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.     

Embryogenesis of a springtail,Tomocerus ishibashii (Collembola,tomoceridae): External morphology

The external features of the developing embryos of the springtail, Tomocerus ishibashii, are described. The clypeolabral anlage arises as a single, unpaired swelling. The entognathy is completed by the ventral growth of the tergal anlagen of mandibular, maxillary, and labial segments. These anlagen also form the posterior part of the cranium. The palpi of maxilla and labium are homologous with the telopodites, and proximal parts of these head appendages are homologous with the coxopodites. The sternal element of the labial segment does not participate in the postmentum formation. The anlagen of abdominal appendages appear in the first to the fourth abdominal segments. The first, third, and fourth appendage anlagen form the ventral tube, tenaculum, and furcula, respectively. The fused proximal parts of the first, third, and fourth appendage anlagen are homologous with the coxopodites, and the distal parts which do not fuse are homologous with the telopodites. The anlagen of the second abdominal appendages become flattened and spread over the ventral side of this segment. The ventral structures of the first four abdominal segments are appendicular in origin.     

The First Complete 3D Reconstruction of a Spanish Fly Primary Larva (Lytta vesicatoria,Meloidae, Coleoptera)

The first detailed anatomical study of a primary larva of Meloidae is presented. Thereby techniques such as three-dimensional reconstructions, microtome sections, SEM (scanning electronic microscopy) and CLSM (confocal laser scanning microscopy) are applied. The structural features are discussed in the context of phylogeny, but also possible correlations with parasitism, phoresy and miniaturisation. The triungulin first instar larva is likely an apomorphy of Meloidae excl. Eleticinae and linked with a specialisation on acridoid eggs or larvae and provisions of bees. The campodeid body shape of Lytta and Meloinae is a groundplan feature of Meloidae, whereas a navicular body is an autapomorphy of the generally phoretic larvae of Nemognathinae. Head structures of Lytta and features of the postcephalic body are largely plesiomorphic. The musculature of the head is only moderately simplified while the one of the postcephalic body is well developed. Its thorax is largely characterised by plesiomorphies. The characteristics of the legs suggest phoretic habits, even though this does not apply to larvae of Lytta. It is conceivable that a phoretic behaviour is secondarily lost, together with some but not all morphological modifications related to it. Derived features of the abdomen of Meloidae are the complete loss of the fixed urogomphi (also missing in Rhipiphoridae and other related groups) and the presence of one or two conspicuous caudal bristles. Only few features of Lytta are shared with the parasitic larvae of Rhipiphoridae and Strepsiptera. These characteristics, which are possibly linked with specialised life habits, have obviously evolved independently. Miniaturisation effects are minimal in the larvae of Lytta.     

Morphogenesis of body segments and appendages during the larval stages of a common spiroboloid milliped

Three-dimensional, histological, and x-ray techniques provide a picture of body segment and limb morphogenesis. Cell multiplication begins in the proliferation region (“meristem”) during the molt from the preceding instar. By four days post-molt, the cells that will form the new segments are well under way in their anterior, lateral, and dorsal migration. It is suggested that after the anlagen for all the new segments are estabilshed, a mitotic wave commences in the most anterior anlagen and moves posteriorly during the remainder of the instar. When cell proliferation is complete, final differentiation of the segments takes place. The process of limb formation is one of cell proliferation and perhaps migration. Each limb develops in a membranous pocket during the instar following the one in which its respective body segment was formed.     

Embryonic development of the scorpionfly Panorpa emarginata Cheng with special reference to external morphology (Mecoptera: Panorpidae)

The Mecoptera are thought to be one of the most primitive groups in the Holometabola, but their embryology is rarely studied. By means of scanning electron microscopy, we studied the external features of the embryo of the scorpionfly Panorpa emarginata in middle and late development. The embryo remains in the superficial position until hatching. Embryonic development can be divided into 10 stages along with the first‐instar larva. The external features are described from the germ band to the first‐instar larva, with special reference to the components and segmentation of the head, the segmentation of abdomen and the formation of abdominal prolegs. Our results confirm that the head consists of an anterior‐most acron and six trunk segments: the labral, antennal, intercalary, mandibular, maxillary, and labial segments. The labrum is confirmed to derive from the paired appendages. Our observations also provide additional direct evidence that the abdominal prolegs are not serially homologous with the thoracic legs. The presence of the eleventh abdominal segment is clarified. J. Morphol. 2009. © 2009 Wiley‐Liss, Inc.     

Postembryonic development and homology of external genitalia in Galloisiana nipponensis (Caudell et king) (Notoptera : Grylloblattidae)

The postembryonic development of genital and postgenital segments of Galloisiana nipponensis (Notoptera : Grylloblattidae) have been described in detail. The coxite, stylus, epiproct and paraproct are already observed in the first nymphal instar, but the sexual characters are almost indistinguishable in the 1st to 4th nymphal instars, because the genital segments in these stages are not yet differentiated. In the 5th nymphal instar in the male, the paired primary phallic lobes start to appear at the hind margin of the 9th sternum. The rudiments of ventral valvulae in the female originate at the hind margin of the 8th sternum. In the 6th nymphal instar, the rudiments of inner valvulae begin to appear on the 9th sternum, in addition to the ventral valvula rudiments. The long dorsal valvulae are formed from the extension of nymphal coxite, while the nymphal styli are more reduced than those of younger instars.The ventral valvulae of the 8th abdominal segment are thought to be homologous with the inner valvulae of the 9th segment, judging from their site and mode of development. Further, the inner valvulae may also be homologous with the primary phallic lobes (phallomeres) of the male. The external genitalia in G. nipponensis originate from 2 structures: the dorsal valvulae (considered of appendicular origin), and the phallomeres; ventral and inner valvulae are regarded as mere integumentary outgrowths.     

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.     

Parasitism of Western Corn Rootworm Larvae and Pupae by Steinernema carpocapsae

Virulence and development of the insect-parasitic nematode, Steinernema carpocapsae (Weiser) (Mexican strain), were evaluated for the immature stages of the western corn rootworm, Diabrotica virgifera virgifera LeConte. Third instar rootworm larvae were five times more susceptible to nematode infection than second instar larvae and 75 times more susceptible than first instar larvae and pupae, based on laboratory bioassays. Rootworm eggs were not susceptible. Nematode development was observed in all susceptible rootworm stages, but a complete life cycle was observed only in second and third instar larvae and pupae. Nematode size was affected by rootworm stage; the smallest infective-stage nematodes were recovered from second instar rootworm larvae. Results of this study suggest that S. carpocapsae should be applied when second and third instar rootworm larvae are predominant in the field.     

COMPARTMENTALIZATION OF THE DEVELOPING MACRONUCLEUS FOLLOWING CONJUGATION IN STYLONYCHIA AND EUPLOTES

The development of the macronucleus following conjugation in the hypotrichous ciliates Euplotes and Stylonychia has been examined with the electron microscope. Banded polytene chromosomes can be seen in thin sections of the macronuclear anlagen during the early periods of exconjugant development. As the chromosomes reach their maximum state of polyteny, sheets of fibrous material appear between the chromosomes and transect the chromosomes in the interband regions. Individual bands of the polytene chromosomes thus appear to be isolated in separate compartments. Subsequently, during the stage when the bulk of the polytenic DNA is degraded (1), these compartments swell, resulting in a nucleus packed with thousands of separate spherical chambers. Individual chromosomes are no longer discernible. The anlagen retain this compartmentalized condition for several hours, at the end of which time aggregates of dense material form within many of the compartments. The partitioning layers disperse shortly before replication bands appear within the elongating anlagen, initiating the second period of DNA synthesis characteristic of macronuclear development in these hypotrichs. The evidence presented here suggests that the "chromatin granules" seen in the mature vegetative macronucleus represent the material of single bands of the polytene chromosomes seen during the earlier stages of macronuclear development. The possibility is also discussed that the degradation of DNA in the polytene chromosomes may be genetically selective, which would result in a somatic macronucleus with a different genetic constitution than that of the micronucleus from which it was derived.     

A scanning electron microscopy study of the embryonic development of Pycnogonum litorale (Arthropoda,Pycnogonida)

The phylogenetic position of the enigmatic Pycnogonida (sea spiders) is still controversial. This is in part due to a lack of detailed data about the morphology and ontogenesis of this, in many aspects, aberrant group. In particular, studies on the embryonic development of pycnogonids are rare and in part contradictory. Here, we present the first embryological study of a pycnogonid species using scanning electron microscopy (SEM). We describe the late embryogenesis of Pycnogonum litorale from the first visible appendage anlagen to the hatchling in 11 embryonic stages. The three pairs of appendage anlagen gain in length by growth, as well as by extension of furrows into the embryo. The opening of the stomodaeum is located far in front of the anlagen of the chelifores and has a Y‐shaped lumen from the onset. During further embryogenesis, the position of the mouth shifts ventrally, until it is located between the chelifores. The proboscis anlage grows out as a circumoral wall‐like structure, which is initially more pronounced ventrally. Hypotheses about the evolution of the proboscis by fusion of originally separated components are critically discussed, because the proboscis anlage of P. litorale shows no indications of a composite nature. In particular, a participation of post‐cheliforal elements in proboscis formation is rejected by our data. Further, no preoral structure and no stage in proboscis formation was found, which could plausibly be homologized with the labrum of othereuarthropods. Thus, our study supports the assumption of a complete lack of a labrum in Pycnogonida. J. Morphol., 2010. © 2010 Wiley‐Liss, Inc.     

The phylogeny of Strepsiptera (Hexapoda)

Previous phylogenetic analyses of Strepsiptera have been limited to characters from only males or first instar larvae, and by poor taxonomic sampling. This investigation is the first cladistic analysis to use more than fourfold as many characters as any prior study, and a broader sampling of taxa. The analysis of 189 morphological characters of all stages of representatives of all extant strepsipteran families and characters of adult males of amber fossils results in the following branching pattern: (?Protoxenos+ (?Cretostylops + (?Mengea + (Mengenillidae + (Corioxenidae + (Bohartillidae + (Halictophagidae + (Elenchidae + (?Protelencholax + (Myrmecolacidae + (Callipharixenidae + (Xenidae + Stylopidae)))))))))))). The basal placement of the Baltic amber fossil ?Protoxenos and the Burmese amber fossil ?Cretostylops is well founded. Even though ?Cretostylops is older than ?Protoxenos it is almost certainly not the most basal strepsipteran group but the sister group of a clade comprising the Baltic amber fossil ?Mengea + Strepsiptera s. str. (excl. stemlineage). Monophyly of Mengenillidae, Stylopidia, Stylopiformia s.l., Corioxenidae, Xenidae, and Stylopidae is confirmed. Mengenillidia is paraphyletic (with respect to ?Mengea (Mengeidae)), Elenchidae (with respect to ?Protelencholax) and the genus Stichotrema (with respect to the Baltic amber fossils). Thus Protelencholacidae fam. n. is described, and S. weitschati and S. triangulum are transferred to Palaeomyrmecolax. A ground plan of adult male Strepsiptera is provided and evolutionary interpretations are presented based on the obtained cladograms. © The Willi Hennig Society 2005.     

Tracking down the "head blob": comparative analysis of wingless expression in the developing insect procephalon reveals progressive reduction of embryonic visual system patterning in higher insects

The evolution of larval head morphology in holometabolous insects is characterized by reduction of antennal appendages and the visual system components. Little insight has been gained into molecular developmental changes underlying this morphological diversification. Here we compare the expression of the segment polarity gene wingless (wg) in the pregnathal head of fruit fly, flour beetle and grasshopper embryos. We provide evidence that wg activity contributes to segment border formation, and, subsequently, the separation of the visual system and protocerebrum anlagen in the anterior procephalon. In directly developing insects like grasshopper, seven expression domains are formed during this process. The activation of four of these, which correspond to polar expression pairs in the optic lobe anlagen and the protocerebral ectoderm, has shifted to postembryonic stages in flour beetle and Drosophila. The remaining three domains map to the protocerebral neuroectoderm, and form by disintegration of a large precursor domain in flour beetle and grasshopper. In Drosophila, the precursor domain remains intact, constituting the previously described “head blob”. These data document major changes in the expression of an early patterning gene correlated with the dramatic evolution of embryonic visual system development in the Holometabola.     

稻虱跗煽的个体发生研究：卵的形成和胚胎发育

捻翅目昆虫是胎生的,胚胎发育和孵化均在母体血腔内进行。稻虱跗蝇Elenchinus japonicus属捻翅目,跗煽科,寄生于白背稻虱、褐稻虱和灰稻虱。本文报道稻虱跗煽卵的形成各阶段：1)雌虫体内无典型的卵巢,所有卵在母体体腔内同步发育和成熟。最早发现的原卵是包囊干细胞,在雌幼虫血腔内。2)每个包囊细胞内含256个姐妹细胞,其中有一个细胞分化成卵母细胞,其余的成为营养细胞。3)成熟卵为椭圆形,大小为95-100X40-50μm。其胚胎发育过程按顺序包括：卵裂、胎盘形成、胚带分节、附肢形成和胚胎背合等阶段。稻虱跗蛔行单胚生殖。     

Prey stage preference and feeding behaviour ofOligota kashmirica benefica (Coleoptera: Staphylinidae), an insect predator of the spider miteTetranychus urticae (Acari: Tetranychidae)

We studied the prey stage preference and feeding behaviour of the first to third instar larvae and adult females ofOligota kashmirica benefica Naomi (Coleoptera: Staphylinidae), a predator of the spider miteTetranychus urticae Koch (red form) (Acari: Tetranychidae), on leaves of the kudzu vine (Pueraria lobata (Wild.) Ohwi (Leguminosae)) under laboratory conditions. The number of mites eaten increased with the growth of predator larvae. Third instar larvae preyed on all stages of spider mite, whereas first instar larvae preyed mainly on immobile stages (eggs and quiescent stages). The predator larvae showed two types of foraging behaviour (active searching and ambush behaviour) when targeting the mobile stages (larval nymph and adult stages of prey). Although no difference was found in the number of prey consumed by adult females and third instar larvae of the predator, the adult females mainly attacked and consumed the immobile stages.     

Numerical allometric growth of the ommatidia,antennal sensilla,and teeth of foretibial combs in the milkweed bug Oncopeltus fasciatus Dallas (Heteroptera : Lygaeidae)

Oncopeltus fasciatus (Heteroptera : Lygaeidae), the milkweed bug, was bred in captivity. Sampling showed that individuals grow exponentially through their 6 developmental stages with an average linear increase per molt of 42% for the females and 41% for the males. The number of ommatidia per eye grows with negative allometry from an average of 30 in the first instar to 860 in the male and 820 in the female adult. The total number of sensilla on the 2 flagellar segments of an antenna increases with negative allometry during the 5 nymphal stages from a mean of 239 in the first instar to 2462 in the last. At this point, this allometric growth pattern is sharply broken by distinct numerical increase to 7163 on the adult flagellum. The number of teeth composing the foretibial comb, the tool for grooming the distal flagellar segment, grows with negative allometry through all 6 developmental stages. Calculations using previously published data from the migratory locust, Locusta migratoria, revealed the same growth pattern of antennal sensilla: uniform allometric growth during the nymphal development, broken by a conspicuous upward jump to the adult number of sensilla. In the American cockroach, Periplaneta americana, this growth pattern of antennal sensilla holds only for the male; the female continues the nymphal allometric growth into the adult stage. These observations on allometric growth fit three theoretical explanations: 1. Smooth allometric growth is evidence for an aut-adaptation to increasing size. 2. Ex-adaptations to novel ecological niches cause breaks in allometric growth patterns. 3. Chapman's rule, which states that increased mobility correlates with greater olfactory sensitivity, correctly predicts the observed breaks in the allometric growth patterns in the abundance of antennal sensilla.