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Background: Centipedes are terrestrial, predatory arthropods with specialized sensory organs. However, many aspects of their sensory biology are still unknown. This also concerns hygroreception, which is especially important for centipedes, as their epicuticle is thin and they lose water rapidly at low humidity. Thus, the detection of humid places is vital but to date no definite hygroreceptor was found in centipedes. House centipedes (Scutigeromorpha) possess a peculiar opening at the base of their antenna, termed 'scape organ', that houses up to 15 cone-shaped sensilla in a cavity. Lacking wall and tip-pores, these socket-less sensilla may be hypothesized to function as hygroreceptors similar to those found in hexapods. Results: The cone-shaped sensilla in the scape organ as well as nearby peg-shaped sensilla are composed of three biciliated receptor cells and three sheath cells. A tip-pore is present but plugged by a highly electron-dense secretion, which also overlays the entire inner surface of the cavity. Several solitary recto-canal epidermal glands produce the secretion. Receptor cell type 1 (two cells in cone-shaped sensilla, one cell in peg-shaped sensilla) possesses two long dendritic outer segments that project to the terminal pore. Receptor cell type 2 (one cell in both sensilla) possesses two shorter dendritic outer segments connected to the first (proximal) sheath cell that establishes a scolopale-like structure, documented for the first time in detail in a myriapod sensillum. Conclusions: The nearly identical configuration of receptor cells 1 with their long dendritic outer segments in both sensilla is similar to hexapod hygroreceptors. In Scutigera coleoptrata, however, the mechanism of stimulus transduction is different. Water vapor may lead to swelling and subsequent elongation of the plug pin that enters the terminal pore, thus causing stimulation of the elongated dendritic outer segments. The interconnection of receptor cell 2 with short outer dendritic segments to a scolopale-like structure potentially suits both sensilla for vibration or strain detection. Thus, both sensilla located at the antennal base of scutigeromorph centipedes fulfill a dual function. [ABSTRACT FROM AUTHOR]

Background: Modern representatives of Polychelida (Polychelidae) are considered to be entirely blind and have largely reduced eyes, possibly as an adaptation to deep-sea environments. Fossil species of Polychelida, however, appear to have well-developed compound eyes preserved as anterior bulges with distinct sculpturation. Methods: We documented the shapes and sizes of eyes and ommatidia based upon exceptionally preserved fossil polychelidans from Binton (Hettangian, United-Kingdom), Osteno (Sinemurian, Italy), Posidonia Shale (Toarcian, Germany), La Voulte-sur-Rhône (Callovian, France), and Solnhofen-type plattenkalks (Kimmeridgian-Tithonian, Germany). For purposes of comparison, sizes of the eyes of several other polychelidans without preserved ommatidia were documented. Sizes of ommatidia and eyes were statistically compared against carapace length, taxonomic group, and outcrop. Results: Nine species possess eyes with square facets; Rosenfeldia oppeli (Woodward, 1866), however, displays hexagonal facets. The sizes of eyes and ommatidia are a function of carapace length. No significant differences were discerned between polychelidans from different outcrops; Eryonidae, however, have significantly smaller eyes than other groups. Discussion: Fossil eyes bearing square facets are similar to the reflective superposition eyes found in many extant decapods. As such, they are the earliest example of superposition eyes. As reflective superposition is considered plesiomorphic for Reptantia, this optic type was probably retained in Polychelida. The two smallest specimens, a Palaeopentacheles roettenbacheri (Münster, 1839) and a Hellerocaris falloti (Van Straelen, 1923), are interpreted as juveniles. Both possess square-shaped facets, a typical post-larval feature. The eye morphology of these small specimens, which are far smaller than many extant eryoneicus larvae, suggests that Jurassic polychelidans did not develop via giant eryoneicus larvae. In contrast, another species we examined, Rosenfeldia oppeli (Woodward, 1866), did not possess square-shaped facets, but rather hexagonal ones, which suggests that this species did not possess reflective superposition eyes. The hexagonal facets may indicate either another type of superposition eye (refractive or parabolic superposition), or an apposition eye. As decapod larvae possess apposition eyes with hexagonal facets, it is most parsimonious to consider eyes of R. oppeli as apposition eyes evolved through paedomorphic heterochrony. Conclusion: Polychelidan probably originally had reflective superposition. R. oppeli, however, probably gained apposition eyes through paedomorphosis. [ABSTRACT FROM AUTHOR]

Superposition and apposition compound eyes are commonly associated with moths and butterflies, respectively. However, recently intermediate eye designs, combining features of both apposition and superposition eyes were found in tiny insects. Here, we examine the eyes of 12 species of moth, ranging from 1.88 to 6.03 mm body size, by scanning and transmission electron microscopy. Correlations between body and eye sizes are discussed with regard to the eyes' functionality. Although all of the species shared an ommatidial organization characteristic of pterygote insects, three optical designs were found: (a) an apposition eye, (b) an eye resembling apposition eyes, but with a unique crystalline cone, and (c) an eye intermediate in structure between apposition and superposition eyes. Our comparisons also revealed a new type of basal matrix for the Lepidoptera. The results show that in most of the examined compound eyes (with the exception of the apposition eye of Micropterix aruncella), a clear distinction between apposition and superposition eyes is not feasible. Due to functional morphological constraints as a consequence of miniaturization, evolutionary transformations from superposition into apposition optics may have occurred several times independently in various 'microlepidopteran' taxa. The Phyllonorycter medicaginella eye appears to illustrate this evolutionary scenario best. [ABSTRACT FROM AUTHOR]

Crustaceans have successfully adapted to a variety of environments including fresh- and saltwater as well as land. Transition from an aquatic to a terrestrial lifestyle required adaptations of the sensory equipment of an animal, particularly in olfaction, where the stimulus itself changes from hydrophilic to mainly hydrophobic, air-borne molecules. Hermit crabs Coenobita spp. (Anomura, Coenobitidae) have adapted to a fully terrestrial lifestyle as adults and have been shown to rely on olfaction in order to detect distant food items. We observed that the specialized olfactory sensilla in Coenobita, named aesthetascs, are immersed in a layer of mucous-like substance. We hypothesized that the mucous is produced by antennal glands and affects functioning of the aesthetascs. Using various microscopic and histochemical techniques we proved that the mucous is produced by aesthetasc-associated epidermal glands, which we consider to be modified rosette-type aesthetasc tegumental glands known from aquatic decapods. These epidermal glands in Coenobita are multicellular exocrine organs of the recto-canal type with tubulo-acinar arrangement of the secretory cells. Two distinct populations of secretory cells were clearly distinguishable with light and electron microscopy. At least part of the secretory cells contains specific enzymes, CUB-serine proteases, which are likely to be secreted on the surface of the aesthetasc pad and take part in antimicrobial defense. Proteomic analysis of the glandular tissue corroborates the idea that the secretions of the aesthetasc-associated epidermal glands are involved in immune responses. We propose that the mucous covering the aesthetascs in Coenobita takes part in antimicrobial defense and at the same time provides the moisture essential for odor perception in terrestrial hermit crabs. We conclude that the morphological modifications of the aesthetasc-associated epidermal glands as well as the functional characteristics of their secretions are important adaptations to a terrestrial lifestyle. [ABSTRACT FROM AUTHOR]

Background: Paired mushroom bodies, an unpaired central complex, and bilaterally arranged clusters of olfactory glomeruli are among the most distinctive components of arthropod neuroarchitecture. Mushroom body neuropils, unpaired midline neuropils, and olfactory glomeruli also occur in the brains of some polychaete annelids, showing varying degrees of morphological similarity to their arthropod counterparts. Attempts to elucidate the evolutionary origin of these neuropils and to deduce an ancestral ground pattern of annelid cerebral complexity are impeded by the incomplete knowledge of annelid phylogeny and by a lack of comparative neuroanatomical data for this group. The present account aims to provide new morphological data for a broad range of annelid taxa in order to trace the occurrence and variability of higher brain centers in segmented worms. Results: Immunohistochemically stained preparations provide comparative neuroanatomical data for representatives from 22 annelid species. The most prominent neuropil structures to be encountered in the annelid brain are the paired mushroom bodies that occur in a number of polychaete taxa. Mushroom bodies can in some cases be demonstrated to be closely associated with clusters of spheroid neuropils reminiscent of arthropod olfactory glomeruli. Less distinctive subcompartments of the annelid brain are unpaired midline neuropils that bear a remote resemblance to similar components in the arthropod brain. The occurrence of higher brain centers such as mushroom bodies, olfactory glomeruli, and unpaired midline neuropils seems to be restricted to errant polychaetes. Conclusions: The implications of an assumed homology between annelid and arthropod mushroom bodies are discussed in light of the 'new animal phylogeny'. It is concluded that the apparent homology of mushroom bodies in distantly related groups has to be interpreted as a plesiomorphy, pointing towards a considerably complex neuroarchitecture inherited from the last common ancestor, Urbilateria. Within the annelid radiation, the lack of mushroom bodies in certain groups is explained by widespread secondary reductions owing to selective pressures unfavorable for the differentiation of elaborate brains. Evolutionary pathways of mushroom body neuropils in errant polychaetes remain enigmatic. [ABSTRACT FROM AUTHOR]

Background: Invertebrate nervous systems are highly disparate between different taxa. This is reflected in the terminology used to describe them, which is very rich and often confusing. Even very general terms such as 'brain', 'nerve', and 'eye' have been used in various ways in the different animal groups, but no consensus on the exact meaning exists. This impedes our understanding of the architecture of the invertebrate nervous system in general and of evolutionary transformations of nervous system characters between different taxa. Results: We provide a glossary of invertebrate neuroanatomical terms with a precise and consistent terminology, taxon-independent and free of homology assumptions. This terminology is intended to form a basis for new morphological descriptions. A total of 47 terms are defined. Each entry consists of a definition, discouraged terms, and a background/comment section. Conclusions: The use of our revised neuroanatomical terminology in any new descriptions of the anatomy of invertebrate nervous systems will improve the comparability of this organ system and its substructures between the various taxa, and finally even lead to better and more robust homology hypotheses. [ABSTRACT FROM AUTHOR]

Background: The Chaetognatha (arrow worms) are a group of marine carnivores whose phylogenetic relationships are still vigorously debated. Molecular studies have as yet failed to come up with a stable hypothesis on their phylogenetic position. In a wide range of metazoans, the nervous system has proven to provide a wealth of characters for analysing phylogenetic relationships (neurophylogeny). Therefore, in the present study we explored the structure of the ventral nerve centre ("ventral ganglion") in Sagitta setosa with a set of histochemical and immunohistochemical markers. Results: In specimens that were immunolabeled for acetylated-alpha tubulin the ventral nerve centre appeared to be a condensed continuation of the peripheral intraepidermal nerve plexus. Yet, synapsin immunolocalization showed that the ventral nerve centre is organized into a highly ordered array of ca. 80 serially arranged microcompartments. Immunohistochemistry against RFamide revealed a set of serially arranged individually identifiable neurons in the ventral nerve centre that we charted in detail. Conclusion: The new information on the structure of the chaetognath nervous system is compared to previous descriptions of the ventral nerve centre which are critically evaluated. Our findings are discussed with regard to the debate on nervous system organisation in the last common bilaterian ancestor and with regard to the phylogenetic affinities of this Chaetognatha. We suggest to place the Chaetognatha within the Protostomia and argue against hypotheses which propose a deuterostome affinity of Chaetognatha or a sister-group relationship to all other Bilateria. [ABSTRACT FROM AUTHOR]

Stemmata of strepsipteran insects represent the smallest arthropod eyes known, having photoreceptors which form fused rhabdoms measuring an average size of 1.69 × 1.21 × 1.04 μm and each occupying a volume of only 0.97-1.16 μm 3 . The morphology of the stemmata of the extremely miniaturized first instar larva of Stylops ovinae (Strepsiptera, Stylopidae) was investigated using serial-sectioning transmission electron microscopy (ssTEM). Our 3D reconstruction revealed that, despite different proportions, all three stemmata maintain the same organization: a biconvex corneal lens, four corneagenous cells and five photoreceptor (retinula) cells. No pigment-containing cell-types were found to adjoin the corneagenous cells. Whereas the retinula cells are adapted to the limited space by having laterally bulged median regions, containing mitochondria and the smallest nuclei yet reported for arthropods (1.37 μm 3 ), special adaptations are found in the corneagenous cells which have cell volumes down to 1 μm 3 . The corneagenous cells lack nuclei and pigment granules and bear only a few mitochondria (up to three) or none at all. Morphological adaptations due to miniaturization are discussed in the context of photoreceptor function and the visual needs of the larva., (Copyright © 2021 Elsevier Ltd. All rights reserved.)

The corona ciliata of Chaetognatha (arrow worms) is a circular or elliptical groove lined by a rim from which multiple lines of cilia emanate, located dorsally on the head and/or trunk. Mechanoreception, chemosensation, excretion, respiration, and support of reproduction have been suggested to be its main functions. Here we provide the first experimental evidence that the cilia produce significant water flow, and the first visualisation and quantification of this flow. In Spadella cephaloptera, water is accelerated toward the corona ciliata from dorsal and anterior of the body in a funnel-shaped pattern, and expelled laterally and caudally from the corona, with part of the water being recirculated. Maximal flow speeds were approximately 140μms -1 in adult specimens. Volumetric flow rate was Q=0.0026μls -1 . The funnel-shaped directional flow can possibly enable directional chemosensation. The flow measurements demonstrate that the corona ciliata is well suited as a multifunctional organ., (Copyright © 2017 Elsevier GmbH. All rights reserved.)

The arthropodium is the key innovation of arthropods. Its various modifications are the outcome of multiple evolutionary transformations, and the foundation of nearly endless functional possibilities. In contrast to hexapods, crustaceans, and even chelicerates, the spectrum of evolutionary transformations of myriapod arthropodia is insufficiently documented and rarely scrutinized. Among Myriapoda, Chilopoda (centipedes) are characterized by their venomous forcipules-evolutionarily transformed walking legs of the first trunk segment. In addition, the posterior end of the centipedes' body, in particular the ultimate legs, exhibits a remarkable morphological heterogeneity. Not participating in locomotion, they hold a vast functional diversity. In many centipede species, elongation and annulation in combination with an augmentation of sensory structures indicates a functional shift towards a sensory appendage. In other species, thickening, widening and reinforcement with a multitude of cuticular protuberances and glandular systems suggests a role in both attack and defense. Moreover, sexual dimorphic characteristics indicate that centipede ultimate legs play a pivotal role in intraspecific communication, mate finding and courtship behavior. We address ambiguous identifications and designations of podomeres in order to point out controversial aspects of homology and homonymy. We provide a broad summary of descriptions, illustrations, ideas and observations published in past 160 years, and propose that studying centipede ultimate legs is not only essential in itself for filling gaps of knowledge in descriptive morphology, but also provides an opportunity to explore diverse pathways of leg transformations within Myriapoda., Competing Interests: The authors declare there are no competing interests.

The primary function of male copulatory organs is depositing spermatozoa directly into the female reproductive tract. Typical male copulatory organs are sensorily active. This is in contrast to the copulatory organs of male spiders (i.e. palpal bulbi), which have been assumed to lack nerves and muscles until recently. Neurons have been found within the bulbus of the spider Hickmania troglodytes, a taxon basal to all Neocribellata. We provide the first evidence for neurons and an internalized multi-sensillar sensory organ in the bulbus of an entelegyne spider (Philodromus cespitum). The sensory organ likely provides mechanical or chemical feedback from the intromitting structure, the embolus. We found further neurons associated with two glands within the bulbus, one of which is likely responsible for sperm extrusion during mating. These findings provide a new framework for studies on reproductive behaviour and sexual selection in spiders.

Harvestmen (Arachnida, Opiliones) are especially dependent on chemical cues and are often regarded as animals that rely mainly on contact chemoreception. Information on harvestman sensilla is scarce when compared to other arachnid orders, especially concerning internal morphology. Using scanning (SEM) and transmission (TEM) electron microscopy, we investigated tarsal sensilla on the distal tarsomeres (DT) of all leg pairs in Heteromitobates discolor (Laniatores, Gonyleptidae). Furthermore, we explored the typological diversity of sensilla present on the DT I and II in members of the suborder Laniatores, which include two thirds of the formally described opilionid fauna, using species from 17 families representing all main laniatorian lineages. Our data revealed that DT I and II of H. discolor are equipped with wall-pored falciform hairs (two types), wall-pored sensilla chaetica (two types) and tip-pored sensilla chaetica, while DT III and IV are mainly covered with trichomes (non-sensory) and tip-pored sensilla chaetica. The ultrastructural characteristics support an olfactory function for all wall-pored sensilla and a dual gustatory/mechanoreceptive function for tip-pored sensilla chaetica. Based on our comparative SEM survey, we show that wall-pored sensilla occur in all investigated Laniatores, demonstrating their widespread occurrence in the suborder and highlighting the importance of both legs I and II as the sensory appendages of laniatorean harvestmen. Our results provide the first morphological evidence for olfactory receptors in Laniatores and suggest that olfaction is more important for harvestmen than previously thought., (Copyright © 2016 Elsevier Ltd. All rights reserved.)

Adaptations to a terrestrial lifestyle occurred convergently multiple times during the evolution of the arthropods. This holds also true for the "true crabs" (Brachyura), a taxon that includes several lineages that invaded land independently. During an evolutionary transition from sea to land, animals have to develop a variety of physiological and anatomical adaptations to a terrestrial life style related to respiration, reproduction, development, circulation, ion and water balance. In addition, sensory systems that function in air instead of in water are essential for an animal's life on land. Besides vision and mechanosensory systems, on land, the chemical senses have to be modified substantially in comparison to their function in water. Among arthropods, insects are the most successful ones to evolve aerial olfaction. Various aspects of terrestrial adaptation have also been analyzed in those crustacean lineages that evolved terrestrial representatives including the taxa Anomala, Brachyura, Amphipoda, and Isopoda. We are interested in how the chemical senses of terrestrial crustaceans are modified to function in air. Therefore, in this study, we analyzed the brains and more specifically the structure of the olfactory system of representatives of brachyuran crabs that display different degrees of terrestriality, from exclusively marine to mainly terrestrial. The methods we used included immunohistochemistry, detection of autofluorescence- and confocal microscopy, as well as three-dimensional reconstruction and morphometry. Our comparative approach shows that both the peripheral and central olfactory pathways are reduced in terrestrial members in comparison to their marine relatives, suggesting a limited function of their olfactory system on land. We conclude that for arthropod lineages that invaded land, evolving aerial olfaction is no trivial task.

A generic character of the genus Spiophanes (Annelida, Sedentaria: Spionidae) is the presence of parapodial glandular organs. Parapodial glandular organs in Spiophanes species include secretory cells with cup-shaped microvilli, similar to those present in deep-sea inhabiting vestimentiferans and frenulate Siboglinidae. These cells are supposed to secrete β-chitin for tube-building. In this study, transverse histological and/or ultrathin sections of parapodial glandular organs and tubes of Spiophanes spp. as well as of Glandulospio orestes (Spionidae) and Owenia fusiformis (Oweniidae) were examined. Fluorescent markers together with confocal laser scanning microscopy, and Raman spectroscopy were used to detect chitin in the parapodial glandular organs of Spiophanes and/or in the glands of Owenia and Glandulospio. Tubes of these taxa were tested for chitin to elucidate the use of it for tube-building. The examinations revealed a distinct labelling of the gland contents. Raman spectroscopy documented the presence of β-chitin in both gland types of Spiophanes. The tubes of Spiophanes were found to have a grid-like structure that seems to be built with this β-chitin. Tests of tubes of Dipolydora quadrilobata (Spionidae) for chitin were negative. However, the results of our study provide strong evidence that Spiophanes species, O. fusiformis and probably also G. orestes produce chitin and supposedly use it for tube-building. This implies that the production of chitin and its use as a constituent part of tube-building is more widespread among polychaetes as yet known. The histochemical data presented in this study support previous assumptions inferring homology of parapodial glandular organs of Spionidae and Siboglinidae based on ultrastructure. Furthermore, transmission electron microscopy-based evidence of secretory cells with nail-headed microvilli in O. fusiformis suggests homology of parapodial grandular organs across annelids including Sipuncula., (© 2015 Wiley Periodicals, Inc.)

Notostigmophora (Scutigeromorpha) exhibit a special tracheal system compared to other Chilopoda. The unpaired spiracles are localized medially on the long tergites and open into a wide atrium from which hundreds of tracheal tubules originate and extend into the pericardial sinus. Previous investigators reported that the tracheal tubules float freely in the hemolymph. However, here we show for the first time that the tracheal tubules are anchored to a part of the pericardial septum. Another novel finding is this part of the pericardial septum is structured as an aggregated gland on the basis of its specialized epithelium being formed by hundreds of oligocellular glands. It remains unclear whether the pericardial septum has a differently structure in areas that lack a connection with tracheal tubules. The tracheal tubules come into direct contact with the canal cells of the glands that presumably secrete mucous substances covering the entire luminal cuticle of the tracheal tubules. Connections between tracheae and glands have not been observed in any other arthropods.

Crustaceans have successfully adapted to a variety of environments including fresh- and saltwater as well as land. Transition from an aquatic to a terrestrial lifestyle required adaptations of the sensory equipment of an animal, particularly in olfaction, where the stimulus itself changes from hydrophilic to mainly hydrophobic, air-borne molecules. Hermit crabs Coenobita spp. (Anomura, Coenobitidae) have adapted to a fully terrestrial lifestyle as adults and have been shown to rely on olfaction in order to detect distant food items. We observed that the specialized olfactory sensilla in Coenobita, named aesthetascs, are immersed in a layer of mucous-like substance. We hypothesized that the mucous is produced by antennal glands and affects functioning of the aesthetascs. Using various microscopic and histochemical techniques we proved that the mucous is produced by aesthetasc-associated epidermal glands, which we consider to be modified rosette-type aesthetasc tegumental glands known from aquatic decapods. These epidermal glands in Coenobita are multicellular exocrine organs of the recto-canal type with tubulo-acinar arrangement of the secretory cells. Two distinct populations of secretory cells were clearly distinguishable with light and electron microscopy. At least part of the secretory cells contains specific enzymes, CUB-serine proteases, which are likely to be secreted on the surface of the aesthetasc pad and take part in antimicrobial defense. Proteomic analysis of the glandular tissue corroborates the idea that the secretions of the aesthetasc-associated epidermal glands are involved in immune responses. We propose that the mucous covering the aesthetascs in Coenobita takes part in antimicrobial defense and at the same time provides the moisture essential for odor perception in terrestrial hermit crabs. We conclude that the morphological modifications of the aesthetasc-associated epidermal glands as well as the functional characteristics of their secretions are important adaptations to a terrestrial lifestyle.

In Chilopoda, solitary epidermal glands are composed of a couple of cells only. These glands are highly abundant on the entire body surface and are distributed throughout the single-layered epidermis. Some authors provided more or less comprehensive observations on the structure of epidermal glands of specific chilopod taxa. However, no information is hitherto available on the ultrastructural diversity of these glands. Furthermore, potential homologies of these chilopod epidermal glands and of their characteristic cellular components remain unknown. Based on our results, we are now able to distinguish two types of epidermal glands in Chilopoda that can be clearly distinguished by their structure and the course of their conducting canal: recto-canal epidermal glands (rceg) and flexo-canal epidermal glands (fceg). In the present paper, we focus on the rceg. We examined the ultrastructural organization of these glands in the head region and on the anterior trunk segments of various representatives of the five extant chilopod orders by light- and electron-microscopy. According to our terminology, rceg consist of up to five different cell types including: a) distal canal cells, b) proximal canal cells, c) intermediary cells, and d) two different types of secretory cells. Intermediary and canal cells form a common conducting canal. The rceg may taxon-specifically differ in relative size and subcellular architecture, but all have the following features in common: 1) a wide distribution on various body regions among all five chilopod subtaxa, 2) the straight, broad and locally dilated conducting canal surrounded by closely packed microvilli or microvilliform infoldings around the apex of the canal cell(s), and 3) the tendency to aggregate to form compound glandular organs of massive size and complexity. Tricellular glandular units established by three different cell types are observed in Scutigeromorpha and Geophilomorpha, whereas four cell types constitute rceg in Lithobiomorpha and Craterostigmomorpha. Five different cell types per glandular unit are found only in Scolopendromorpha. The partial cuticularization of the lower part of the conducting canal formed by the intermediary cell, as found in Chilopoda, differs from the pattern described for equivalent euarthropod epidermal glands, as for instance in Hexapoda. Their wide distribution in Chilopoda and Progoneata makes it likely that tricellular rceg were at least present in the last common ancestor of the Myriapoda. Concerning Chilopoda, the evolution of highly diverse rceg is well explained on the basis of the Pleurostigmophora concept. Glands of the recto-canal type are also found in other arthropods. The paper discusses cases where homology of rceg and also fceg may be assumed beyond Myriapoda and briefly evaluates the potentials and the still-to-be-solved issues prior to use them as an additional character system to reconstruct the phylogeny of the Euarthropoda., (Copyright © 2013 Elsevier Ltd. All rights reserved.)

Emerging evidence suggests that Chaetognatha represent an evolutionary lineage that is the sister group to all other Protostomia thus promoting these animals as a pivotal model for our understanding of bilaterian evolutionary history. We have analyzed the proliferation of neuronal progenitor cells in the developing ventral nerve center (VNC) of Spadella cephaloptera hatchlings. To that end, for the first time in Chaetognatha, we performed in vivo incorporation experiments with the S-phase specific mitosis marker bromodeoxyuridine (BrdU). Our experiments provide evidence for a high level of mitotic activity in the VNC for ca. 3 days after hatching. Neurogenesis is carried by presumptive neuronal progenitor cells that cycle rapidly and most likely divide asymmetrically. These progenitors are arranged in a distinct grid-like geometrical pattern including about 35 transverse rows. Considering Chaetognaths to be an early offshoot of the protostome lineage we conclude that the presence of neuronal progenitor cells with asymmetric division seems to be a feature that is rooted deeply in the Metazoa. In the light of previous evidence indicating the presence of serially iterated peptidergic neurons with individual identities in the chaetognath VNC, we discuss if these neuronal progenitor cells give rise to distinct lineages. Furthermore, we evaluate the serially iterated arrangement of the progenitor cells in the light of evolution of segmentation., (Copyright © 2013 Wiley Periodicals, Inc.)

Parapodial glandular organs (PGOs) of Spiophanes (Polychaeta: Spionidae) were studied using light and electron microscopy. These organs are found in parapodia of the mid body region, starting on chaetiger 5 and terminating with the appearance of neuropodial hooks (chaetiger 14 or 15 in adult individuals). Large PGOs in anterior chaetigers display different species-specific types of openings whereas small PGOs in posterior parapodia of the mid body region always open in a simple vertical slit. Each PGO is composed of three main complexes: (1) the glandular sac with several distinct epithelia of secretory cells and secretory cell complexes and the reservoir filled with fibrous material, (2) the gland-associated chaetal complex (including the region of chaetoblasts and follicle cells, follicular canals, two chaetal collector canals, the combined conducting canal, the chaetal spreader including the opening of the glandular organ with associated type-1 secretory cells, and the gland-associated chaetae), and (3) a bilayered musculature surrounding the gland. A considerable number of different cell types are involved in the secretory activity, in the guidance of the gland-associated chaetae, and in the final expulsion of the fibrous secretion at the opening slit. Among these different cell types the type-5 secretory cells of the proximal glandular complex with their cup-shaped microvilli emanating thick microfibrils into the lumen of the glandular sac are most conspicuous. Secretory cells with cup-shaped microvilli being involved in the production of β-chitin microfibrils have so far only been reported from some representatives of the deep-sea inhabiting Siboglinidae (Polychaeta). We suggest that the gland-associated chaetae emerging from inside the PGOs of Spiophanes are typical annelid chaetae formed by chaetoblasts and follicle cells. Functional morphology implies the crucial role of PGOs in tube construction. Furthermore, the PGOs are discussed in consideration of phylogenetic aspects., (Copyright © 2011 Wiley Periodicals, Inc.)

With a body length of only 0.3-0.4 mm, the parasitoid wasp Trichogramma evanescens (Westwood) is one of the smallest insects known. Yet, despite its diminutive size, it possesses compound eyes that are of oval shapes, measuring across their long axes in dorsoventral direction 63.39 and 71.11 μm in males and females, respectively. The corresponding facet diameters are 5.90 μm for males and 6.39 μm for females. Owing to the small radii of curvature of the eyes in males (34.59 μm) and females (42.82 μm), individual ommatidia are short with respective lengths of 24.29 and 34.97 μm. The eyes are of the apposition kind, and each ommatidium possesses four cone cells of the eucone type and a centrally fused rhabdom, which throughout its length is formed by no more than eight retinula cells. A ninth cell occupies the place of the eighth retinula cell in the distal third of the rhabdom. The cone is shielded by two primary and six secondary pigment cells, all with no apparent extensions to the basement membrane, unlike the case in larger hymenopterans. The regular and dense packing of the rhabdoms reflects an effective use of space. Calculations on the optics of the eyes of Trichogramma suggest that the eyes need not be diffraction limited, provided they use mostly shorter wavelengths, that is, UV light. Publications on the visual behavior of these wasps confirm Trichogramma's sensitivity to UV radiation. On the basis of our findings, some general functional conclusions for very small compound eyes are formulated.

The eyes of adult Phryssonotus platycephalus (Synxenidae) and Polyxenus lagurus (Polyxenidae) were investigated by light and electron microscopy. At each side of the head, various numbers of eye cups are situated on projections, the eye hills. The eye cups of P. platycephalus and P. lagurus are similarly structured and considered homologous sense organs. Each corneal lens is biconvex and formed by four to six pigmented corneagenous cells with their nuclei displaced towards the mid-periphery of the eye cup. The corneal surface displays a conspicuous nanostructure of fingerprint-like ridges in P. platycephalus. However, the corneal surface appears smooth in P. lagurus. In P. platycephalus. A rudimentary crystalline cone is observed in each eye cup, always produced by a constant number of three eucone cells. The crystalline cone is wedged between the corneal lens and the distal rhabdom and consists of three distinct compartments. Each cone compartment is connected to the voluminous proximal nuclear region by one elongated cytoplasmic process, which runs through the infraretinular space. A dual type retinula is always arranged in two distinct horizontal cell layers. The distal retinula contains an unfixed number of four to five cells in P. lagurus, whereas it contains five to eight cells in P. platycephalus. The distal retinula cells form a large and fused axial rhabdom. A constant number of three proximal retinula cells give rise to a small axial rhabdom, which looks more or less triangular in cross sections. The basal matrix is rather thin, inconspicuous and lines the bases of the eye cups. The ultrastructure of the eye cups of P. platycephalus resembles that observed in the ommatidia of the centipede Scutigera coleoptrata. The present study lends additional support to the homology of mandibulate ommatidia, because of the common possession of crystalline cone cells and a bilayered dual type retinula in the eye cups of P. platycephalus. Ommatidia or unicorneal eyes that include eucone cells with nuclei displaced outside the cone compartments, as found in Scutigeromorpha and Penicillata, might also be interpreted as an additional autapomorphy of the Myriapoda. The suggested homology of scutigeromorph and penicillate eyes implies that penicillate eye cups have to be considered modified, probably miniaturized ommatidia.

The lateral lens eye of adult Craterostigmus tasmanianus Pocock, 1902 (a centipede from Australia and New Zealand) was examined by light and electron microscopy. An elliptical, bipartite eye is located frontolaterally on either side of the head. The nearly circular posterior part of the eye is characterized by a plano-convex cornea, whereas no corneal elevation is visible in the crescentic anterior part. The so-called lateral ocellus appears cup-shaped in longitudinal section and includes a flattened corneal lens comprising a homogeneous and pigmentless epithelium of cornea-secreting cells. The retinula consists of two kinds of photoreceptive cells. The distribution of the distal retinula cells is highly irregular. Variable numbers of cells are grouped together in multilayered, thread-like unions extending from the ventral and dorsal margins into the center of the eye. Around their knob-like or bilobed apices the distal retinula cells give rise to fused polymorphic rhabdomeres. Both everse and inverse cells occur in the distal retinula. Smaller, club-shaped proximal retinula cells are present in the second (limited to the peripheral region) and proximal third of the eye, where they are arranged in dual cell units. In its apical region each unit produces a small, unidirectional rhabdom of interdigitating microvilli. All retinula cells are surrounded by numerous sheath cells. A thin basal lamina covers the whole eye cup, which, together with the distal part of the optic nerve, is wrapped by external pigment cells filled with granules of varying osmiophily. The eye of C. tasmanianus seemingly displays very high complexity compared to many other hitherto studied euarthropod eyes. Besides the complex arrangement of the entire retinula, the presence of a bipartite eye cup, intraocellar exocrine glands, inverse retinula cells, distal retinula cells with bilobed apices, separated pairs of proximal retinula cells, medio-retinal axon bundles, and the formation of a vertically partitioned, antler-like distal rhabdom represent apomorphies of the craterostigmomorph eye. These characters therefore collectively underline the separate position of the Craterostigmomorpha among pleurostigmophoran centipedes. The remaining retinal features of C. tasmanianus agree with those known from other chilopod eyes and, thus, may be considered plesiomorphies. Characters like the unicorneal eye cup, sheath cells, and proximal rhabdomeres with interdigitating microvilli were already present in the ground pattern of the Pleurostigmophora. Other retinal features were developed in the ancestral lineage of the Phylactometria (e.g., large elliptical eyes, external pigment cells, polygonal sculpturations on the corneal surface). The homology of all chilopod eyes (including Notostigmophora) is based principally on the possession of a dual type retinula., ((c) 2006 Wiley-Liss, Inc.)

We describe the ultrastructural organization of the anal organs of Craterostigmus tasmanianus, which are located on the ventral side of the bivalvular anal capsule. Each part of the capsule bears four pore fields with several anal pores. The pores lead into a pore canal, which is surrounded by the single-layered epithelium of the anal organs. Each anal organ is composed of four different cell types: transporting cells of the main epithelium, junctional cells, isolated epidermal glands, and the cells forming the pore canal. The transporting cells exhibit infoldings of the outer cell membranes, forming a basal labyrinth and a poorly developed apical complex. The cells are covered by a specialized cuticle with a widened subcuticular layer. Only the cuticle of the main epithelium is covered by a mucous layer, secreted by the epidermal glands. The ultrastructural organization of the anal organ is comparable to the coxal and anal organs of other pleurostigmophoran Chilopoda. It is likely that the coxal and anal organs of the Pleurostigmophora are homologous, due to their identical ultrastructural organization. Differences concerning the location on the trunk of Pleurostigmophora are not sufficient to reject a hypothesis of homology. Anal organs are found not only in Craterostigmomorpha, but also in most adult Geophilomorpha, and in larvae and most adults of Lithobiomorpha. The anal organs of C. tasmanianus are thought to play an important role in the uptake of atmospheric water. J. Morphol.