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.     

The sperm pumps of Strepsiptera and Antliophora (Hexapoda)

Male genital structures of representatives of Strepsiptera, Siphonaptera and Diptera are described in detail, with special emphasis on sperm pumps. The parts involved in the apparatus are evaluated with respect to their homology. Functional interpretations are presented based on the morphological observations. The phylogenetic significance of characters related to the male genital apparatus is discussed. The sperm pumps differ strongly in Strepsiptera and Antliophora (s.s.) and are not homologous. The strepsipteran type, which lacks any sclerotized parts, has evolved independently. Autapomophies of the male genital apparatus are the compact testes, the large balloon‐shaped vesicula seminalis, the strongly developed musculature of the proximal ductus ejaculatorius, the strongly simplified copulatory organ, the unusual muscles of segments VIII and IX, and the complete absence of accessory glands. The median fusion and almost globular shape of the vesicula are potential autapomorphies of Corioxenidae. The absence of the furrow separating the testes from the vesicula seminalis is a derived condition found in Xenos and Myrmecolax. A sperm pump is absent in Boreus (Mecoptera) and Culicomorpha and the functionally relevant parts and their arrangement differ strongly in Siphonaptera, Pistillifera and Diptera (excl. Culicomorpha). The presence of a functional and homologous pumping apparatus does not belong to the groundplan of Antliophora, which implies that this alleged autapomorphy of the clade is invalid. A sperm pump belongs to the groundplan of Diptera and was secondarily reduced in Culicomorpha, many representatives of Bibionomorpha, and in Diopsidae. It was very likely primarily absent in Mecoptera. However, the precise reconstruction of the groundplan depends on the position of Nannochoristidae within Mecoptera and on the possible affinities of Siphonaptera and Boreidae. Sperm pumps should be considered as a functional term and not be used as a character for phylogenetic reconstruction, unless specific similarities are included in the character definition.     

Rediscovered parasitism of Andrena savignyi Spinola (Hymenoptera,Andrenidae) by Stylops (Strepsiptera,Stylopidae) and revised taxonomic status of the parasite

Parasitism of Andrena (Suandrena) savignyi Spinola (Hymenoptera: Andrenidae) by Stylops Kirby (Strepsiptera: Stylopidae) has been recorded only once, and from an individual collected in Egypt almost a century ago, with the parasite described as Stylops savignyi Hofeneder. The recent rediscovery of this Stylops from an individual of Andrena savignyi permits a reinterpretation of the species and its affinities among other Stylops. The bee was collected at flowers of Zilla spinosa (Turra) Prantl. (Brassicaceae) in Amariah, Riyadh, Kingdom of Saudi Arabia. Based on DNA barcode sequences from material sampled across Africa, Asia, and Europe, it is apparent that Stylops savignyi is conspecific with Stylops nassonowi Pierce, and we accordingly synonymize this name (syn. n.), with the latter representing the senior and valid name for the species. A differential diagnosis is provided for Stylops nassonowi and the morphology of the female is described, as well as the first instars.     

Zum Feinbau des Komplexauges von Stylops spec. (Insecta,Strepsiptera)

Zusammenfassung Unter den Cornealinsen des Komplexauges von Stylops befindet sich ein Kristallkegel vom pseudoconen Typ, der von zahlreichen Pigmentzellen umhüllt wird. An seinem proximalen Ende liegen 6 meist pigmentfreie Zellen (Sempersche Zellen).Das Ommatidium besteht aus etwa 60 Retinulazellen. Ihre distal kranzartig miteinander verbundenen Mikrovillisäume bilden ein einziges offenes Rhabdom, das extrazelluläres (?) granuläres Material und die Basis der Semperschen Zellen umgibt. Stellenweise wird das Rhabdom samt granulärem Material von homogen erscheinenden distalen Ausläufern einzelner Retinulazellen überlagert. Proximad zerfällt das Rhabdom zunehmend in kleinere Rhabdomteile. Im zentralen Teil des Ommatidiums liegen 1–2 auffallend große Retinulazellen, die meist weniger elektronendicht erscheinen und kleinere Pigmentgrana haben.Die einzelnen Ommatidien werden von ungemein zahlreichen, sehr pigmentarmen Stützzellen umhüllt. Diese werden — wie die basalen Teile der Retinulazellen — teilweise durch Gliazellfortsätze isoliert.Bei Stylops, einem Vertreter der Strepsipteren, handelt es sich nicht um ocelläre Komplexaugen (Strohm, 1910), auch nicht um eucone Ommatidien (Kinzelbach, 1967), sondern um Ommatidien vom pseudoconen Typ. Zumindest der Bau des Rhabdoms ähnelt dem des Larvenauges (Stemma), dessen rezeptorischer Teil entgegen den Annahmen früherer Autoren in der Imago nicht reduziert wird.

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On the fine structure of the compound eye of Stylops spec. (Insecta, Strepsiptera)

Summary In the compound eye of Stylops a crystalline cone of the pseudocone type is found beneath the corneal lens. It is enveloped by several pigment cells. At the proximal part of the cone there are 6 cells (Semper cells) mostly pigment-free.The ommatidium consists of approximately 60 retinula cells. Their rhabdomeres distally rim-like connected to another form a single open rhabdom which encircles extracellular granular material as well as the bases of the Semper cells. Here and there the rhabdom plus granular material is overlain with distal protrusions of single retinula cells which appear to be homogeneous. Towards the proximal part the rhabdom increasingly divides up into smaller rhabdomal segments. One or two conspicuous large retinula cells were found in the central part of the ommatidium, appearing to be less electron-dense and containing pigment granules of a smaller size. Each ommatidium is surrounded by numerous cells (Stützzellen) lacking in pigment. These cells are partially insulated from another—as well as the basal parts of retinula cells—by protrusions of glia cells.Our investigations show that the eyes of Stylops (as a representative of Strepsiptera) are not of the ocellar complex eye type. At least the structure of the rhabdom resembles to that of the larval eye (stemma), the receptor part of which is not reduced in the imago.

Herrn Prof. Dr. Helmcke danke ich für die freundliche Unterstützung am Raster-Elektronenmikroskop.     

The spermatozoon of Mengenilla moldrzyki (Strepsiptera,Mengenillidae): Ultrastructure and phylogenetic considerations

Even though the spermatozoa of several strepsipteran species were described earlier, no data were available for the basal family Mengenillidae. Well-fixed material of the recently described Tunisian species Mengenilla moldrzyki was used for a detailed examination of the sperm ultrastructure. The total length is c. 30 μm. The head region contains a conical acrosome vesicle (0.3-0.35 μm) and an elongated nucleus (7.3 μm) with dense chromatin. Some granular material along with a uniformely dense centriole adjunct and two mitochondrial derivatives are visible at the posterior end of the nucleus. The material of the centriole adjunct does not extend along the flagellum and accessory bodies are absent. The mitochondrial derivatives are elongated structures crossed by a longitudinal crista but lacking parallel transverse cristae and paracrystalline material in the dense matrix. The mitochondrial derivatives gradually reduce their size and end at the most posterior tail region. The flagellar axoneme has a 9 + 9 + 2 pattern and originates beneath the nucleus. In the terminal tail region the axoneme gradually disintegrates. Despite the extreme specialization of the endoparasitc group, strepsipteran spermatozoa are mostly characterized by plesiomorphies. The pattern within the order is largely uniform, but Mengenilla displays several apomorphic features compared to the presumptive strepsipteran groundplan (e.g., absence of crystallizations and cristae in the mitochondrial derivatives). The subdivision of the intertubular material into two compartments with a dense beak-like structure adhering to the tubular wall supports a clade Coleopterida (=Strepsiptera + Coleoptera) + Neuropterida.     

The ultrastructure of the preformed ecdysial 'line of weakness' in the puparium cap of Elenchus tenuicornis (Kirby) (Insecta: Strepsiptera).

This study describes the ultrastructure at 'the line of weakness' in the male puparium of Elenchus tenuicornis (Kirby) (Insecta: Strepsiptera). Superficially this line looked like an area of untanned cuticle but ultrastructurally it had an undifferentiated epicuticle, an untanned exocuticle and a loose textured endocuticle. It is speculated that the E. tenuicornis pharate adult male prior to emergence smears a chemical solution on the inner rim of the 'line of weakness' which dissolves the exo- and endocuticles. Following this, the slightest pressure exerted by the ptilinum of the male breaks open the cap of the puparium.     

Effects of Nu-Serum® on in vitro development of goat (Capra hircus) embryos

To evaluate the effects of a commercially produced serum substitute on the in vitro development of caprine embryos, registered Nubian doelings were synchronized with norgestomet-impregnated implants (Synchromate-B®: CEVA) and superovulated with descending doses of FSH-p® (Schering). A total of 246 embryos was collected and placed in Tissue Culture Medium 199 (TCM 199, Gibco Laboratories) containing Nu-Serum® (NuS) at concentrations of 2.5%, 5.0%, 10%, or 20%. Control treatments consisted of TCM 199 alone or TCM 199 plus 10% heat-inactivated fetal bovine serum (FBS). Embryos developed in all concentrations of NuS to the morula, blastocyst, and expanded blastocyst stages. The TCM 199 plus 10% NuS had significantly higher percentages of embryos developing to the expanded blastocyst stage than TCM 199 plus 10% FBS. Time to expanded blastocyst development in NuS was shorter than in the TCM 199 plus FBS. No stage-specific block to development was observed with embryos collected and cultured in vitro for any of the treatments. These results demonstrate that NuS, when compared to FBS, allowed a higher percentage (P < 0.05) of caprine embryos to develop to the expanded blastocyst stage, thus providing a valuable substitute for FBS.     

The embryonic development of the rhabdocoel flatworm Mesostoma lingua (Abildgaard, 1789)

The embryonic development of the flatworm Mesostoma lingua was studied using a combination of life observation and histological analysis of wholemount preparations and sections (viewed by both light and electron microscopy.) We introduce a series of stages defined by easily recognizable morphological criteria. These stages are also applicable to other platyhelminth taxa that are currently under investigation in our laboratory. During cleavage (stages 1 and 2), the embryo is located in the center of the egg, surrounded by a layer of yolk cells. After cleavage, the embryo forms a solid, disc-shaped cell cluster. During stage 3, the embryo migrates to the periphery of the egg and acquires bilateral symmetry. The side where it contacts the egg surface corresponds to the future ventral surface of the embryo. Stage 4 is the emergence of the first organ primordia, the brain and pharynx. Gastrulation, as usually defined by the appearance of germ layers, does not exist in Mesos-toma; instead, organ primordia emerge ”in situ” from a mesenchymal mass of cells. Organogenesis takes place during stages 5 and 6. Cells at the ventral surface form the epidermal epithelium; inner cells differentiate into neurons, somatic and pharyngeal muscle cells, as well as the pharyngeal and protonephridial (excretory) epithelium. A junctional complex, consisting initially of small septate junctions, followed later by a more apically located zonula adherens, is formed in all epithelial tissues at stage 6. Beginning towards the end of stage 6 and continuing throughout stages 7 and 8, cytodifferentiation of the different organ systems takes place. Stage 7 is characterized by the appearance of eye pigmentation, brain condensation and spindle-shaped myocytes. Stage 8 describes the fully dorsally closed and differentiated embryo. Muscular contraction moves the body in the egg shell. We discuss Mesostoma embryogenesis in comparison to other animal phyla. Particular attention is given to the apparent absence of gastrulation and the formation of the epithelial junctional complex. Received: 10 February 2000 / Accepted: 10 April 2000     

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.     

Developmental morphology of the Asian one-leaf plant, Monophyllaea glabra (Gesneriaceae) with emphasis on inflorescence morphology

We examined the developmental morphology of the tropical Asian one-leaf plant Monophyllaea glabra, which is believed to have diverged first in the phylogenetic tree of the genus. The embryo within the seed consists of two cotyledons and a hypocotyl with no shoot or root apical meristems. The endogenous root meristem is formed nearer the hypocotyl end than in other examined Monophyllaea species. One of the cotyledons grows to form the macrocotyledon by means of the basal meristem. The groove meristem arises between the anisocotyledons, shifts toward the macrocotyledon, and is transformed to the inflorescence apex, which produces inflorescence axes in the axils of all ventral bracts of two rows, and secondary inflorescences in the axils of the lower dorsal bracts of the other two rows. The macrocotyledon may act as a ventral bract for the first inflorescence axis at the reproductive stage. This organization suggests that a common ancestor of Monophyllaea and Whytockia with decussate inflorescences diverged in one direction to become Monophyllaea and in another to become Whytockia.     

Apolysis without ecdysis in larval development of a strepsipteran, Elenchus tenuicornis (Kirby)

Previous uncertainty concerning the number of larval stages in a strepsipteran ( Elenchus tenuicornis ) was resolved by transmission and scanning electron microscope observations. The first larval instar (triungulinid) is free-living: after entering the host (a delphacid) it undergoes a normal ecdysis. The resulting second instar larva undergoes two further apolyses without ecdysis. In the male, the cuticle of the fourth instar larva tans to form the puparium, while the female becomes sexually mature without a further moult after extrusion of the cephalothorax through the host cuticle. The discarded exuviae of the triungulinid remain in the host abdomen; subsequent larval staging is made possible by the retention of the exuviae ensheathing the larva as each apolysis is accomplished.     

Pollen morphology of Microstachys (Euphorbiaceae) with emphasis on neotropical species

Abstract

In Euphorbiaceae tribe Hippomaneae is the pantropical genus Microstachys, comprised by 24 species, mostly from Brazil. Palynological studies with several representatives of the genus are scarce. The purpose of this study was to describe the pollen morphology of Microstachys and find possible differentiating characteristics between its species. Ten species of the genus were analysed, represented by 21 herbarium collections. Their pollen grains were acetolysed, measured and photographed under light microscopy (LM) and scanning electron microscopy (SEM). The analysed species possess pollen in monads, of small to medium size, isopolars, amb distincticly trilobed, oblate-spheroidal, prolate-spheroidal or subprolate, tricolporate with narrow and very long colpi, almost fused at the poles sometimes forming an invagination with tapering ends and margo of the colporus wide to very wide and psilate; endoapertures lalongate to very lalongate with tapering ends, presence of a narrow costae distinct in the endoaperture; exine microreticulate homobrochate, walls with simple columella; at the mesocolpus the sexine separates from the nexine forming cavea; and sexine always thicker than the nexine. Microstachys is a stenopalynous genus, despite its species varying in pollen size, shape and width of the cavea, margo of the colporus, and length of the endoapertures, demonstrating taxonomic value, and indicating a contribution towards future systematic analyses.     

Smaller nests of the hornet Vespa analis (Hymenoptera, Vespidae) are more severely affected by the strepsipteran parasite Xenos moutoni (Strepsiptera, Stylopidae) than are larger nests

The parasite Xenos moutoni inactivates workers and castrates reproductives of the Japanese hornet Vespa analis. Using over 500 nests, we investigated the relationship between hornet nest size (number of cells) and the proportion of parasitized adults (i.e., prevalence) in central Japan. Over 3 years, 36–48% of nests had more than one parasitized adult, and approximately 3–5% of female adults (mostly workers) and 1–3% of male adults were parasitized within the population. The number of cells did not differ between nests with and without parasitized workers; therefore, the effects of parasite-caused losses within the worker force on nest size appear to be negligible. However, the prevalence of parasitized workers tended to decrease with an increase in the number of cells. Our analysis indicated that the number of parasitized workers was consistently low and did not increase with nest size, resulting in a relatively high prevalence in small nests.     

External morphology of limb development in the amphipod <Emphasis Type="Italic">Orchestia cavimana</Emphasis> (Crustacea,Malacostraca, Peracarida)

The external morphology of limb development in Orchestia cavimana is examined by scanning electron microscopy and fluorescence staining from the appearance of the first limb buds until hatching. As other amphipods, O. cavimana undergoes direct development and the degree of segmental differentiation shows a more or less continual decrease in anteroposterior direction. Limbs form ventrally as small buds, which elongate and divide into podomeres early in development. This early subdivision largely corresponds with the limb segmentation of the hatchling. When the post-naupliar limbs start to develop, the germ band begins to split into two halves along the midline, so that the trunk limbs transiently occupy a very dorsolateral position. After the germ band has closed again, the differentiation into the characteristic amphipodan tagmata (cephalothorax, pereon, pleon) takes place and the limb podomeres lose their round-shape. The late embryo is covered by a so-called intermediate cuticle, which is formed after an embryonic moult and shed after hatching. The early development of O. cavimana reveals the Anlage of a vestigial seventh pleonic segment that is assumed to belong to the ground pattern of malacostracans, but is retained as a free, limbless segment only in adult Leptostraca. A transient subdivision of the proximal segment of the pleopods suggests the occurrence of a coxa and a basis in these limbs. The mandible attains its upright, adult position via a characteristic bending process that is strikingly similar to that in Archaeognatha (Insecta).     

Formation of the hindgut cuticular lining during embryonic development of Porcellio scaber (Crustacea,Isopoda)

The hindgut and foregut in terrestrial isopod crustaceans are ectodermal parts of the digestive system and are lined by cuticle, an apical extracellular matrix secreted by epithelial cells. Morphogenesis of the digestive system was reported in previous studies, but differentiation of the gut cuticle was not followed in detail. This study is focused on ultrastructural analyses of hindgut apical matrices and cuticle in selected intramarsupial developmental stages of the terrestrial isopod Porcellio scaber in comparison to adult animals to obtain data on the hindgut cuticular lining differentiation. Our results show that in late embryos of stages 16 and 18 the apical matrix in the hindgut consists of loose material overlaid by a thin intensely ruffled electron dense lamina facing the lumen. The ultrastructural resemblance to the embryonic epidermal matrices described in several arthropods suggests a common principle in chitinous matrix differentiation. The hindgut matrix in the prehatching embryo of stage 19 shows characteristics of the hindgut cuticle, specifically alignment to the apical epithelial surface and a prominent electron dense layer of epicuticle. In the preceding embryonic stage – stage 18 – an electron dense lamina, closely apposed to the apical cell membrane, is evident and is considered as the first epicuticle formation. In marsupial mancae the advanced features of the hindgut cuticle and epithelium are evident: a more prominent epicuticular layer, formation of cuticular spines and an extensive apical labyrinth. In comparison to the hindgut cuticle of adults, the hindgut cuticle of marsupial manca and in particular the electron dense epicuticular layer are much thinner and the difference between cuticle architecture in the anterior chamber and in the papillate region is not yet distinguishable. Differences from the hindgut cuticle in adults imply not fully developed structure and function of the hindgut cuticle in marsupial manca, possibly related also to different environments, as mancae develop in marsupial fluid. Bacteria, evenly distributed within the homogenous electron dense material in the hindgut lumen, were observed only in one specimen of early marsupial manca. The morphological features of gut cuticle renewal are evident in the late marsupial mancae, and are similar to those observed in the exoskeleton.     

The embryonic development of the triclad <Emphasis Type="Italic">Schmidtea polychroa</Emphasis>

Triclad flatworms are well studied for their regenerative properties, yet little is known about their embryonic development. We here describe the embryonic development of the triclad Schmidtea polychroa, using histological and immunocytochemical analysis of whole-mount preparations and sections. During early cleavage (stage 1), yolk cells fuse and enclose the zygote into a syncytium. The zygote divides into blastomeres that dissociate and migrate into the syncytium. During stage 2, a subset of blastomeres differentiate into a transient embryonic epidermis that surrounds the yolk syncytium, and an embryonic pharynx. Other blastomeres divide as a scattered population of cells in the syncytium. During stage 3, the embryonic pharynx imbibes external yolk cells and a gastric cavity is formed in the center of the syncytium. The syncytial yolk and the blastomeres contained within it are compressed into a thin peripheral rind. From a location close to the embryonic pharynx, which defines the posterior pole, bilaterally symmetric ventral nerve cord pioneers extend forward. Stage 4 is characterized by massive proliferation of embryonic cells. Large yolk-filled cells lining the syncytium form the gastrodermis. During stage 5 the external syncytial yolk mantle is resorbed and the embryonic cells contained within differentiate into an irregular scaffold of muscle and nerve cells. Epidermal cells differentiate and replace the transient embryonic epidermis. Through stages 6–8, the embryo adopts its worm-like shape, and loosely scattered populations of differentiating cells consolidate into structurally defined organs. Our analysis reveals a picture of S. polychroa embryogenesis that resembles the morphogenetic events underlying regeneration.Edited by D. Tautz     

Extremely miniaturised and highly complex: The thoracic morphology of the first instar larva of Mengenilla chobauti (Insecta,Strepsiptera)

Thoracic structures of the extremely small first instar larva of the strepsipteran species Mengenilla chobauti (ca. 200 μm) were examined, described and reconstructed 3-dimensionally. The focus is on the skeletomuscular system. The characters were compared to conditions found in other insect larvae of very small (Ptiliidae) or large (Dytiscus) size (both Coleoptera) and features of “triungulin” larvae, first instar larvae of Rhipiphoridae, Meloidae (both Coleoptera), and Mantispidae (Neuroptera).The specific lifestyle and the extreme degree of miniaturisation result in numerous thoracic modifications. Many sclerites of the exo- and endoskeleton are reduced. Cervical sclerites, pleural ridges, furcae and spinae are absent. Most of the longitudinal muscles are connected within the thorax, and a pair of ventral longitudinal muscles is present in the pleural region of the meso- and metathorax. This results in a high intersegmental flexibility. Due to the size reduction and the correlated shift of the brain to the thorax, with 94 identified muscles the thoracic musculature appears highly compact. Compared to larger larvae the number of both the individual muscles and the muscle bundles are distinctly reduced. The thorax of the first instar larvae displays many additional strepsipteran autapomorphies. At least partly due to the highly specialised condition, potential synapomorphies with other groups were not found.     

Ovule and gametophyte development in Nicotiana glauca Graham (Solanaceae)

Abstract

The development of ovule and megagametophyte is examined in Nicotiana glauca, using light microscopy. The ovules proved unitegmic, tenuinucellate and endothelial as in all the Solanaceae so far studied. The ovule primordia are of the three-zonate type. The integument, which is of dermal origin, is at first two-layered but later produces additional intermediate cells whose origin is not constant. The nucellus, whose initial curvature bears no relation to the origin of the integument, has, like other Solanaceae, a one or two-celled archesporium from which a single meiocyte develops. The gametophyte is confirmed to be bisporic in origin and its development follows the Allium type. Furthermore, the hypostase, which is rare in the family, is observed below the antipodal cells.