All Strepsiptera and Neuroptera have large insert regions in 18S and 28S. The secondary structure of 18S insert 23 is composed of long stems that are GC rich in the basal Strepsiptera and AT rich in the more derived Strepsiptera. A matrix of 176 morphological characters was analyzed for holometabolous orders. Incongruence length difference tests indicate that the 28S + morphological data sets are incongruent but that 28S + 18S, 18S + morphology, and 28S + 18S + morphology fail to reject the hypothesis of congruence. Phylogenetic trees were generated by parsimony analysis, and clade robustness was evaluated by branch length, Bremer support, percentage of extra steps required to force paraphyly, and sensitivity analysis using the following parameters: gap weights, morphological character weights, methods of data set combination, removal of key taxa, and alignment region. The following are monophyletic under most or all combinations of parameter values: Holometabola, Polyphaga, Megaloptera + Raphidioptera, Neuroptera, Hymenoptera, Trichoptera, Lepidoptera, Amphiesmenoptera (Trichoptera + Lepidoptera), Siphonaptera, Siphonaptera + Mecoptera, Strepsiptera, Diptera, and Strepsiptera + Diptera (Halteria). Antliophora (Mecoptera + Diptera + Siphonaptera + Strepsiptera), Mecopterida (Antliophora + Amphiesmenoptera), and Hymenoptera + Mecopterida are supported in the majority of total evidence analyses. Mecoptera may be paraphyletic because Boreus is often placed as sister group to the fleas; hence, Siphonaptera may be subordinate within Mecoptera. The 18S sequences for Priacma (Coleoptera: Archostemata), Colpocaccus (Coleoptera: Adephaga), Agulla (Raphidioptera), and Corydalus (Megaloptera) are nearly identical, and Neuropterida are monophyletic only when those two beetle sequences are removed from the analysis. Coleoptera are therefore paraphyletic under almost all combinations of parameter values. Halteria and Amphiesmenoptera have high Bremer support values and long branch lengths. The data do not support placement of Strepsiptera outside of Holometabola nor as sister group to Coleoptera. We reject the notion that the monophyly of Halteria is due to long branch attraction because Strepsiptera and Diptera do not have the longest branches and there is phylogenetic congruence between molecules, across the entire parameter space, and between morphological and molecular data.     

Genomic and morphological evidence converge to resolve the enigma of Strepsiptera

The phylogeny of insects, one of the most spectacular radiations of life on earth, has received considerable attention. However, the evolutionary roots of one intriguing group of insects, the twisted-wing parasites (Strepsiptera), remain unclear despite centuries of study and debate. Strepsiptera exhibit exceptional larval developmental features, consistent with a predicted step from direct (hemimetabolous) larval development to complete metamorphosis that could have set the stage for the spectacular radiation of metamorphic (holometabolous) insects. Here we report the sequencing of a Strepsiptera genome and show that the analysis of sequence-based genomic data (comprising more than 18 million nucleotides from nearly 4,500 genes obtained from a total of 13 insect genomes), along with genomic metacharacters, clarifies the phylogenetic origin of Strepsiptera and sheds light on the evolution of holometabolous insect development. Our results provide overwhelming support for Strepsiptera as the closest living relatives of beetles (Coleoptera). They demonstrate that the larval developmental features of Strepsiptera, reminiscent of those of hemimetabolous insects, are the result of convergence. Our analyses solve the long-standing enigma of the evolutionary roots of Strepsiptera and reveal that the holometabolous mode of insect development is more malleable than previously thought.     

9-Genes Reinforce the Phylogeny of Holometabola and Yield Alternate Views on the Phylogenetic Placement of Strepsiptera

Background

The extraordinary morphology, reproductive and developmental biology, and behavioral ecology of twisted wing parasites (order Strepsiptera) have puzzled biologists for centuries. Even today, the phylogenetic position of these enigmatic “insects from outer space” [1] remains uncertain and contentious. Recent authors have argued for the placement of Strepsiptera within or as a close relative of beetles (order Coleoptera), as sister group of flies (order Diptera), or even outside of Holometabola.

Methodology/Principal Findings

Here, we combine data from several recent studies with new data (for a total of 9 nuclear genes and ∼13 kb of aligned data for 34 taxa), to help clarify the phylogenetic placement of Strepsiptera. Our results unequivocally support the monophyly of Neuropteroidea ( = Neuropterida + Coleoptera) + Strepsiptera, but recover Strepsiptera either derived from within polyphagan beetles (order Coleoptera), or in a position sister to Neuropterida. All other supra-ordinal- and ordinal-level relationships recovered with strong nodal support were consistent with most other recent studies.

Conclusions/Significance

These results, coupled with the recent proposed placement of Strepsiptera sister to Coleoptera, suggest that while the phylogenetic neighborhood of Strepsiptera has been identified, unequivocal placement to a specific branch within Neuropteroidea will require additional study.     

The thoracic skeleto-muscular system of Mengenilla (Strepsiptera: Mengenillidae) and its phylogenetic implications

The thorax of Mengenilla was examined using traditional morphological techniques and its features were documented in detail using scanning electron microscopy and computer-based 3D reconstructions. The results were compared to conditions found in other holometabolan insects. The implications for the systematic placement of Strepsiptera are discussed. The observations are interpreted in the light of the recently confirmed sistergroup relationship between Strepsiptera and Coleoptera (Coleopterida). The synapomorphies of the thorax of Strepsiptera and Coleoptera are partly related with posteromotorism (e.g., increased size of the metathorax), partly with a decreased intrathoracic flexibility (e.g., a fused pronotum and propleurum), and partly independent from these two character complexes (e.g., not connected profurca and propleuron). Strepsiptera are more derived than Coleoptera in some thoracic features (e.g., extremely enlarged metathorax) but have also preserved some plesiomorphic conditions (e.g., tegulae in both pterothoracic segments). All potential apomorphies of Mecopterida are missing in Strepsiptera. The last common ancestor of Coleopterida had already acquired posteromotorism but the wings were still largely unmodified. Several reductions in the mesothorax likely occurred independently.     

Phylogeny of the Strepsiptera based on morphological data of the first instar larvae

This investigation was the first cladistic analysis using morphological data of first instar larvae of Strepsiptera. The analysis of representatives of nearly all known families of Strepsiptera supports the division of Strepsiptera into Mengenillidia and Stylopidia. Corioxenidae and Elenchidae are placed at the base of Stylopidia. Halictophagidae is the sister group to Xeninae + Myrmecolacidae + Stylopinae. Xeninae is placed as the sister group to Myrmecolacidae + Stylopinae. Stylopidae are paraphyletic. Thus, Xenidae stat. n. is re-established. A sister-group relationship between Myrmecolacidae and Elenchidae is not supported on characters of first instar larvae.     

The morphology and terminology of the hindwing articulation and wing base of the Coleoptera, with specific reference to the Scarabaeoidea

Abstract. Characters of the hindwing articulation and wing base are important for contributing to the solution of phylogenetic and systematic problems in the Coleoptera. In the Scarabaeoidea morphological terms proposed by previous authors do not cover many structures in sufficient detail and additional terms are needed to describe and utilize all characters used in systematic considerations; these can be used for all Coleoptera.
In this paper we identify new structures, the first basal plate and the second basal plate (two subdivisions of the coleopteran wing base), name the various yokes, braces and reinforcements found on them and propose names for various projections, lobes, indentations and embayments on the axillary sclerites (first axillary, head, neck and tail; second axillary, arm and body; third axillary, prong; basalare).     

Head structures of males of Strepsiptera (Hexapoda) with emphasis on basal splitting events within the order

Internal and external head structures of males of Strepsiptera were examined and the head of a species of Mengenilla is described in detail. The results suggest a reinterpretation of some structures. The head of basal extant strepsipterans is subprognathous, whereas it is strictly orthognathous in the groundplan of Strepsiptera s.l. The labrum and hypopharynx are not part of the mouthfield sclerite. The labial palps are absent in all strepsipterans. A very slightly modified mandibular articulation is preserved in Eoxenos, whereas it is distinctly reduced in other extant groups. A salivary duct, salivary glands, and a cephalic aorta are absent. The cladistic analysis of 44 characters of the head results in the following branching pattern: (Protoxenos + (Mengea + (Eoxenos + (Mengenilla [Austr.] + Mengenilla) + (Elenchus + Dundoxenos + Xenos + Stylops)))). Most apomorphies of males are associated with the necessity of finding females within a short time span and with a reduced necessity to consume food: large "raspberry" eyes, flabellate antennae with numerous dome-shaped chemoreceptors, Hofeneder's organ, an ovoid sensillum of the maxillary palp, and the simplified condition of the maxilla and the labium. Strepsiptera excl. Protoxenos are supported by the dorsomedian frontal impression, the dorsally shifted antennal insertions, a reduced number of antennal segments, absence of the galea, and probably by the presence of the mouthfield sclerite, which is a unique apomorphic feature. The balloon-gut combined with an unusual air-uptake apparatus is another possible autapomorphy of this clade. It is likely that the last common ancestor of Strepsiptera excl. Protoxenos did not process food. Strepsiptera s.str. are characterized by the strongly reduced condition of the labrum and the absence of the epistomal suture. Eoxenos is the sister group of the remaining Strepsiptera s.str. Synapomorphies of Mengenilla + Stylopidia are the advanced reduction of the mandibular articulation and the secondary absence of the ovoid sensillum. The monophyly of Mengenilla is confirmed, even though a small free labrum is present in Australian species. Derived features of Stylopidia are the absence of the coronal suture and the reduced condition of the frontal suture. Apomorphies that have evolved within Stylopidia are the membranization of parts of the head, the fusion of antennal segments, the increase or decrease of the number of flabellate flagellomeres, reductions and modifications of the mandibles, and modifications of the mouthfield sclerite. The monophyly of Stylopiformia is not unambiguously supported. A position of the mandibles posterior to the mouthfield sclerite (when adducted) is a possible synapomorphy shared by Xenos, Stylops, and other "higher Stylopidia." The blade-like distal part of the mandibles suggests a closer relationship of Elenchus with these taxa.     

The morphology and evolution of the female postabdomen of Holometabola (Insecta)

In the present article homology issues, character evolution and phylogenetic implications related to the female postabdomen of the holometabolan insects are discussed, based on an earlier analysis of a comprehensive morphological data set. Hymenoptera, the sistergroup of the remaining Holometabola, are the only group where the females have retained a fully developed primary ovipositor of the lepismatid type. There are no characters of the female abdomen supporting a clade Coleopterida + Neuropterida. The invagination of the terminal segments is an autapomorphy of Coleoptera. The ovipositor is substantially modified in Raphidioptera and distinctly reduced in Megaloptera and Neuroptera. The entire female abdomen is extremely simplified in Strepsiptera. The postabdomen is tapering posteriorly in Mecopterida and retractile in a telescopic manner (oviscapt). The paired ventral sclerites of segments VIII and IX are preserved, but valvifers and valvulae are not distinguishable. In Amphiesmenoptera sclerotizations derived from the ventral appendages VIII are fused ventromedially, forming a solid plate, and the appendages IX are reduced. The terminal segments are fused and form a terminal unit which bears the genital opening subapically. The presence of two pairs of apophyses and the related protraction of the terminal unit by muscle force are additional autapomorphies, as is the fusion of the rectum with the posterior part of the genital chamber (cloaca). Antliophora are supported by the presence of a transverse muscle between the ventral sclerites of segment VIII. Secondary egg laying tubes have evolved independently within Boreidae (absent in Caurinus) and in Tipulomorpha. The loss of two muscle associated with the genital chamber are likely autapomorphies of Diptera. The secondary loss of the telescopic retractability of the postabdomen is one of many autapomorphies of Siphonaptera.     

Goodbye Halteria? The thoracic morphology of Endopterygota (Insecta) and its phylogenetic implications

Characters of the thorax of 30 representatives of all endopterygote orders and four hemimetabolous outgroup taxa were examined. In total, 126 characters potentially useful for phylogenetic reconstruction are discussed and presented as a data matrix. The thoracic features were analysed with different approaches combined with an additional large set of morphological data. Endopterygota were confirmed as monophyletic and new morphological autapomorphies of the group are suggested. The highly controversial Strepsiptera are not placed as sistergroup of Diptera (Halteria‐concept) but consistently as sistergroup of Coleoptera. This clade was mainly supported by characters associated with posteromotorism. The traditionally proposed relationship of Neuropterida + Coleoptera was not confirmed. Hymenoptera was placed as sistergroup of all remaining orders in parsimony analyses. The inclusion of Strepsiptera + Coleoptera in Mecopterida in parsimony analyses is probably artificial and potential thoracic autapomorphies of Mecopterida in the traditional sense are suggested. Mecopterida are confirmed as a clade in Bayesian analyses. Amphiesmenoptera and Antliophora are well supported. The paraphyly of Mecoptera is due to a clade comprising Nannochoristidae and Siphonaptera + Diptera. The phylogenetic reconstruction using characters of the thorax is impeded by functional constraints, parallel losses, a general trend to reinforce the skeleton and to simplify the muscular apparatus, and also by different specializations occurring in potential outgroup taxa. The addition of a large additional morphological data set only partly compensated for these problems. It is apparent that the inclusion of more outgroup and ingroup taxa is required, notably presumably basal representatives of Mecoptera, Trichoptera, and Diptera. This may reduce the effect of an artificial attraction of branches caused by homoplasy, notably character losses occurring within different lineages.© The Willi Hennig Society 2010.     

The beetle tree of life reveals that Coleoptera survived end‐Permian mass extinction to diversify during the Cretaceous terrestrial revolution

Here we present a phylogeny of beetles (Insecta: Coleoptera) based on DNA sequence data from eight nuclear genes, including six single‐copy nuclear protein‐coding genes, for 367 species representing 172 of 183 extant families. Our results refine existing knowledge of relationships among major groups of beetles. Strepsiptera was confirmed as sister to Coleoptera and each of the suborders of Coleoptera was recovered as monophyletic. Interrelationships among the suborders, namely Polyphaga (Adephaga (Archostemata, Myxophaga)), in our study differ from previous studies. Adephaga comprised two clades corresponding to Hydradephaga and Geadephaga. The series and superfamilies of Polyphaga were mostly monophyletic. The traditional Cucujoidea were recovered in three distantly related clades. Lymexyloidea was recovered within Tenebrionoidea. Several of the series and superfamilies of Polyphaga received moderate to maximal clade support in most analyses, for example Buprestoidea, Chrysomeloidea, Coccinelloidea, Cucujiformia, Curculionoidea, Dascilloidea, Elateroidea, Histeroidea and Hydrophiloidea. However, many of the relationships within Polyphaga lacked compatible resolution under maximum‐likelihood and Bayesian inference, and/or lacked consistently strong nodal support. Overall, we recovered slightly younger estimated divergence times than previous studies for most groups of beetles. The ordinal split between Coleoptera and Strepsiptera was estimated to have occurred in the Early Permian. Crown Coleoptera appeared in the Late Permian, and only one or two lineages survived the end‐Permian mass extinction, with stem group representatives of all four suborders appearing by the end of the Triassic. The basal split in Polyphaga was estimated to have occurred in the Triassic, with the stem groups of most series and superfamilies originating during the Triassic or Jurassic. Most extant families of beetles were estimated to have Cretaceous origins. Overall, Coleoptera experienced an increase in diversification rate compared to the rest of Neuropteroidea. Furthermore, 10 family‐level clades, all in suborder Polyphaga, were identified as having experienced significant increases in diversification rate. These include most beetle species with phytophagous habits, but also several groups not typically or primarily associated with plants. Most of these groups originated in the Cretaceous, which is also when a majority of the most species‐rich beetle families first appeared. An additional 12 clades showed evidence for significant decreases in diversification rate. These clades are species‐poor in the Modern fauna, but collectively exhibit diverse trophic habits. The apparent success of beetles, as measured by species numbers, may result from their associations with widespread and diverse substrates – especially plants, but also including fungi, wood and leaf litter – but what facilitated these associations in the first place or has allowed these associations to flourish likely varies within and between lineages. Our results provide a uniquely well‐resolved temporal and phylogenetic framework for studying patterns of innovation and diversification in Coleoptera, and a foundation for further sampling and resolution of the beetle tree of life.     

Bahiaxenidae, a "living fossil" and a new family of Strepsiptera (Hexapoda) discovered in Brazil

An adult male of a newly discovered strepsipteran species from Brazil— Bahiaxenos relictus— is described. A new family Bahiaxenidae is suggested based on cladistic analyses of comprehensive morphological data sets with a broad taxon sampling including the stem group. It is unambiguously placed as the sister group of all other extant families of Strepsiptera. Bahiaxenos relictus is the only species of basal, i.e. non-stylopidian, Strepsiptera occurring in the New World. It appears to be a relict taxon that has survived in the fossil sand dunes of the São Francisco River (Bahia State). The loss of the 8th antennomere and the greatly reduced labrum are autapomorphies of Strepsiptera s.s . excluding Bahiaxenidae. The sister group relationship between†Protoxenidae and the remaining Strepsiptera, and between † Cretostylops and a clade comprising † Mengea and Strepsiptera s.s. , is confirmed, as is the monophyly of Stylopidia and Stylopiformia.     

The embryonic development of Stylops ovinae (Strepsiptera,Stylopidae) with emphasis on external morphology

External features of the embryonic development of Stylops ovinae (Strepsiptera) were examined. Eighteen distinct embryological stages are suggested. Many embryological traits are closely correlated to the parasitic life style of the first instar larvae or to vivipary. The high number of eggs, their small size, the characteristic egg membrane, and the lack of micropyles are derived groundplan features of Strepsiptera. The development with a semi-long germ embryo is shared with several other groups of Holometabola. The reduction of the labrum and antennae are autapomorphies of Strepsiptera. The cephalic ventral plate of the first instar larva of S. ovinae is formed by parts of the head capsule and the anlagen of the maxillae and labium. It is involved in the formation of the specific entognathous condition, and the entire character complex is autapomorphic for Stylopidae. The trochanter is recognizable in the anlagen of all three legs. Its fusion with the femur in the later stages is an autapomorphy of Stylopidia. The extreme spiralization and compression of the abdomen during blastokinesis is a derived feature, like the reduction of the anlagen of the anterior abdominal appendages. The caudal bristles on segment XI are possibly re-activated cerci. The same is likely in the case of segment XI.     

†Kinzelbachilla ellenbergeri – a new ancestral species,genus and family of Strepsiptera (Insecta)

A single male specimen of a new species (†Kinzelbachilla ellenbergeri gen. et sp.n. ) of a new family of the endoparasitic Strepsiptera (†Kinzelbachillidae fam.n.) from Burmese amber is described and evaluated with respect to its systematic placement. Its features come very close to the presumptive groundplan of the order suggested in recent studies. Preserved plesiomorphic features are the following: fully sclerotized head with long coronal suture, small ommatidia not separated by chitinous bridges, absence of microtrichia between ommatidia, eight antennomeres, robust mandibles with dicondylic articulation, galea distinctly developed, free pro‐ and mesotrochanters, slender five‐segmented tarsi without adhesive soles, and equally sclerotized abdominal tergites and sternites. An important character that is not recognizable due to damage is the shape of the metapostnotum. This structure is transverse in the groundplan of Strepsiptera and in †Protoxenos, but elongated and shield‐like in all other known strepsipterans. In a cladistic analyses of 82 characters of adult males and additional characters for females and immatures (scored as unknown for all included fossils) †Kinzelbachilla is placed as sister group of all remaining strepsipterans except for †Protoxenos, followed by †Cretostylops and †Mengea as the third and fourth branches in the stem group, respectively. The position of †Protoxenos as first branch is suggested by three unambiguous apomorphic features of all remaining Strepsiptera, the reduced size of less than 6 mm, mandibles distinctly narrowing distad the basalmost part, and fan‐shaped hindwings which are broader than they are long. The hitherto known fossil stem group strepsipterans do not distinctly narrow the large morphological gap separating this order from its sister group, the Coleoptera. This published work has been registered in ZooBank, http://zoobank.org/urn:lsid:zoobank.org:pub:07554C01-DEC3-4080-A337-B1F46BC9070F .     

Phylogenetic Relationships of the Suborders of Coleoptera (Insecta)

One hundred seven external and internal characters of larval and adult representatives of 28 genera of the coleopteran suborders were analyzed cladistically. Four groups of Neuropterida were introduced as outgroup. The analysis yielded 18 trees with a minimum of 194 steps (CI 0.691). All trees support the monophyly of all four suborders and a branching pattern (Archostemata + (Adephaga + (Myxophaga + Polyphaga))). The presence of elytra with meso- and metathoracic locking devices, the specific hind-wing folding, the close connection of exposed sclerites, the absence of the mera, the absence of eight thoracic muscles, the reduced abdominal sternite I, and the invagination of terminal segments are autapomorphies of Coleoptera. The monophyly of Coleoptera excl. Archostemata is supported by further transformations of the thoracic sclerites such as absence of the mesothoracic discriminal line and katepisternal joint, by an internalized or absent metathoracic trochantin, by the presence of a bending zone in the hind-wing, and by eight further muscle losses. Fusion of tibia and tarsus and presence of a single claw are larval synapomorphies of Myxophaga and Polyphaga. Adults are characterized by fusion of protrochantin and propleura and by the rigid connection of the meso- and metathoracic ventrites. The eucinetoid lineage of Polyphaga is characterized by the secondary absence of the bending zone of the alae. This results in a distinctly simplified wing folding mechanism. The monophyly of Cucujiformia (+ Bostrichoidea) is supported by the presence of cryptonephric Malpighian tubules. Transformations of fore-and hind-wings, reinforcement and simplification of the thoracic exoskeleton, and an efficient use of a distinctly reduced set of thoracic muscles play an important role in the early evolution of Coleoptera. Many different larval character transformations take place in the earlier Mesozoic within the suborders.     

鞘翅目系统演化关系研究进展

现生鞘翅目类群是昆虫纲中种类最多的一个类群,由4个亚目组成。形态学研究表明,鞘翅目是一个单系,但在加入了分子数据之后,其结果证明鞘翅目为并系类群。最早的甲虫起源于二叠纪,与现生的原鞘亚目具有许多相似特征。大多数学者认为与鞘翅目亲缘关系最近的类群是脉翅类昆虫,但是也有的学者认为鞘翅目与捻翅目互为姐妹群。本文对鞘翅目高级阶元4个亚目的系统关系和有关鞘翅目的起源和演化等研究进行了总结,并对相关问题做了进一步探讨和展望。     

The evolution of Strepsiptera (Hexapoda)

An evolutionary scenario for the enigmatic group Strepsiptera is provided, based on the results of a comprehensive cladistic analysis of characters of all life stages. A recently described fossil--+Protoxenos janzeni--the most archaic strepsipteran, sheds new light on the early evolution of the group and reduces the "morphological gap" between Strepsiptera and other insects. It weakens both current hypotheses--Coleoptera+Strepsiptera and Diptera+Strepsiptera (="Halteria"). The splitting into +Protoxenos (Protoxenidae) and the remaining Strepsiptera was linked with a distinct size reduction and many morphological changes. Unlike males of extant strepsipteran species, +Protoxenos was still able to process food. Mengeidae (+Mengea), with two small species, is the sister group of extant Strepsiptera. A unique characteristic of extant males (Strepsiptera s. str.) is the mouthfield sclerite. It is part of an air uptake apparatus which belongs to an extremely modified air-filled "balloon gut". Besides this, male strepsipterans possess specialised antennae and compound eyes, a strongly developed flight apparatus, large testes, and a sperm pump, whereas other organ systems are strongly reduced (e.g., fat body, malpighian tubules). Males are designed to find females within a few hours and to copulate. A dramatic change is linked with the split into Mengenillidae and Stylopidia. The change to pterygote hosts and the permanent endoparasitism of the females are evolutionary novelties acquired by the latter clade, and linked with far-reaching morphological transformations, e.g. the presence of unique brood organs. Hairy tarsal adhesive devices are present in males and guarantee efficient attachment to the host during copulation. A well-founded clade within Stylopidia is Stylopiformia, which are characterised by a unique fissure-shaped birth opening. The evolutionary development towards the most specialised and successful forms (parasites of aculeate Hymenoptera [e.g., Xenidae+Stylopidae], ca. 46% of the species) is a stepwise process. The presented evolutionary scenario comprises a complex network of functionally correlated morphological changes in primary larvae, secondary larvae, females and males.