Adults of European ant‐like stone beetles (Coleoptera: Staphylinidae: Scydmaeninae) Scydmaenus tarsatus Müller & Kunze and Scydmaenus hellwigii (Herbst) prey on soft‐bodied arthropods

Scydmaenine beetles are commonly described as predators specialized in capturing and feeding on armored mites of the order Oribatida, and documented cases of feeding on other live arthropods have not been known. Based on laboratory observations and a broad choice of Acari (armored and soft‐bodied) and other soil arthropods, food preferences and associated behavior of two scydmaenine species are clarified and described. Adults of Scydmaenus tarsatus ignored oribatid and mesostigmatan mites, but readily attacked and fed on a soft‐bodied Rhizoglyphus sp. (Acaridae), and on small springtails, especially on Ceratophysella denticulata (Hypogastruridae). A water drinking behavior was observed for this species, not reported previously in any Staphylinidae. Scydmaenus hellwigii ignored all tested Acari (including Rhizoglyphus) and scavenged on dead neanurine collembolans or freshly cut pieces of large springtails; a long term culture was maintained by feeding beetles with isotomid springtails. Previously reported strict specialization of Scydmaenus as a predator on Oribatida was not confirmed and it is concluded that the studied species feed on live soft‐bodied organisms and scavenge on dead arthropods.     

Cephalic modifications in dimorphic dwarf spiders of the genus oedothorax (Erigoninae,Linyphiidae, Araneae) and their evolutionary implications

Reproductive competition among males selects for a broad variety of strategies and traits from mate guarding to nuptial food gifts. Males of many dwarf spider species possess conspicuous secondary cephalic modifications, and the few studies available suggest that these cephalic structures are connected to extensive glandular tissue. Because females were observed to contact the male head structures during mating, these traits may have evolved in the context of sexual selection. We investigated the structure, glandular equipment, and sensory equipment of the cephalic regions of several species of the dwarf spider genus Oedothorax with varying degrees of sexual dimorphism using light and electron microscopy. In one Oedothorax species, there are two male morphs that exhibit a cephalic modification (O. gibbosus gibbosus) or not (O. gibbosus tuberosus). Our study demonstrates that all males investigated produce cephalic secretions, irrespective of the morphology of their cephalic region, however, they may differ in amount of secretion and in cellular organization. In males of O. apicatus, O. gibbosus gibbosus and O. retusus the gland cells are very abundant in the area of a cephalic hump, whereas in the less conspicuous O. agrestis, and O. gibbosus tuberosus the gland cells are restricted to a small area behind the ocular region or include the ocular region as in O. fuscus. The glandular tissue consists of two gland types in O. agrestis, O. fuscus, O. gibbosus tuberosus and O. retusus and of only one type in O. apicatus and O. gibbosus gibbosus. The setae present on the head structure of all species seem to function as mechano‐ and/or chemoreceptors. The implications of our findings for the evolution of secretory head structures are discussed along with their potential for driving speciation. J. Morphol. 2011. © 2011Wiley‐Liss, Inc.     

Response by two species of crabs to a rhizocephalan extract

Repeated injections of crude aqueous extracts of roots of the sacculinid, Loxothylacus panopei, were performed in two xanthid species: Rhithropanopeus harrisii, the rhizocephalan's usual crab host, and Panopeus herbstii, a nonparasitized crab in North Carolina waters but which is frequently infected by the same parasite along southern Atlantic coasts. Testes, androgenic glands, and the different parts of the central nervous system were observed histologically. The rhizocephalan extracts induced alterations of the testicular germinative zone for both crab species; mesodermal cells and gonia degenerate. The sinus glands of the central nervous system appear devoid of secretions. The comparison of the sensitivity of the two species with respect to the parasite shows that, at the level of the genital apparatus, the modifications are more severe in P. herbstii. Hypotheses concerning the noninfestation of P. herbstii in brackish waters of North Carolina are proposed following a brief survey of the physiological and ecological characteristics of both species.     

Cephalic anatomy of Sinentomon erythranum Yin (Protura : Sinentomidae)

The structure of the head capsule, endoskeletal structures, oral folds, mouthparts, foregut, maxillary and labial glands, and the central nervous system of the Chinese proturan Sinentomon erythranum Yin (Protura : Sinentomidae) have been described. The most significant features are the unusual thickness of the cuticle with numerous serrated lines, the reduction of sutures and the absence of linea ventralis, the simplification of gnathal pieces, and the reduction of the musculature. A comparison with other Protura and Apterygota leads to some morphological and phylogenetical interpretations of the cephalic structures, in particular those concerning the entotrophy.     

Profound head modifications in Claviger testaceus (Pselaphinae,Staphylinidae, Coleoptera) facilitate integration into communities of ants

Clavigeritae is a group of obligate myrmecophiles of the rove beetle subfamily Pselaphinae (Staphylinidae). Some are blind and wingless, and all are believed to depend on ant hosts through feeding by trophallaxis. Phylogenetic hypotheses suggest that their ancestors, as are most pselaphines today, were free-living predators. Morphological alterations required to transform such beetles into extreme myrmecophiles were poorly understood. By studying the cephalic morphology of Claviger testaceus, we demonstrate that profound changes in all mouthpart components took place during this process, with a highly unusual connection of the maxillae to the hypopharynx, and formation of a uniquely transformed labium with a vestigial prementum. The primary sensory function of the modified maxillary and labial palps is reduced, and the ventral mouthparts transformed into a licking/‘sponging’ device. Many muscles have been reduced, in relation to the coleopteran groundplan or other staphylinoids. The head capsule contains voluminous glands whose appeasement secretions are crucial for the beetle survival in ant colonies. The brain, in turn, has been shifted into the neck region. The prepharyngeal dilator is composed of an entire series of bundles. However, the pharynx does not show any peculiar adaptations to taking up liquid food. We demonstrate that far-reaching cephalic modifications characterize C. testaceus, and that the development of appeasement glands and adaptation of the mouthparts to trophallaxis determine the head architecture of this extreme myrmecophile.     

Generation gland morphology in cordylid lizards: An evolutionary perspective

To elucidate the functional significance of the three distinct types of generation glands that have been identified among cordylid lizards, we mapped gland type to the terminal taxa in the most recent phylogenetic tree for the Cordylidae. We used the phylogenetic programme Mesquite and applied the principle of parsimony to infer character states for the ancestral nodes in the tree. For those species where information on gland type was not available from the literature, we conducted a histological investigation of generation gland morphology, using standard histological techniques. We included two species of the sister family Gerrhosauridae in the analysis to serve as outgroups. In both Gerrhosaurus typicus and G. flavigularis, scales immediately anterior to the femoral pores displayed glandular activity, but differed from generation glands of cordylids in the absence of mature glandular generations. Among the cordylids investigated, we identified a fourth type of generation gland in Pseudocordylus subviridis, P. spinosus, and in the two Hemicordylus species, one where the glands consistently comprise of two mature glandular generations. In H. capensis, both single‐ and two‐layer type glands are present. Our reconstruction of ancestral character states suggests a minimum of six transformations from one gland type to another during the evolutionary history of the family. The reconstruction furthermore suggests that the single‐layer type gland reappeared at least once (in Hemicordylus) in the Cordylinae after having been lost. The reconstruction also unequivocally shows that the pit‐like multiple‐layer type gland evolved directly from the single‐layer type and not from the protruding multiple‐layer type. The two‐layer type gland appears to be an intermediary condition between the multiple‐layer and single‐layer types. The evolutionary transformation of generation gland type appears to be linked to changes in lifestyle and associated changes in degree of territoriality and the need for chemical communication. J. Morphol. 275:456–464, 2014. © 2013 Wiley Periodicals, Inc.     

Ultrastructure of the cerebral glands in Scolopendra cingulata Latr., Cryptops savignyi leach and C. hortensis Leach (Myriapoda : Scolopendromorpha)

The fine structure of the cerebral glands (cephalic neurohaemal organs) was investigated in the scolopendromorphs, Scolopendra sp. and Cryptops sp. (Myriapoda Scolopendromorpha). As in other Chilopoda, these organs are composed of glandular cells and axons, originating from the brain. Differences between the 2 genera studied concern: (i) The general structure (axons and glandular cells are mixed in Scolopendra and separated in Cryptops). (ii) The different cell types (I type of glial cell and 2 of glandular cells in Scolopendra while respectively 2 and 3 are persent in Cryptops. Fruthermore, this last genus has a less-developed Golgi apparatus and shows in one of the glandular cell types an intranuclear vacuole, sometimes related to a dilated perinuclear space. Physiological significance of such a structure remains unknown). (iii) The number of axonal types (less numerous in Scolopendra than in Cryptops). The 2 genera show little exocytosis profiles, both in axons and in glandular cells.     

Cephalic secretion release in the male dwarf spider Oedothorax retusus (Linyphiidae: Erigoninae): An ultrastructural analysis

Secondary sexual traits in males can extend to glandular structures that play a role during courtship and mating. In dwarf spiders (Linyphiidae, Erigoninae), glandular secondary sexual traits are particularly common. Males are characterized by cephalic modifications which produce secretions that females contact with their mouthparts during courtship and/or copulation. We used the dwarf spider Oedothorax retusus as a model species to investigate if and when the contents of the glands are released during a mating sequence and if so, if the gland reservoirs are refilled after mating. To this aim, we quantitatively compared the glandular tissue on the ultrastructural level between a) inexperienced males, b) males that performed courtship, c) males immediately after copulation, and d) males three days after mating. We assessed whether the treatment groups differed in the filling state of the conducting canals and receiving canals (reservoir regions) of the glandular units. Our study shows that courting males as well as males three days after mating did not differ significantly from control (inexperienced) males in the presence of secretions. However, males exhibited significantly less secretion immediately after mating. This strongly suggests that the main function of the secretions is gustatorial courtship and not the emission of volatile pheromones for mate attraction as was previously assumed.     

Female sex pheromone of a lichen moth Eilema japonica (Arctiidae, Lithosiinae): Components and control of production

Seven candidates for components of the female sex pheromone of Eilema japonica (Arctiidae, Lithosiinae) were detected in an extract of pheromone glands with a gas chromatograph-electroantennographic detector. The compounds were identified as (Z,Z)-6,9-icosadiene (D20), (Z,Z)-6,9-henicosadiene (D21), (Z,Z,Z)-3,6,9-henicosatriene (T21), (Z,Z)-6,9-docosadiene (D22), (Z,Z,Z)-3,6,9-docosatriene (T22), (Z,Z)-6,9-tricosadiene (D23), and (Z,Z,Z)-3,6,9-tricosatriene (T23). Assays using synthetic lures in a wind tunnel showed that D21 (proportion, 0.39), T21 (0.08), D22 (0.27), and T22 (0.26) are important for evoking full behavioral responses from the males. Titers of the pheromone components did not show clear temporal fluctuations. Moreover, decapitation of the female moth had no effect on the titers of pheromone components in the pheromone gland, suggesting that cephalic endocrine factors such as pheromone biosynthesis activating neuropeptide (PBAN) are not involved in the control of pheromone biosynthesis in this species.     

The head of Merope tuber (Meropeidae) and the phylogeny of Mecoptera (Hexapoda)

External and internal features of the head of adults of Merope tuber were examined and described in detail. The results were compared to conditions found in other members of Mecoptera and other antliophoran lineages. A list of characters of different body parts and life stages is presented. The parsimony analysis and a recent evaluation of thoracic features suggest a basal placement of Merope within monophyletic Pistillifera. The monophyly of Mecoptera was not supported by our data set. Nannochoristidae (Nannomecoptera) was placed as sistertaxon of a clade comprising Diptera and Siphonaptera. Cephalic features supporting this group are modifications of the mouthparts linked to feeding on liquid substrates. Considering recent results of extensive morphological and molecular investigations we consider this placement of Nannochoristidae and the implied mecopteran paraphyly as a possible artefact. Potential cephalic autapomorphies of Mecoptera are the presence of a tooth-like projection of the gena and a prepharyngeal tube, the absence of M. frontolabralis, and the origin of M. tentoriooralis on the middle region of the anterior tentorial arm. Despite of the conspicuous morphological differences between Caurinus and the other boreid genera the family forms a well supported clade. A sistergroup relationship between Boreidae and Pistillifera is confirmed. A unique synapomorphy is the presence of specialized dilator muscles of the salivary duct. The reconstruction of the relationships of the pistilliferan taxa is strongly impeded by a serious lack of morphological data. However, a group comprising Eomeropidae, Choristidae, Apteropanorpidae, Panorpidae and Panorpodidae is supported in our analyses. Further well documented anatomical data are needed for a reliable reconstruction of mecopteran relationships. The collecting and morphological study of larvae should also have high priority. Inherent problems are extreme secondary modifications of cephalic features of Caurinus and Nannochorista.     

Intervention possible d'un facteur cephalique dans la regeneration des pattes chez les embryons du Phasme,Carausius morosus Br. (Phasmida : Lonchodidae)

Suppressions of endocrine cephalic areas (which contain the presumptive endocrine glands) were performed on young (stage IV₃-V₁) or differentiated (stages VI₈, or VII₁) embryos of Carausius morosus Br.(Phasmida : Lonchodidae). In all these operated embryos (= cephaloprives), amputation of a thoracic leg during the second cuticle formation (stage VI₈), or even after this event (stage VII₁), prevented regeneration. On the other hand, regenerative abilities were preserved when the amputation of the leg was made on embryos deprived of abdominal segments, with similar conditions as for cephaloprives. Consequently, operative shock did not modify regenerative abilities. Because very similar ecdysteroid levels were assayed both in cephaloprives and normal eggs, interference of a possible cephalic factor (perhaps neurosecretion) in the regeneration of the embryonic leg could be envisaged.     

The ultrastructure of the cuticle of Nematoda. II. The cephalic cuticle of Stilbonematinae (Adenophorea,Desmodoridae)

Summary  The ultrastructure of the cephalic cuticle of 6 of 7 representative genera of Stilbonematinae (Eubostrichus, Catanema, Laxus, Robbea, Leptonemella, Stilbonema) is investigated using SEM and TEM techniques. Additionally, one species of Spirinia (Spiriniinae) and one of Desmodora (Desmodorinae) were studied for outgroup comparison. Most of the studied species show modifications of the cephalic cuticle. Furthermore, at least four different pathways have been developed to reinforce the head within Stilbonematinae. Species with a coarsely annulated somatic cuticle (Leptonemella sp., Stilbonema majum, and Desmodora ovigera) developed a rigid, non-annulated cephalic capsule by modifying the main constructing element of the median zone, the ’’ring body.’’ In faintly annulated Laxus oneistus, the annulated cephalic capsule results from a newly inserted ’’block layer’’ between the median and basal zone. The non-annulated cephalic capsule of Robbea sp. is formed by both the block layer and the ring body element. The annulated capsule of Catanema sp. stems from a doubled number of fiber layers within the basal zone. In Spirinia sp., only the amphidial region is strengthened in what could be termed an amphidial shield. All forms with cephalic capsules show mechanisms to keep the oral region pliable. Only Eubostrichus topiarius lacks a reinforcement of the cephalic cuticle. A comparison with the literature is made to elucidate corresponding structures within the cephalic capsules of representatives of Desmodorida. It is demonstrated that the presence of a cephalic cuticle is of no systematic value above the genus level. Accepted: 3 March 1996     

A conserved genetic mechanism specifies deutocerebral appendage identity in insects and arachnids

The segmental architecture of the arthropod head is one of the most controversial topics in the evolutionary developmental biology of arthropods. The deutocerebral (second) segment of the head is putatively homologous across Arthropoda, as inferred from the segmental distribution of the tripartite brain and the absence of Hox gene expression of this anterior-most, appendage-bearing segment. While this homology statement implies a putative common mechanism for differentiation of deutocerebral appendages across arthropods, experimental data for deutocerebral appendage fate specification are limited to winged insects. Mandibulates (hexapods, crustaceans and myriapods) bear a characteristic pair of antennae on the deutocerebral segment, whereas chelicerates (e.g. spiders, scorpions, harvestmen) bear the eponymous chelicerae. In such hexapods as the fruit fly, Drosophila melanogaster, and the cricket, Gryllus bimaculatus, cephalic appendages are differentiated from the thoracic appendages (legs) by the activity of the appendage patterning gene homothorax (hth). Here we show that embryonic RNA interference against hth in the harvestman Phalangium opilio results in homeonotic chelicera-to-leg transformations, and also in some cases pedipalp-to-leg transformations. In more strongly affected embryos, adjacent appendages undergo fusion and/or truncation, and legs display proximal defects, suggesting conservation of additional functions of hth in patterning the antero-posterior and proximo-distal appendage axes. Expression signal of anterior Hox genes labial, proboscipedia and Deformed is diminished, but not absent, in hth RNAi embryos, consistent with results previously obtained with the insect G. bimaculatus. Our results substantiate a deep homology across arthropods of the mechanism whereby cephalic appendages are differentiated from locomotory appendages.     

Ultrastructure,functional morphology and evolution of recto-canal epidermal glands in Myriapoda

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.     

Observations on lesser-known flatworms: temnocephala

Temnocephala novae-zealandie, a flatworm epizoic on crayfish, was examined by light and electron microscopy to investigate the variation within rhabdocoel turbellarians and to provide information on the possible structural modifications in the evolution of parasitism. The sucker is clearly glandular; the tentacular glands are eosinophilic and at the surface store and often release mucus as a coiled thread; the epidermis is clearly not reduced in structure and contains septate junctions in the anterior portion. Light microscope studies documented the presence of ten pair of paranephrocytes (athrocytes). In the laboratory, egg deposition, survival on and apart from the host and another crayfish, and behavior during the host molt were observed.     

The cephalic morphology of free-living and cave-dwelling species of trechine ground beetles from China (Coleoptera,Carabidae)

Morphological adaptations of cave-dwelling organisms including different groups of Coleoptera have fascinated researchers since a long time. Nevertheless, very few detailed documentations of the anatomy of cave beetles using modern techniques are available. In this study, we describe and illustrate external and internal cephalic features of free-living and cave-dwelling trechine carabid beetles using digital microscopy, SEM, micro-CT, and computer-based 3D reconstruction. Morphological characteristics found in three selected species with different habitat preferences are compared. The following derived features distinguish a troglobite species (Sinaphaenops wangorum Ueno et Ran 1998) from a species living in the entrance of caves (Trechiotes perroti Jeannel 1954) and from a fully epigean species (Bembidion sp.): (1) the optic lobes are completely missing; (2) the head capsule, mouth parts, and antennae are distinctly elongated; (3) some muscle attachment areas are shifted; and (4) an additional dilator of the pharynx is present. Despite of conspicuous differences likely related with subterranean habits, the link of some cephalic features to this specific habitat preference remains uncertain.     

Differential expression of Prominin-1 (CD133) and Prominin-2 in major cephalic exocrine glands of adult mice

The major cephalic exocrine glands share many morphological and functional features and so can be simultaneously affected in certain autoimmune- and inherited disorders leading to glandular hypofunction. Phenotypic characterization of these exocrine glands is not only an interesting biological issue, but might also be of considerable clinical relevance. The major salivary and lacrimal glands might therefore be potential subjects of future cell-based regenerative/tissue engineering therapeutic approaches. In the present study, we described the expression of the stem and progenitor cell marker Prominin-1 and those of its paralogue, Prominin-2, in the three pairs of major salivary glands, i.e., submandibular-, major sublingual-, and parotid glands in adult mice. We have also documented their expression in the extraorbital lacrimal and meibomian glands (Glandulae tarsales) of the eyelid (Palpebra). Our analysis revealed that murine Prominin-1 and Prominin-2 were differentially expressed in these major cephalic exocrine organs. Expression of Prominin-1 was found to be associated with the duct system, while Prominin-2 expression was mostly, but not exclusively, found in the acinar compartment of these organs with marked differences among the various glands. Finally, we report that Prominin-2, like Prominin-1, is released into the human saliva associated with small membrane particles holding the potential for future diagnostic applications.     

An active principle from the corpora cardiaca,corpora allata and suboesophageal ganglion of the locust,inducing egg diapause in Bombyx mori

The activity of the substance(s) which are contained in the cephalic endocrine organs of the locust which induce egg diapause in Bombyx mori was examined by implantation and injection of saline extracts of these organs. Extracts from the median and lateral neurosecretory parts of the locust brain were not effective in inducing egg diapause. Extracts of the corpora cardiaca, corpora allata, and suboesophageal ganglion of the locust induced diapause eggs in Bombyx pharate adults from which the suboesophageal ganglion had been removed. The first two extracts could induce egg diapause even in isolated abdomens of pharate adults of Bombyx. In the locust corpora cardiaca, the activity was present only in the glandular lobe and not in the nervous region. This activity decreased when the nervi corporis cardiaci I and II and of nervi corporis allati I were cut. Allatectomy also brought about a decrease in the activity in the glandular lobe which could not be restored by the injection of juvenile hormone. The activity in the corpora allata was enhanced slightly by the disconnection though not significantly.From these results, it is assumed that the corpora cardiaca, corpora allata and suboesophageal ganglion of the locust contain and active principle(s) capable of inducing egg diapause in Bombyx mori. The nervous connections between the brain, corpora cardiaca, and corpora allata are essential for the accumulation of the active substance(s) in the glandular lobes of the corpora cardiaca.