Arthropod Structure & Development

Published by Elsevier
Online ISSN: 1467-8039
Publications
Article
Brain morphology is an important character in the discussion of arthropod relationships. While a large body of literature is available on the brains of Hexapoda and Malacostraca, the structure of the brain has been rarely studied in representatives of the Entomostraca. This account examines the morphology and development of the nervous system in the brine shrimp Artemia salina Linnaeus, 1758 (Crustacea, Branchiopoda, Anostraca) by classical histology and immunohistochemistry against synaptic proteins (synapsins), and the neurotransmitters serotonin and histamine. The results indicate that the shape of the developing larval brain in A. salina (a circumstomodeal ring of neuropil) closely resembles that in malacostracan embryos. Furthermore, the organization of the central complex as well as the tritocerebral innervation pattern of the labrum is homologous in this species and in Malacostraca. Nevertheless, differences exist in the layout of the deutocerebrum, specifically in the absence of olfactory glomeruli in A. salina while the glomerular organization of the olfactory lobe is a character in the ground pattern of Malacostraca. These findings are compared to the brain structure in other Euarthropoda and possible phylogenetic implications are discussed.
 
Article
This study represents the first sperm description of a Vespinae species (Vespa crabro). The acrosome consists of an acrosomal vesicle and a perforatorium. The nucleus has compact chromatin and shows lenticular structures on the nuclear envelope. These structures, which have never been observed in a hymenopteran sperm, could be clusters of nuclear pores. The centriolar adjunct has an asymmetric pattern and shows a structured periphery. The centriole consists of 9 accessory microtubules and 9 doublet microtubules devoid of arms and spokes. The axoneme has a 9+9+2 microtubule pattern and the accessory microtubules have 16 protofilaments. The mitochondrial derivatives differ in length and diameter. The larger one is adjacent to the nuclear base, while the smaller one begins below the centriolar adjunct. They possess three distinct areas and a large paracrystalline region, which occurs only in the large one. The large mitochondrial derivative ends first, followed by the small one. The axoneme gradually disorganizes: first the central microtubules disappear, then the doublets, which show opened B-tubules, and finally the accessory microtubules. The sperm morphology of V. crabro is very similar to that of the polistine wasp, Agelaia vicina. This can indicate that, in Vespidae, sperm morphology is maintained without important variations among subfamilies and/or that this similarity indicates close phylogenetic relationship between these two subfamilies. Although Vespidae phylogenetically related to Formicidae, these data suggest that the former more closely related to Apoidea than to Formicidae.
 
Article
Female Ixodes ricinus (ticks) are able to consume several times as much as their own weight in food. The body volume increases without the animal moulting. Special structural and material characteristics of the cuticular exoskeleton render this phenomenon possible. Only the alloscutum, which has a very pronounced lamellar structure, extends. The individual lamellae correspond to surface folds in the outer endocuticle and the epicuticle covering it. The degree of folding is correlated to the nutritional state of the animal, which can be differentiated into three significant phases. The phase-dependent morphology of the surface structure and the cuticular layers of the alloscutum are described qualitatively and quantitatively in this paper. Furthermore, the presence of systematic deposits of resilin in the alloscutum, which are significant for the flexibility, is demonstrated.
 
Article
We report our investigations on the embryonic development of Gryllus assimilis, with particular attention to the head. Significant findings revealed with scanning electron microscopy (SEM) images include: (1) the pre-antennal lobes represent the anterior-most segment that does not bear any appendages; (2) each of the lobes consists of central and marginal regions; (3) the central region thereof develops into the protocerebrum and the optic lobes, whereas the marginal region thereof becomes the anterior portion of the head capsule; (4) the initial position of the antennal segment is posterior to the mouth region; (5) appendage anlagen are transitorily present in the intercalary segment, and they later vanish together with the segment itself; (6) a bulged sternum appears to develop from the ventral surface of the mandibular, maxillary and labial segments. Embryonic features are then compared across the Insecta and further extended to the embryos of a spider (Araneae, Chelicerata). Striking similarities shared by the anterior-most region of the insect and spider embryos lead the authors to conclude that such comparison should be further undertaken to cover the entire Euarthropoda. This will help us to understand the embryology and evolution of the arthropod head.
 
Article
The antennal lobes of different castes of the ant species Camponotus compressus show a marked diversity in the organization of their olfactory glomeruli. Notably, there is a significant difference in the number and size of glomeruli between the reproductives and the workers and among the different worker castes. In this report, we investigate the notion that these caste-specific differences in glomerular number might be accounted for, at least in part, by the differences in numbers of olfactory sensilla that target the antennal lobe. For this, we examine the number of sensilla on the antennal flagella of all the individual castes of C. compressus. This analysis reveals a striking correlation between sensillar number and the number of antennal glomeruli in a given caste. As a first step in investigating the causal mechanisms that might give raise to this correlation, we carry out an initial characterization of olfactory system development in the minor workers of C. compressus. We analyze the temporal pattern of innervations of the developing antennal lobe by olfactory sensory neuron axons. We document the development of the olfactory glomeruli in the antennal lobe during this process, which occurs during early pupal stages. Our findings provide the basis for future manipulative developmental studies on the role of sensory afferent number in glomerular development of different castes within the same species.
 
Article
We use monoclonal antibodies against synaptic proteins and anterograde tracing with neurobiotin to describe the architecture of the antennal lobes in different castes of two ant species -Camponotus sericeus and Camponotus compressus. The reproductives and worker classes are readily categorized based on size and external morphology. The overall organization of brain neuropile is comparable between castes with differences only in the visual ganglia. Males have a larger fraction of neuropile occupied by the medulla and lobula than females. In the diurnal species, C. sericeus these regions are more highly represented, than in the nocturnal species C. compressus. The most striking differences are in the antennal lobe where males possess a macroglomerulus, which is about ten times larger in volume than the other glomeruli; such a specialization is absent in females. Minor workers possess a significantly larger number of glomeruli than the majors despite the smaller overall volume of the lobe. These caste-specific differences occur mainly within glomerular clusters that receive input from sensory neurons that project in tracts - T4 and T5 - within the antennal nerve. The comparative anatomy of different castes of ants provides an entry point into a future systematic analysis of how divergent brain architectures can arise within a single species.
 
Article
Detailed studies of larval development of Octolasmis angulata and O. cor are pivotal in understanding the larval morphological evolution as well as enhancing the functional ecology. Six planktotrophic naupliar stages and one non-feeding cyprid stage are documented in details for the first time for the two species of Octolasmis. Morphologically, the larvae of O. angulata and O. cor are similar in body size, setation patterns on the naupliar appendages, labrum, dorsal setae-pores, frontal horns, cyprid carapace, fronto-lateral gland pores, and lattice organs. Numbers of peculiarities were observed on the gnathobases of the antennae and mandible throughout the naupliar life-cycle. The setation pattern on the naupliar appendages are classified based on the segmentation on the naupliar appendages. The nauplius VI of both species undergoes a conspicuous change before metamorphosis into cyprid stage. The cyprid structures begin to form and modify beneath the naupliar body towards the end of stage VI. This study emphasises the importance of the pedunculate barnacle larval developmental studies not only to comprehend the larval morphological evolution but also to fill in the gaps in understanding the modification of the naupliar structures to adapt into the cyprid life-style. Copyright © 2015. Published by Elsevier Ltd.
 
Article
Johann H.L. Flögel (1834-1918) was an amateur scientist and self-taught microscopist in Germany who 130years ago pioneered comparative arthropod neuroanatomy. He was fascinated by innovations in optical instrumentation, and his meticulous studies of the insect supraoesophageal ganglia were the first to use serial sections and photomicrographs to characterize the architecture of circumscribed regions of brain tissue. Flögel recognized the interpretative power resulting from observations across various species, and his comparative study of 1878, in particular, provided a baseline for subsequent workers to evolve a secure nomenclature of insect brain structures. His contributions stand out from contemporary accounts by virtue of their disciplined descriptions and emphasis on identifying comparable elements in different taxa. Here we give a biographical sketch of his life and summarize his remarkable achievements.
 
Article
The midgut of unfed larvae and adult mites of Platytrombidium fasciatum (C.L. Koch, 1836) and Camerotrombidium pexatum (C.L. Koch, 1937) (Acariformes: Microtrombidiidae) was investigated by electron microscopy. The sac-like midgut occupies the entire body volume, ends blindly and is not divided into functionally differentiated diverticula or caeca. The midgut walls are composed of one type of digestive cell that greatly varies in shape and size. In larvae, the lumen of the midgut is poorly recognizable and its epithelium is loosely organized, although yolk granules are already utilized. In adults, the midgut forms compartments as a result of deep folds of the midgut walls, and the lumen is well distinguished. The epithelium is composed of flat, prismatic or club-like cells, which may contain nutritional vacuoles and residual bodies in various proportions that depend on digestive stages. In both larvae and adult mites, parts of cells may detach from the epithelium and float within the lumen. The cells contain a system of tubules and vesicles of a trans-Golgi network, whereas the apical surface forms microvilli as well as pinocytotic pits and vesicles. Lysosome-like bodies, lipid inclusions and some amount of glycogen particles are also present in the digestive cells. Spherites (concretions) are not found to be a constant component of the digestive cells and in adult mites occur for the most parts in the midgut lumen.
 
Article
Organization of dermal glands in adult water mites Teutonia cometes (Koch, 1837) was studied using light-optical, SEM and TEM methods for the first time. These glands are large and occur in a total number of ten pairs at the dorsal, ventral and lateral sides of the body. The slit-like external openings of the glands (glandularia) are provided with a cone-shaped sclerite, and are combined with a single small trichoid seta (hair sensillum), which is always situated slightly apart from the anterior aspect of the gland opening. Each gland is formed by an epithelium encompassing a very large lumen (central cavity) normally filled with secretion that stains in varying intensity on toluidine blue stained sections. The epithelium is composed of irregularly shaped secretory cells with an electron-dense cytoplasm and infolded basal portions. The cells possess a large irregularly shaped nucleus and are filled with tightly packed slightly dilated cisterns and vesicles of rough endoplasmic reticulum (RER) with electron lucent contents. Dense vesicles are also present in the apical cell zone. Some cells undergo dissolution, occupy upper position within the epithelium and have a lighter cytoplasm with disorganized RER. Muscle fibers are regularly present in the deep folds of the basal cell portions and may serve to squeeze the gland and eject the secretion into the external milieu. The structure of these dermal glands is compared with the previously described idiosomal glands of the same species and a tentative correlation with the glandularia system of water mites is given. Possible functions of the dermal glands of T. cometes are discussed.
 
Article
The mandibles of decapod zoea-I larvae are robustly built masticating mouthparts equipped with several processes and spines. Superficial examination of these sturdy, inflexible structures can suggest that they are lacking sensory receptors. However, detailed TEM analysis of their ultrastructure revealed up to 11 sensillar cell clusters on the gnathal edges of the mandibles of the zoea-I in Palaemon elegans Rathke, 1837. Based on ultrastructural criteria we distinguish 7 types of sensilla: mechanoreceptors, chemoreceptors and mechano- and chemoreceptors. One sensory unit located at the base of the 'lacinia mobilis' exhibits the typical features of a crustacean mechanosensitive sensillum with an external seta and corresponding ultrastructure. Another unit shows features indicating bimodal contact chemosensitivity. A third one is similar to known olfactory chemoreceptors. Using the concept of modality-specific structures we analyse the structure and functional morphology of each sensillum, and give a comprehensive overview of the sensory abilities of zoea mandibles. We take a closer look at the ultrastructure of the 'lacinia mobilis', providing further features to trace its evolutionary history in Decapoda, and thus contributing to a better understanding of malacostracan phylogeny.
 
Article
Detailed information on sensory organs of Diplopoda especially on antennal sensilla are still sparse and fragmentary. The present study on the antennae of Oranmorpha guerinii (Polydesmida, Paradoxosomatidae) utilizing scanning electron microscopy revealed the presence of six sensillar types: (1) apical cones, (2) sensilla trichodea, (3) sensilla microtrichodea, (4) sensilla chaetica, (5) sensilla basiconica bacilliformis, and (6) sensilla basiconica spiniformis. External structure and distribution of cuticular antennal sensilla are compared with data from other diplopod species. We moreover discuss possible functions of antennal sensilla in millipedes.
 
Article
We document the early morphogenesis of Latrodectus geometricus, particularly of the anterior body region. Significant changes in the development of the external prosomal structures revealed with scanning electron microscopy (SEM) images include: (1) reorganisation of each pre-cheliceral lobe by subdivision and internalisation of its central area; (2) shortening of the ventro-median bridge connecting the pre-cheliceral lobes and its eventual disappearance; (3) appearance and expansion of a prospective mouth region between the pre-cheliceral lobes with a recessed median area surrounded by lip-like borders, the anterior lip-part developing into the hypostome; (4) reduction of the mouth region to an area around the hypostome and the lip-like latero-posterior border of the mouth opening; (5) change of the position of the mouth region from anterior to the insertions of the chelicerae to posterior to them; (6) eventual shortening of the mouth opening to a slit overhung by the hypostome; (7) origination of the prosomal shield from the anterior margin of the pre-cheliceral lobes and the tergal portions of the four posterior-most prosomal segments; and (8) expansion of a 'ventral sulcus' from the cheliceral to the fifth opisthosomal segment separating the sides of these segments. Embryonic features are compared across the Chelicerata and discussed briefly in a phylogenetic context.
 
Article
Inhibitory motoneurons which supply the leg musculature are identified and characterized in the scorpion, Vaejovis spinigerus (Wood, 1863) (Vaejovidae, Scorpiones, Arachnida). (1) Successive intracellular muscle fiber recordings from antagonists, and correlation of the monitored inhibitory postsynaptic potentials with spikes in motor nerves, suggest supply of the scorpion leg musculature by common inhibitory motoneurons. (2) Anti-GABA immunohistochemistry is combined with transmission electron microscopy to estimate the number of inhibitory motor axons present in the main leg nerve. The number of immunoreactive axons decreases toward more distal leg segments, from 14 to 18 in the basis to 6-8 in the tibia. No immunoreactive axons are detected beyond the tibia. (3) The distribution of putative inhibitory neurons in the subesophageal ganglion mass is determined by anti-GABA immunohistochemistry, revealing notable similarities to the situation in pterygote insects. This provides a framework for the characterization of the inhibitory motoneurons. (4) Backfills from leg nerves are combined with anti-GABA immunocytochemistry to identify inhibitory motoneurons in the central nervous system. Putative inhibitory motoneurons occur in three clusters per hemi-segment. Two clusters are located near the posterior edge of the neuromere, one lateral, the other more medial, and both contain ca. 8-10 cell bodies. The third cluster consists of two somata located contralaterally, just off the ganglion midline.
 
Article
(1) The musculature of the walking legs is analysed with regard to both morphology and function in the scorpion, Vaejovis spinigerus (Wood, 1863) (Vaejovidae, Scorpiones, Arachnida), and selected other species. Conspicuous features are multipartite muscles, muscles spanning two joints, and partial lack of antagonistic muscles. The muscle arrangement is compared to that in the walking limbs of other Arthropoda and possible phylogenetic implications are discussed. (2). Histochemical characterisation of selected leg muscles indicates that these are composed of layers of slow, intermediate and fast muscle fibres. Anti-GABA immunohistochemistry shows that mainly the intermediate fibres receive innervation from putative inhibitory motoneurons. (3). Intracellular recording from muscle fibres reveals both excitatory and inhibitory muscle innervation. Individual muscle fibres may receive input from more than one inhibitory motoneuron, as indicated by different IPSP amplitudes. (4). The motoneuron supply of the leg muscles is analysed by retrograde fills of motor nerves. The general arrangement of leg motoneurons in the central nervous system and motoneuron anatomy conforms to the situation in pterygote insects and decapod crustaceans. For example, there are an anterior and a posterior group of leg motoneurons in each hemineuromere, and two contralateral somata near the ganglion midline. Between 12 and 20 motoneurons are found to supply each muscle. Most motoneuron cell bodies supplying a given muscle are arranged in a single cluster with a specific location.
 
Article
The crustacean family Gnathiidae Leach, 1814 (Peracarida: Isopoda) comprises 12 genera known mostly from marine environments. Juvenile gnathiid isopods are fish ectoparasites, feeding on blood and tissue fluids in order to complete their life cycles. Gnathiid juvenile development generally includes three moults, the last involving metamorphosis to non-feeding, adult stages. The blood meal ingested by juveniles provides resources for adult survival, reproduction and embryological development. Reproductive biology in the brackish water gnathiid, Paragnathia formica (Hesse, 1864), is unusual amongst crustaceans, since brooding females have paired internal uterine sacs, rather than an external brood pouch. Known embryological development for P. formica includes three post gastrulation stages. In the current study, brooding and embryological development in this gnathiid were reexamined using histological and fluorescence methods, and by scanning electron microscopy. Novel observations were made of the blastodisc and germ cell migration within developing eggs, release of Stage 2 embryos by rupture of embryonic membranes, the in utero moult of Stage 2 to Stage 3 embryos, and the asynchronous development of the brood within the paired uterine sacs. These findings highlight the remarkable nature of brooding in P. formica and expand the paucity of knowledge of embryological development in gnathiids in general.
 
Article
Transmission electron microscopy was conducted on the digestive epithelium of the crustacean ectoparasite Dolops ranarum to elucidate its ultrastructure for the first time, both in a nourished and starved condition. Specimens were collected from the Limpopo Drainage System in South Africa, and the specimens were killed and dissected in Todd's fixative. The anterior midgut is composed mostly of absorptive cells or R cells, while the diverticula are composed of R cells and of F cells, which are moderately abundant in rough endoplasmic reticulum. They are probably responsible for producing digestive enzymes. The posterior midgut is composed of papilliform B cells with large apically located vesicles and R' cells devoid of cell inclusions. Under starvation, specimens survive for a maximum of 12 days; R cells show the most conspicuous changes in ultrastructural characteristics. It is concluded that D. ranarum has adapted to short-term survival only without a host.
 
Article
The ultrastructure of the cells of the digestive system of Argulus japonicus is described with the use of transmission electron microscopy. Specimens of Argulus japonicus were collected from the Vaal Dam in South Africa and fixed in Todd's fixative. The samples were post fixed in osmium tetroxide and embedded in resin. The anterior midgut is composed mostly of R cells while the enteral diverticula are composed mainly of R cells in the proximal diverticules and of F cells in the distal diverticula. The posterior midgut is composed of very large papilliform B cells and of R cells. The R cells in the anterior midgut probably absorb nutrients including lipids. The F cells are filled mostly with rough endoplasmic reticulum, suggesting enzyme synthesis, while the B cells portrayed endocytotic vesicles, indicating intracellular digestion of predigested food. The R cells of the posterior midgut are less active than cells present in the anterior midgut. E cells and peritrophic membrane were not observed.
 
Article
The Malpighian tubules of workers of the fire ant Solenopsis saevissima (Myrmicinae) were analyzed by scanning and transmission electron microscopy in order to determine their functional organization and association with the hindgut epithelium. The ants showed six Malpighian tubules with three segments morphologically and structurally different. The proximal segment was long and its cells showed abundant smooth endoplasmic reticulum and lipid droplets, which suggest their role in lipid secretion. The mid segment was long and undulated and it was composed by the cells that showed the typical features of ion transporting epithelia. The distal segment, short and flattened, adheres to the rectum wall. The cells of this segment showed the basal lamina fused to that of the rectum, it is probable that this part of the tubule may play a role in ion and water uptake from the feces.
 
Article
The embryogenesis of a collembolan, Tomocerus cuspidatus, was examined and described, with special reference to the development of serosa and its developmental potential. As a result of cleavage, which starts with holoblastic cleavage and changes to the superficial type, the blastoderm forms. At the center of the dorsal side of the egg, the primary dorsal organ develops. The mesoderm is segregated beneath the entire blastoderm, excluding the primary dorsal organ. The mesoderm then migrates to the presumptive embryonic area, and the embryonic and extra-embryonic areas differentiate. The area lined with mesoderm is the embryo, and that devoid of it is the serosa. Owing to blastokinesis completion, the extra-embryonic area or the serosa is highly stretched, and the serosal cells are often found to undergo mitosis. The serosa possesses the ability to differentiate into the body wall. It was confirmed, in contrast to the previous understanding, that the serosal cells do not degenerate, but participate in the formation of the body wall or definitive dorsal closure. Integrating this newly obtained information and other embryological evidence, the basal splitting of Hexapoda was phylogenetically discussed and reconstructed, and a phylogeny formulated as "Ellipura (= Protura + Collembola) + Cercophora (= Diplura and Ectognatha)" was proposed. Copyright © 2015. Published by Elsevier Ltd.
 
Tetraphalerus bruchi, male.
Tetraphalerus bruchi , male abdomen, habitus. Dorsal (left) and ventral (right) 
T. bruchi, male postabdomen, hyperextended, SEM images (left) and corresponding line drawings (right). (A, B) Dorsal view; (C, D) lateral view; (E, F) ventral view. bp, Basal piece; de, ejaculatory duct; par, paramere; pe, penis; s, segment; st, sternite; and tg, tergite.
Article
External and internal features of the male postabdomen of Tetraphalerus bruchi were examined with a broad spectrum of morphological techniques and are described in detail. The conditions found in males of Tetraphalerus are compared to those in other archostematan beetles and members of other coleopteran suborders. The far-reaching reduction of the sternite I, structural modifications of sternite II, the retracted condition of the terminal segments, and ventromedially fused apodemes arising from the anterior margin of tergite IX are likely autapomorphies of Coleoptera. The male postabdomen of Tetraphalerus is less derived than in most other groups of Coleoptera. The sclerotized elements are symmetrical. In contrast to earlier statements on the archostematan male genital apparatus a distinctly developed, sclerotized basal piece is present. The aedeagus is trilobed and all elements of the copulatory apparatus are distinct. The muscular equipment is simple and moderately developed. All muscles (except the transverse muscles 61 and 62) occur pairwise and symmetrically. The distinct increase of the number of postabdominal muscles in representatives of the higher lineages of Coleoptera is likely linked with a torsion of the copulatory apparatus, which also results in asymmetries of the sclerotised parts. The testes of Tetraphalerus are long, multi-coiled tubes like in other archostematans, Myxophaga (Torridincola) and Adephaga. The presence of a deep notch on the parameres is a synapomorphy of Tetraphalerus and Omma. Curved parameres, a shortened distal portion, and a distinctly shortened penis are potential synapomorphies of Omma rutherfordi and Omma mastersi. The large size of the sclerotized part of the phallobase ('basal piece') and the division of the sclerotization of sternum IX are potential ground-plan autapomorphies of Archostemata, with secondary modification of the latter feature in Cupedidae. The reduced condition of the sclerotization of sternum VIII is an apomorphic condition which has likely evolved independently in Tetraphalerus and Paracupes. Further anatomical investigation of the male genital apparatus of Coleoptera and holometabolous insects in general is required for a reliable morphological and phylogenetic interpretation. Concerning the presence or absence of particular sclerotizations (e.g., 'basal piece' of phallobase) histological section series and Confocal Laser Scanning Microscopy can add more precise information to what can be observed using permanent preparations of macerated specimens.
 
Article
Aspects of pantopod ontogeny have been known for a long time, but specific information is available for only a few species. Our account of the postembryonic development of Pycnogonum litorale is based on laboratory-reared individuals and SEM studies. We documented particularly all early developmental stages, with emphasis on morphogenetic changes of head structures and appendages. In P. litorale the protonymphal limbs, the chelicerae and two more uniramous legs, degenerate already during the larval phase; only the third one, the ovigers, reappears in male juveniles. Other Pantopoda vary in this aspect from retention of all three protonymphal appendages to their complete reduction, as in P. litorale. Accordingly, the two post-cheliceral larval appendages are separate legs in front of the walking legs in the adults, the 'parapalps' and the 'ovigers', but they do not occur in all pantopods. The scarcity of studies of the ontogeny of Pantopoda prevents us from a more conclusive picture, but our data are promising to state that additional such studies will increase the usability of ontogenetic data for a phylogenetic analysis of Pantopoda, the crown group of the Pycnogonida. We also discuss the phylogenetic implications of our data in the light of new information from Hox genes and developmental-biological data on body segmentation and tagmosis of the Chelicerata. These suggest the homology of chelicerae and antenn(ul)ae of other euarthropods. Accepting this, we conclude that the adult pycnogonid/pantopod head, the cephalosoma, corresponds to the euarthropod head and that the protonymph with three appendage-bearing segments may represent an even shorter, possibly phylogenetically older larval type than the euarthropod 'head larva' bearing four pairs of appendages. In further consequence, the fourth walking legs of Pycnogonida/Pantopoda should correspond to the first opisthosomal appendages, the chilaria, of euchelicerates. This implies that within Pycnogonida the post-prosomal region became compacted during evolution to a single leg-bearing segment plus a tubular end piece. Accordingly, neither the anterior nor the posterior functional boundaries of the walking-leg region correspond to the original tagma borders.
 
Article
The sperm of Microstigmus arlei and Microstigmus nigrophthalmus are twisted in a spiral and consist of two regions: the head, formed by an acrosome and a nucleus, and the flagellum, formed by two asymmetric mitochondrial derivatives, a long centriolar adjunct, an axoneme (9+9+2) and two accessory bodies. The head shows a characteristic morphology. The acrosome is very long and is basically made up of a paracrystalline structure. In the central head region, the acrosome is inserted into the nucleus, which is observed coiling laterally around the paracrystalline structure. In the subsequent part of the spermatozoon the nucleus appears round in transverse sections, and over some length it is still penetrated by the acrosome until shortly distal to the flagellar insertion. At this point the nucleus forms an inverted cone-shaped projection. These morphological characteristics of acrosome and nucleus of the Microstigmus wasp have not been previously described in Apoidea and are useful for phylogenetic evaluation of this superfamily.
 
Article
Ultrastructural details of spermiogenesis, spermatozoa and the spermatophore of the early derived actinedid mite Saxidromus delamarei are described. Spermatids and mature sperm cells are provided with up to four acrosomal complexes and nuclei derivatives (chromatin bodies). Due to this reason, the sperm cells may be classified as synspermia, a sperm type found only in some spiders until now. The acrosomal complex is composed of a remarkably complicated vacuole and filament. Other peculiarities of sperm structure correspond to those found in prostigmatic mites, i.e. penetration of the chromatin body by the acrosomal filament and the presence of peripheral invaginations of the plasmalemma. The sperm cells are covered by a thin secretion layer of probably proteinaceous material. Stalked spermatophores are rather large, but simply structured and contain relatively few sperm cells. The results are discussed taking systematical and behavioural aspects into account. In particular, it is suggested that the peculiar mating behaviour of these mites secures both sperm transfer and first male's sperm priority and that this allowed reduction of sperm numbers.
 
Article
We present here a description of early development in the onychophoran Euperipatoides kanangrensis with emphasis on processes that are ambiguously described in older literature. Special focus has been on the pattern of early cleavage, blastoderm and germinal disc development and gastrulation. The formation of the blastopore, stomodeum and proctodeum is described from sectioned material using light and transmission electron microscopy as well as whole-mount material stained for nuclei and gene expression. The early cleavages were found to be superficial, contrary to earlier descriptions of cleavage in yolky, ovoviviparous onychophorans. Also, contrary to earlier descriptions, the embryonic anterior-posterior axis is not predetermined in the egg. Our data support the view of a blastopore that becomes elongated and slit-like, resembling some of the earliest descriptions. From gene expression data, we concluded that the position of the proctodeum is the most posterior pit in the developing embryo. This description of early development adds to our knowledge of the staging of embryonic development in onychophorans necessary for studies on the role of developmental changes in evolution.
 
Semischematic models of ommatidia: (a) ark clam; (b) sabellid polychaete; (c) xiphosurid; (d) myriapod; (e) scutigeromorph; (f) non-malacostracan crustacean; (g) malacostracan crustacean; (h) insect.
Receptive fields of individual receptor cells in the ommatidium, illustrated by lines through the posterior nodal point. In the myriapod type of ommatidium (a), distal cells will have wide off-axis sensitivities, whereas proximal receptors will have narrow angular sensitivities close to the ommatidial axis. In the crustacean type of ommatidium (b), light guiding provides all receptors with identical receptive fields.
Evolution of arthropod ommatidia, suggested on the basis of functional arguments.
Article
New data on the phylogenetic relationships of various arthropod groups have spurred interesting attempts to reconstruct the evolution of arthropod nervous and visual systems. Some of the relevant new data are cell identities and developmental processes in the nervous and sensory systems, which is particularly useful for reconstructing the evolution of these systems. Here, we focus on the structure of compound eye ommatidia, and make an evolutionary analysis with functional arguments. We investigate possible routes of evolution that can be understood in terms of selection for improved visual function, and arrive at a number of conclusions that are discussed in the light of recent phylogenetic hypotheses. On the basis of ommatidial focusing structures and the arrangement of receptor cells we show that the evolution of compound eyes proceeded largely independently along at least two lineages from very primitive ancestors. A common ancestor of insects and crustaceans is likely to have had ommatidia with focusing crystalline cones, and colour and/or polarization vision. In contrast, the compound eyes in myriapods and chelicerates are likely to date back to ancestors with corneal lenses and probably without the ability to discriminate colour and polarization.
 
Article
Arthropod photoreceptor evolution is a prime example of how evolution has used existing components in the origin of new structures. Here, we outline a comparative approach to understanding the mutational origins of novel structures, describing multiple examples from arthropod photoreceptor evolution. We suggest that developmental mechanisms have often split photoreceptors during evolution (field-splitting) and we introduce "co-duplication" as a null model for the mutational origins of photoreceptor components. Under co-duplication, gene duplication events coincide with the origin of a higher level structure like an eye. If co-duplication is rejected for a component, that component probably came to be used in a new photoreceptor through regulatory mutations. If not rejected, a gene duplication mutation may have allowed the component to be used in a new structure. In multiple case studies in arthropod photoreceptor evolution, we consistently reject the null hypothesis of co-duplication of genetic components and photoreceptors. Nevertheless, gene duplication events have in some cases occurred later, allowing divergence of photoreceptors. These studies provide a new perspective on the evolution of arthropod photoreceptors and provide a comparative approach that generalizes to the study of any evolutionary novelty.
 
Article
Identifying the unambiguous sister group to the hexapods has been elusive. Traditional concepts include the Myriapoda (the Tracheata/Atelocerata hypothesis), but recent molecular studies consistently indicate it is the Crustacea, either in part or entirety (the Pancrustacea/Tetraconata hypothesis). The morphological evidence in support of Tracheata is reviewed, and most features are found to be ambiguous (i.e., losses, poorly known and surveyed structures, and probable convergences), though some appear to be synapomorphic, such as tentorial structure and the presence of styli and eversible vesicles. Other morphological features, particularly the structure of the eyes and nervous system, support Pancrustacea, as does consistent molecular evidence (which is reviewed and critiqued). Suggestions are made regarding hexapod-crustacean limb homologies. Relationships among basal (apterygote) hexapods are reviewed, and critical Paleozoic fossils are discussed. Despite the scarceness of Devonian hexapods, major lineages like Collembola and even dicondylic Insecta appeared in the Early Devonian; stem-group and putative Archaeognatha are known from the Carboniferous through Permian and the Late Devonian, respectively. Thus, the earliest divergences of hexapods were perhaps Late Silurian, which is considerably younger than several estimates made using molecular data.
 
Article
Most individuals of the Australian 'fire-beetle' Merimna atrata have two pairs of IR receptors which are located ventrolaterally on the second and third abdominal sternite. An IR receptor consists of a specialized IR absorbing area, which is innervated by a neural complex. This complex contains one thermoreceptive multipolar neuron with a unique terminal dendritic mass (TDM) and two scolopidia and was termed 'sensory complex'. However, also individuals with one pair of IR receptors on the second sternite and beetles with three pairs on the second, third, and fourth sternites were found. Additionally, beetles having one or two pairs of IR receptors may have preliminary stages of IR receptors on the third and fourth sternite, respectively. We found two kinds of preliminary stages, both of which are characterized by a much less pronounced absorbing area. In all five abdominal sternites segmental nerves are attached to the cuticle with a neural complex. Investigation of complexes of non-IR sternites suggests that the sensory cells inside the sensory complex of an IR receptor have developed from common internal stretch receptors. From our results it can be hypothesized that the IR sensory system in Merimna atrata has not yet reached a stage, which can be regarded as evolutionary stable.
 
SEM of L. cruzi and L. longipalpis. Fig. 3. L. cruzi. Detail of tergite IV, showing that the macrotrichia bases are located at the tergite periphery (asterisks), around the region where the disseminating structures are present. Fig. 4. L. cruzi. Detail of a pheromone disseminating structure on tergite IV. Fig. 5. L. longipalpis. The pheromone disseminating structures (arrowhead) are situated amidst the microtrichia (arrow) and can be easily distinguished from the macrotrichia bases (B). Fig. 6. L. longipalpis pupa. The arrows indicate the spot localization on abdominal tergite IV.
Article
The fine structure of male sex pheromone disseminating structures on abdominal segments of five species of Lutzomyia sandflies (Diptera: Psychodidae) was analyzed. Scanning electron microscopy showed that in Lutzomyia cruzi [Mem. Inst. Oswaldo Cruz 33 (1938) 349] and Lutzomyia longipalpis [Mem. Inst. Oswaldo Cruz 4 (1912) 84], the disseminating structures are located in pale spots on abdominal tergites IV or III and IV and are morphologically similar, appearing as small round cuticular elevations with a central pore. Observation of abdominal tergites of L. longipalpis pupae showed that the spots, but not the structures, are already present in this stage. In Lutzomyia lenti [Mem. Inst. Oswaldo Cruz 33 (1938) 349] and Lutzomyia carmelinoi [Mem. Inst. Oswaldo Cruz 81 (1986) 323] adult males, the disseminating structures are present on tergal segments V and VI, where pale spots could not be observed, and appear as apple-like elevations with a central pore. In Lutzomyia renei, a single disseminating structure is found at the anterior region of tergal segment VI. Transmission electron microscopy was used to analyze the gland cell fine structure in L. cruzi. Several class III gland cells are located side by side in the fourth abdominal segment. Each epidermal secretory cell contains a small reservoir and a short outlet channel through the cuticle.
 
Article
A study was conducted to determine whether the mandibles of the detrito-/bryophagous groundhopper Tetrix tenuicornis are subject to mechanical wear as a result of feeding, as is the case for grasshoppers that feed on silica-rich grasses. Abrasion was evaluated by measuring the length and width of the 3rd incisor and length of the 4th incisor in adults of different ages collected under natural conditions during one season. Although T. tenuicornis and other groundhoppers avoid feeding on grasses, we found that mandible abrasion increased with T. tenuicornis age. Age-related abrasion of the incisors of left and right mandibles was statistically significant in both sexes but the degree of abrasion was greater for females than males, apparently reflecting differences in the frequency and magnitude of feeding. Degree of abrasion also differed between right and left mandibles, probably because of differences in how each mandible is used during food processing. Abrasion of chitinous mandible structures may reduce the effectiveness of food processing late in the season.
 
Article
Pericardial cells are present in a wide variety of insects and are thought to constitute the majority of 'athrocytes (nephrocytes)'. In contrast, peritracheal athrocytes have only been observed in Bombyx mori L. Although peritracheal athrocytes have a distinct morphology, it is unknown whether these cells are common to all lepidopterans. We anatomically compared eight lepidopteran species: Bombyx mori L. and Bombyx mandarina M. (Bombycidae); Samia cynthia ricini D. (Saturniidae); Agrius convolvuli L. (Sphingidae); Spodoptera litura F. and Mythimna separata W. (Noctuidae); Pieris rapae L. (Pieridae); and Glyphodes pyloalis W. (Crambidae). Of these species, only Bombyx mori L. and Bombyx mandarina M. possess peritracheal athrocytes.
 
Article
First generation egg clusters of Elasmucha grisea are more closely guarded than second generation clusters. The ovaries of this species are structured to enhance this behavior. The population of E. grisea from S-W Poland breeds in the spring (May-June) and late summer (July-August). The second generation clutches contain fewer eggs and are destroyed 3-4 days after oviposition by predators and parasitoids. The ovary structure in the studied species differs from that found in other Heteroptera. The average number of ovarioles per ovary is 24 while in the other investigated species the number of ovarioles per ovary is 6-7. Lateral oviducts are elongated and the ovarioles are arranged in a pennate pattern. Each ovariole contains only one growing ovarian follicle. Differentiation of the ovarioles and ovarian follicles is synchronised thus enabling simultaneous oviposition. A comparative analysis of the ovary structure during the life cycle, particularly the presence of atresive ovarian follicles in the ovarioles of egg- and nymph guarding females, as well as the shape and structure of the apical part of the tropharium all support the hypothesis of cooperation between females in E. grisea. A similar ovary structure has been observed in the Coccoidea (Hemiptera, Homoptera) which indicates presocial behavior.
 
Article
The mode of cleavage in the Acari is generalized as superficial or intralecithal, with a preceding phase of total (holoblastic) cleavage, but the knowledge is fragmentary and conclusions have been inconsistent, even when relating to the same species. Since no data about early embryology is available for the speciose group Oribatida, we studied Archegozetes longisetosus using transmission electron microscopy. We focused on early cleavages and the formation of the blastula, as these are the important and controversial points in early embryology of the Acari. We expected, as postulated for other acarine eggs, the early cleavages to be holoblastic and followed by a superficial preblastoderm stage. The early cleavages of A. longisetosus are holoblastic and blastomeres give rise to yolk-free micromeres and macromeres containing all the yolk. In contrast to expectations, the micromeres do not form a superficial preblastoderm layer. They are scattered along the embryonic surface and form an external, monocellular layer that covers the whole surface of the embryo. Since each of the existing TEM studies of mites shows this same pattern, and since this specialized form of total cleavage seems to be unique in Chelicerata, it may be the general mode of cleavage in Acari. However, the question will require much more investigation, especially since most data relate to the Actinotrichida and very few are currently available for species in the other major group, the Anactinotrichida.
 
Article
Mitochondria are well-characterized intracellular organelles usually concentrated in locations of high energy consumption. Light microscopic and transmission electron microscopic observations of the internal anatomy of the feather mites Diplaegidia columbae and Falculifer rostratus were conducted. In the anterior half of the bodies of the mites, we found several dozen of distinctive mitochondria-rich (MR) cells filled with abundant, large mitochondria. Mitochondria are placed individually or enclosed in small groups within an elaborated lamellar system forming a mitochondria-lamellae complex (MLC). The role of the MLC as well as the MR cells is not clear at present, but their involvement in heat generation is hypothesized and briefly discussed.
 
Article
Testes, spermatogenesis and spermatozoa are described in the mite Carpoglyphus lactis (L.), the first representative of the Hemisarcoptoidea superfamily studied ultrastructurally. Paired testes are located posteriorly in the idiosoma, with germaria situated dorsolaterally. The germarium consists of a compact group of spermatogonia; no testicular central cell was found. The remainder of the gonad is occupied by germ cells in different stages of spermatogenesis, distributed separately rather than in cysts, and embedded in a few large somatic cells filling the remaining space. Spermatocytes are covered by a spongy layer, a product of the Golgi apparatus. Spermatids are anucleate. Their chromatin condenses into granular and then tubular threads. As spermiogenesis progresses, the spongy layer assembles at a single site and forms a structure termed the spongy body; mitochondria become electron dense, elongate and gather forming a bundle; a narrow ER cistern, promptly transforming into a dense lamella, appears between the mitochondria and chromatin. Mature spermatozoa are small, highly electron-dense cells interdigitating with others via superficial protrusions. They possess chromatin threads, electron-dense lamella and mitochondria, but do not have an acrosome. Our results support the monophyly of Astigmata, but do not explain the phylogenetic affinities of Hemisarcoptoidea to other superfamilies of astigmatic mites.
 
Article
The coxal glands of M. murismusculi consist of the proximal tubular portion (tubulus), the distal glandular sac and the terminal excretory duct. The tubulus comprises looped proximal and distal tubes that run in close association with each other. The cells of the proximal tube form numerous short protrusions that project into the neighbouring organs through the pores in their basal lamina. The sac is a distal part of the gland and so it cannot be considered as a homologue of the proximal filter sacculus of other arthropods. A large number of pinocytotic vesicles and lysosome-like bodies in the epithelial lining of the sac imply that the main functions of this organ may be the absorption of substances from the lumen of the gland and their subsequent intracellular transformation. In addition the sac of females was shown to produce dense secretory granules. The ultrastructural features of the glands are discussed and compared to other representatives of Acari.
 
Article
The pedicel is a structure that connects the phoretic deutonymph of Uropodina mites with its carrier and enables dispersal. The shapes, lengths and diameters of pedicels formed by Uropoda orbicularis, Trichouropoda ovalis, Uroobovella pulchella and Uroobovella nova were studied by scanning and light microscopy. Pedicels of U. orbicularis and T. ovalis have the shape of a straight stalk. In U. pulchella, the pedicel is extremely short, irregularly shaped and composed of homogenous material. The longest pedicel is found in U. nova and it may be helically coiled in this species. The length of the pedicel is positively correlated with deutonymph body size between species, but not within species. Pedicels of U. orbicularis and U. pulchella have the largest diameter. The pedicel diameter in U. orbicularis and T. ovalis is inversely proportional to its length, but not in U. nova and U. pulchella. The constituent of pedicel stems in U. pulchella is homogenous, whereas in U. orbicularis and T. ovalis it contains a bundle of tightly packed fibres. In U. nova coiled pedicels are comprised of two layered materials of different electron density, one of which is electron lucid and located peripherally. Hypotheses on the origin of the pedicel are proposed.
 
Article
Phoretic deutonymphs of uropodid mites are attached to their carrier via an anal pedicel which is formed by a secretion from the pedicellar gland. Since the ultrastructure of the pedicel and pedicellar gland has never been investigated, we studied these structures in three species, Uropoda orbicularis (Müller), Uroobovella marginata (Koch), and Trichouropoda ovalis (Koch) by light (LM) and electron microscopy (TEM, SEM). In addition, the pedicel in Uroobovella nova (Oudemans) was documented in SEM. The pedicellar gland is a distinct globular structure comprised of three types of secretory cells (A-, B-, and C-types) with apical parts directed towards a junctional zone of postcolon and anal atrium. Secretory cells of the A-type are located dorsally, whereas B-type cells are central, and C-type cells are distributed ventrally or both ventrally and dorsally. Protrusions of visceral muscle cells are distributed on the external surface of the gland. The cuticle-lined anal atrium is large and located between the gland and dorso-anal muscles. The pedicel is composed of a main stalk and two extended extremities: one adhering to the anal region of the deutonymph and a second connected to the cuticle of the carrier. In each case, the anal atria were empty, whereas the pedicellar material was located outside of the mite body.
 
Article
Ptychoidy is a mechanical predator defence in some groups of Oribatida (Acari), where the animals can retract their legs into the idiosoma and encapsulate. This mechanism is enabled by a number of morphological adaptations. We used the non-invasive technique of synchrotron X-ray microtomography to compare muscular elements involved in ptychoidy of two species from the Euphthiracaroidea (Oribotritia banksi and Rhysotritia ardua) which differ in a number of cuticular elements involved in ptychoidy. We hypothesize that a strong functional correlation exists between these cuticular structures and their corresponding musculature. We found a number of distinct differences concerning quantitative and qualitative muscle morphology. For testing the functional impact of different muscle configurations we simulated two conditions in silico (encapsulated / opened) and analysed the spatial relative force vectors of the prodorsum lateral adjustor muscles (pla) which are responsible for retraction and adjustment of the prodorsum during encapsulation. We show that the functional morphology of these muscles strongly differs between the two species and that this can be explained by the structure of corresponding cuticular elements. Furthermore, the dynamics of pla, as measured by the extent of contraction during encapsulation, is more than two times higher in R. ardua than in O. banksi.
 
Article
Males of Parasitina and Dermanyssina (Gamasida=Mesostigmata) have chelicerae modified to function as gonopods. The slit-like spermatotreme in the movable digit of the chela in males of Parasitina was studied in three species: in Pergamasus quisquiliarum and Holoparasitus sp. a rather simple slit is indeed present, whereas in Vulgarogamasus kraepelini the structure is represented by a fine duct traversing the movable digit. The spermatodactyl studied in two phytoseioid species (Phytoseiulus persimilis, Blattisocius dentriticus) of Dermanyssina is a slender process arising from the movable digit and containing a fine duct which is formed by cuticular folds. The spermatodactyl of these species thus differs remarkably from that described in Veigaia sp. The diversity of these structures seen in the few taxa studied up to now is discussed under functional and systematic aspects.
 
Article
The development of male and female gonads in arrhenotokous and thelytokous species of Histiostoma was studied using transmission electron microscopy (TEM). All instars were examined: larvae, protonymphs, facultative heteromorphic deutonymphs (=hypopi), tritonymphs, and adults. In testis primordium, spermatogonia surrounding a testicular central cell (TCC) with a gradually enlarging, branched nucleus are present already at the larval stage. Spermatogonia and the TCC are connected via narrow, tubular intercellular bridges revealing that the TCC is a germline cell. Spermatocytes appear at the protonymphal stage. At the heteromorphic deutonymph stage, the testis primordium is similar to that of the protonymph, but in the tritonymph it is much larger and composed as in the adult: spermatids as well as sperm cells are present. The latter are congregated ventrally in the testis at the entrance of the deferent duct.
 
Article
Astigmatans are a large group of mites living in nearly every environment and exhibiting very diverse reproductive strategies. In spite of a uniform anatomical organization of their reproductive systems, gametogenesis in each sex is highly variable, leading to gamete formation showing many peculiar features and emphasizing the distinct position of Astigmata. This review summarizes the contemporary knowledge on the structure of ovaries and testes in astigmatic mites, the peculiarities of oogenesis and spermatogenesis, as well as provides new data on several species not studied previously. New questions are discussed and approaches for future studies are proposed.
 
Article
Ultrastructure of male reproductive accessory glands and ejaculatory duct in the Queensland fruit fly (Q-fly), Bactrocera tryoni, were investigated and compared with those of other tephritid flies. Male accessory glands were found to comprise one pair of mesodermic glands and three pairs of ectodermic glands. The mesodermic accessory glands consist of muscle-lined, binucleate epithelial cells, which are highly microvillated and extrude electron-dense secretions by means of macroapocrine transport into a central lumen. The ectodermic accessory glands consist of muscle-lined epithelial cells which have wide subcuticular cavities, lined with microvilli. The electron-transparent secretions from these glands are first extruded into the cavities and then forced out through small pores of the cuticle into the gland lumen. Secretions from the two types of accessory glands then flow into the ejaculatory duct, which is highly muscular, with epithelial cells rich in rough endoplasmic reticulum and lined with a thick, deeply invaginated cuticle. While there are some notable differences, reproductive accessory glands of male Q-flies generally resemble those of the olive fruitfly, Bactrocera oleae, and to a lesser extent the Mediterranean fruit fly, Ceratitis capitata.
 
Article
The morphology and the ultrastructure of the male accessory glands and ejaculatory duct of Ceratitis capitata were investigated. There are two types of glands in the reproductive apparatus. The first is a pair of long, mesoderm-derived tubules with binucleate, microvillate secretory cells, which contain smooth endoplasmic reticulum and, in the sexually mature males, enlarged polymorphic mitochondria. The narrow lumen of the gland is filled with dense or sometimes granulated secretion, containing lipids. The second type consists of short ectoderm-derived glands, finger-like or claviform shaped. Despite the different shape of these glands, after a cycle of maturation, their epithelial cells share a large subcuticular cavity filled with electron-transparent secretion. The ejaculatory duct, lined by cuticle, has epithelial cells with a limited involvement in secretory activity. Electrophoretic analysis of accessory gland secretion reveals different protein profiles for long tubular and short glands with bands of 16 and 10kDa in both types of glands. We demonstrate that a large amount of accessory gland secretion is depleted from the glands after 30min of copulation.
 
Article
The development of the female accessory glands in Teleogryllus commodus was studied by detailed morphometric and stereological analyses. In addition, a microsurgical method was developed to quantify the gland secretion. The morphometric results yield evidence that the glands are subject to a significant growth during peak differentiation, starting immediately after the adult moult. The gland growth is exclusively caused by a hypertrophy of single gland cells with respective volume gains between 400 and 700%. According to the stereological results, the volume of mitochondria per cell is marked by an up to fourfold increase during peak differentiation. Other cell structures (rER, sER) are characterized by a similar propagation behavior. The nucleus and nucleolus grow simultaneously with the cell, indicating high production of site-specific macromolecules. Infolds of the basal cell membrane cause a progressive enlargement of the basal cell surface ensuring an increased uptake of secretory precursors from the hemolymph. Quantitative studies show that the total production of secretion increases with proceeding age and can be correlated with a rising egg-laying activity, starting on the eighth day of adult's life. This underlines the main function of the secretion as a lubricant for a facilitated transport of the eggs through the ovipositor.
 
Top-cited authors
Rolf Beutel
  • Friedrich Schiller University Jena
Wolfgang Rössler
  • University of Wuerzburg
Manfred Schmidt
  • Georgia State University
Romano Dallai
  • Università degli Studi di Siena
Harald Krenn
  • University of Vienna