The chemosensory behaviour of Lithobius forficatus. 1. Evidence for a pheromone released by the coxal organs (Myriapoda: Chilopoda)

The behaviour of the Common brown centipede, Lithobius forficatus (.L.) is investigated with reference to the function of the coxal organs which are a peculiar and characteristic feature of Chilopoda. Previous studies of these organs in centipedes have been largely morphological and have relied on ultrastructural comparisons with other arthropod epithelia of disparate functions. Evidence is presented which suggests that the coxal pores release a sex-specific pheromone. The nature of intraspecific interactions in L. forficatus is discussed.     

Coxal organs in geophilomorpha (Chilopoda). Organization and fine structure of the transporting epithelium

Summary The coxal organs of different Geophilomorpha were studied by scanning and by transmission electron microscopy.1) The coxae of the last trunk-segment contain pores in different arrangements and numbers. They are the openings of the coxal organs.2) The coxal organs are formed by four different cell types: the main epithelium consists of radially arranged transporting cells, surrounded by junctional cells, gland cells, and the cells of the pore channel.3) The cells of the transporting epithelium show an enlargement of the apical and basal surface. Deep and narrow extracellular channels of the apical infoldings are closely associated by mitochondria (plasmalemma-mitochondrial complexes). The epithelium is covered by a prominent cuticle with a spacious subcuticle.4) A distinct mucous layer covers the cuticle of the transporting epithelia, and is secreted by the gland cells.5) A small cellular sheath separates the epithelium of the coxal organ against the haemolymph.6) The possible function of the coxal organs in ion and fluid transport is discussed.     

The water relations of Lithobius forficatus and the role of the coxal organs (Myriapoda: Chilopoda)

The water relations of the common brown centipede, Lithobius forficatus (L.), are examined with regard to the possible involvement of the coxal organs in water uptake. Centipedes with blocked and open coxal pores are subjected to a variety of dehydration and rehydration regimes and weight change over time measured. Weight loss and regain is related to body surface area and is not influenced by the coxal organs.
The conclusion of the paper is that the coxal organs of L. forficatus do not contribute significantly to the ability of these animals to recover water lost through cuticular transpiration.     

The chemosensory behaviour of Lithobius forficatus (Myriapoda: Chilopoda). 2. Bioassay and chemistry of the coxal pheromone

The common brown centipede, Lithobius jorJcatus (L.), releases a sex-specific pheromone which emanates from distinctive cuticular openings on the coxae of the hind legs-the coxal pores (Littlewood & Blower, 1987). The pheromone can be isolated from tissue extracts of the coxal organs (surrounding the coxal pores) and from substrata which have been 'marked by male or female L. forjcatus and presented to centipedes in the form of a behavioural bio-assay.
In this paper, further evidence is presented for a centipede pheromone, the chemical components of male and female secretions are separated using Gas Layer Chromatography (GLC), a qualitative chemical analysis is described and the nature of the pheromone is discussed.     

The coxal glands of geophilomorpha (chilopoda): Organs of osmoregulation

Summary The organs terminating at the coxal pores of the tug-legs of Geophilomorpha are not repugnatorial glands, but possess typical transport epithelia with deep apical and basal infoldings of the cell membranes, between which numerous large mitochondria are located. Many transport vesicles are found in the basal region but fewer in the apical cytoplasm. The apex is characterized by bundles of longitudinally oriented microtubules, sparse endoplasmic reticulum and free ribosomes. Single neurosecretory axons with synaptoid areas are scattered among the cells. It is suggested that the coxal organs have a diuretic function in moist habitats and an antidiuretic effect in arid environments. The switch-over is evidently controlled by a neuroendocrine mechanism.     

The contributions of diverse sense organs to the control of leg movement by a walking insect

Summary During locomotion, stick insectsCarausius morosus, place the tarsus of the rear leg near the tarsus of the ipsilateral middle leg, whatever the position of the latter. This adjustment by the hind leg requires that it receive information on the actual position of the middle leg tarsus. It is shown by ablation experiments that such information is contributed by the following proprioceptors of the middle leg: the ventral and dorsal coxal hairplates, the coxal hair rows, the trochanteral hairplate and the femoral chordotonal organ. Additional information comes from other, as yet unidentified, sense organs. Several alternatives are considered to explain how the signals from the diverse sense organs of the subcoxal joint might be combined in computing the target position for the protracting hind leg. The experimental results support the hypothesis that the signals are added nonlinearly and that a signal deviating from the majority pattern is weighted less.Abbreviations cxHPu ventral coxal hairplate - cxHPd dorsal coxal hairplate - trHP trochanteral hairplate - HR hair row - feCO femoral chordotonal organ - AEP anterior extreme position     

Ultrastructural organization of the anal organs in the anal capsule of Craterostigmus tasmanianus Pocock, 1902 (Chilopoda, Craterostigmomorpha)

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.     

The ultrastructure of the sternal gills forming a striking contrast with the coxal gills in a fresh-water amphipod (Crustacea)

Sternomoera yezoensis has specialized sterna with 21 sternal gills in addition to six pairs of coxal gills. Despite a common high permeability to chloride ions, the cpithelia of these two kinds of gills are diametrically opposed in the polarity of the cell membranemitochondria complex. The coxal gill epithelium (4-6 mum thick) is characterized by a welldeveloped AIS (apical infolding system) associated with a huge number of large mitochondria. The AIS exceeds two-thirds of the epithelial thickness and forms a highly sophisticated, subcuticular labyrinth. On the contrary, the sternal gill epithelium, an extension of the sternal epithelium proper, is extremely thick (10-15 mum) and is characterized by a very deep BIS (basolateral infolding system) associated with numerous slender mitochondria. The BIS reaches nine-tenths of the epithelial thickness and forms a giant, baso-lateral labyrinth. Shallower, less elaborate AIS and BIS without mitochondrial association originate from the opposite sides of these epithelia. Although AIS and BIS interpenetrate in the sternal gill epithelium, they never communicate. The results indicate that in addition to the coxal gills, the sterna with Ihe sternal gills function as transporting as well as respiratory organs, though the functional difference between these two kinds of gills remains to be elucidated.     

Coxal organs of Lithobius forficatus (Myriapoda,Chilopoda)

Summary In Lithobius forficatus each of the coxae of the four posterior trunk segments bear a pore field with several coxal pores. The surrounding single-layered epithelium is composed of four different cell types: the main epithelial cells having a fine-structural organization of transport cells with deep apical and basal folds of the cell surfaces and plasmalemma-mitochondrial complexes, junctional cells, exocrine glands, and the wall cells of the pore channel. The entire epithelium is separated from the hemolymph by an inner cellular sheath. It is assumed that the coxal organs participate in fluid uptake.     

Comparative morphology,functions, and homologies of the coxal glands in oribatid mites (Arachnida: Acari)

1. Forty-eight species of oribatids in 37 families representing most of the superfamilies were collected from various environments (littoral, salt marsh, litter, sod, and freshwater) and sectioned. 2. The coxal gland is composed of a sacculus and a labyrinth in all stages of all oribatid species. Muscles, originating on the body wall, insert at several points on the thin-walled sacculus which opens into the labyrinth. The labyrinth has an internal, chitinous supporting skeleton. The type A labyrinth has 3–180° bends, producing four parallel regions, and occurs in all inferior oribatids. The type B labyrinth has 1–180° bend, producing two parallel regions, and occurs in all superior oribatids. The coxal gland duct and the lateral gland duct join, penetrate the body wall, and empty into the posterior end of the podocephalic canal. All oribatids have lateral accessory glands, but only inferior oribatids have rostral and medial glands. Three ductless coxendral bodies are always present. 3. The labyrinth length in oribatids is correlated with body size and the environment of the species. Oribatids from sod, leaflitter, or moss show a simple correlation of labyrinth length (X) to total body length (Y) where Y = 4.64X. Freshwater species have a labyrinth length greater than that of comparably sized terrestrial species and salt water (littoral) species have a labyrinth length less than that of comparably sized terrestrial species. There is a greater reduction in labyrinth length in species restricted to salt marshes than in species not restricted to salt marshes. 4. The probable function of oribatid coxal glands is osmoregulation. Hemolymph filtration would occur across the sacculus by positive hemolymph pressure and contraction of the sacculus muscles. Resorption of ions would occur in the labyrinth, which is noncollapsible due to the internal skeleton. The hypothesis is that in freshwater species the rate of filtration is high and resorption of ions would have to be very efficient, therefore they have an elongated labyrinth; but in salt water species water loss must be minimized and preservation of ions would be a disadvantage, therefore they have a shortened labyrinth. Excre ion may also be a function of the coxal glands. The lateral gland may possibly function as an endocrine gland involved with production of a molting hormone. The rostral glands in inferior oribatids may have a salivary function. 5. The coxal glands of Peripatus, some millipedes, apterygote insects, decapod crustaceans, and all arachnid orders are homologous. The Tetrastigmata, Notostigmata, Cryptostigmata, and soft ticks have typical arachnid coxal glands. The coxal glands of higher Prostigmata may be modified into salivary, silk, or venom glands. The coxal glands in Mesostigmata, Astigmata, and hard ticks are lacking or highly modified.     

The proprioceptive function of a complex chordotonal organ associated with the mesothoracic coxa in locusts

Summary A system of chordotonal organs in the locust mesothorax consists of four subunits one of which connects to the coxa. Proprioceptive afferents from the scoloparia record the rotatory movements of the coxa. Mechanical stimulation of the sensory system by sinusoidal stretch or movements mimicking stretch as in natural walking of the locust elicits reflex activation of coxal motoneurones. Both assistance and resistance reflexes to imposed movements occur, but their intensity can vary from periods of suppression below firing threshold in a motoneurone to recruitment of additional motoneurones to the same muscle. It is concluded that some of these reflexes recorded in isolated preparations can also occur in freely walking animals where they should contribute to the muscular coordination of transitions between antagonistic movements.Abbreviations aCO, cCO, pCO, vCO anterior, coxal, posterior, ventral chordotonal organ - COS chordotonal organ system - pm-al postero-median to anterior-lateral     

Transport-active organs within the ’ano-genital’ capsule of Craterostigmus tasmanianus (Chilopoda, Craterostigmomorpha)

 In Craterostigmus tasmanianus, first results of the cellular organization of anal organs within the ’ano-genital’ capsule are presented. Each valve of the ’ano-genital’ capsule bears four pore fields ventrally, each of them consisting of several pore openings of the anal organs. The pores lead into a cuticle-lined pore channel, the base of which is surrounded by a single-layered epithelium that is composed of three different cell types. The main epithelium consists of radially arranged transport-active cells surrounded by exocrine cells, and the cells of the pore channel. The cells of the transporting epithelium show deep invaginations of the apical and basal cell surfaces and plasmalemma-mitochondrial complexes. These cells are covered by a specialized cuticle with a prominent subcuticle. Exocrine glands secrete a mucous layer on the cuticle of the main epithelium. The type of anal organ present in Craterostigmus tasmanianus shows similarities to coxal and anal organs found in other Pleurostigmophora in the chilopods. The possible function of the anal organs in uptaking water vapour is discussed. It is appropriate to call the organs within the ’ano-genital’ capsule of Craterostigmus tasmanianus ”anal organs”, as components of the genital segments are not involved. Accepted: 17 November 1996     

Fine structure and function of the coxal glands of lithobiomorph centipedes: Lithobius forficatus and L. crassipes (Chilopoda,Lithobiidae)

The ultrastructure of the coxal glands and associated tissues in the centipedes Lithobius forficatus and Lithobius crassipes has been examined in the light of two contrasting functional hypotheses postulated by different authors. Lithobiomorph chilopods possess eight sets of pores on the posterioventral border of the coxal podomeres of leg pairs 12–15 in adult (maturus) and subadult (pseudomaturus) stadia. A modified cuticular hypodermis, known as the coxal gland, surrounds the distal portion of each blindended pore. Each gland is made up of cells which contain large numbers of hypertrophied mitochondria and a highly folded apical and basal plasma membrane. The similarity of the coxal gland to so called “transporting epithelia” is discussed and further comparisons are made between these and secretory glands in arthropods. A careful consideration of both functional hypotheses (osmoregulation or pheromone release) has revealed the possibility that the coxal gland may encompass both functions.     

Fine structure and function of the prosomal glands of the two-spotted spider mite,Tetranychus urticae (Acari,Tetranychidae)

Summary The prosomal glands of Tetranychus urticae (Acari, Tetranychidae) were examined light and electron microscopically. Five paired and one unpaired gland are found both in females and males. The silk spinning apparatus consists of paired silk glands which extend laterally on both sides of the esophagus into the pedipalps. There, they enter the terminal silk gland bag which opens into a silk bristle at the apex of the pedipalps. The salivary secretions are formed in three paired glands which have an interconnecting duct, the podocephalic canal. The dorsal podocephalic glands may produce a serous secretion, the anterior podocephalic glands a mucous secretion, and the coxal organ may add a liquid, ion-rich secretion. These secretions pass the podocephalic canal and reach the mouth at the apex of the gnathosome. The function of the paired tracheal organs and the unpaired tracheal gland is still unclear. The tracheal gland may produce a secretion which facilitates the movement of the fused chelicerae and the stylets.This study was financed by a grant from the Deutsche Forschungsgemeinschaft (DFG Se 162/12)     

The peripheral and central nervous organization of the locust coxo–trochanteral joint

The micromorphology of the locust coxo–trochanteral joint was examined in cobalt-stained material. Peripheral nervous system, musculature, and internal proprioceptors—two strand receptors and a muscle receptor organ—of the metathoracic coxa are compared with those of the pro- and mesothoracic legs. The number and position of trochanter levator and depressor motoneurons as well as the central projections of coxal sense organs are described. Evidence for a femoro–tibial strand receptor was obtained by tracing the path of a particular nerve branch.     

Detecting taxonomic signal in an under-utilised character system: geometric morphometrics of the forcipular coxae of Scutigeromorpha (Chilopoda)

To date, the forcipules have played almost no role in determining the systematics of scutigeromorph centipedes though in his 1974 review of taxonomic characters Markus Würmli suggested some potentially informative variation might be found in these structures. Geometric morphometric analyses were used to evaluate Würmli's suggestion, specifically to determine whether the shape of the forcipular coxa contains information useful for diagnosing species. The geometry of the coxae of eight species from the genera Sphendononema, Scutigera, Dendrothereua, Thereuonema, Thereuopoda, Thereuopodina, Allothereua and Parascutigera was characterised using a combination of landmark- and semi-landmark-based sampling methods to summarize group-specific morphological variation. Canonical variates analysis of shape data characterizing the forcipular coxae indicates that these structures differ significantly between taxa at various systematic levels. Models calculated for the canonical variates space facilitate identification of the main shape differences between genera, including overall length/width, curvature of the external coxal margin, and the extent to which the coxofemoral condyle projects laterally. Jackknifed discriminant function analysis demonstrates that forcipular coxal training-set specimens were assigned to correct species in 61% of cases on average, the most accurate assignments being those of Parascutigera (Parascutigera guttata) and Thereuonema (Thereuonema microstoma). The geographically widespread species Thereuopoda longicornis, Sphendononema guildingii, Scutigera coleoptrata, and Dendrothereua linceci exhibit the least diagnostic coxae in our dataset. Thereuopoda longicornis populations sampled from different parts of East and Southeast Asia were significantly discriminated from each other, suggesting that, in this case, extensive synonymy may be obscuring diagnosable inter-species coxal shape differences.     

The ultrastructural investigation of the midgut in the quill mite Syringophilopsis fringilla (Acari,Trombidiformes: Syringophilidae)

The midgut of the females of Syringophilopsis fringilla (Fritsch) composed of anterior midgut and excretory organ (=posterior midgut) was investigated by means of light and transmission electron microscopy. The anterior midgut includes the ventriculus and two pairs of midgut caeca. These organs are lined by a similar epithelium except for the region adjacent to the coxal glands. Four cell subtypes were distinguished in the epithelium of the anterior midgut. All of them evidently represent physiological states of a single cell type. The digestive cells are most abundant. These cells are rich in rough endoplasmic reticulum and participate both in secretion and intracellular digestion. They form macropinocytotic vesicles in the apical region and a lot of secondary lysosomes in the central cytoplasm. After accumulating various residual bodies and spherites, the digestive cells transform into the excretory cells. The latter can be either extruded into the gut lumen or bud off their apical region and enter a new digestive cycle. The secretory cells were not found in all specimens examined. They are characterized by the presence of dense membrane-bounded granules, 2–4 μm in diameter, as well as by an extensive rough endoplasmic reticulum and Golgi bodies. The ventricular wall adjacent to the coxal glands demonstrates features of transporting epithelia. The cells are characterized by irregularly branched apical processes and a high concentration of mitochondria. The main function of the excretory organ (posterior midgut) is the elimination of nitrogenous waste. Formation of guanine-containing granules in the cytoplasm of the epithelial cells was shown to be associated with Golgi activity. The excretory granules are released into the gut lumen by means of eccrine or apocrine secretion. Evacuation of the fecal masses occurs periodically. Mitotic figures have been observed occasionally in the epithelial cells of the anterior midgut.     

Ultrastructural evidence for an osmoregulatory effect of nikkomycin in the mite Tetranychus urticae

Histological effects of the microbial metabolite and chitin synthesis inhibitor complex Nikkomycin (AMS 0896 Bayer Leverkusen) on osmoregulatory organs of all developmental stages of Tetranychus urticae are described. The metabolite, in a concentration of 100 ppm, was applied via the nutritive plant. Mites fed for 2 to 14 days, and then were collected and immediately fixed. Two osmoregulatory organs occur in T. urticae. The Malpighian complex, differentiated only in females, shows an increased number of apical microvilli in the epithelium of the distal regions after metabolite application, thus resulting in an enlargement of the surface area. Changes in the second osmoregulatory organ, the coxal organ, after Nikkomycin application include depositions of membranous bodies in the lumen as well as in cytoplasmic vacuoles of the proximal tubule. Additionally, an increase in the luminal diameter occurs. Numerous vacuoles of different contents are observed in the cytoplasm of the distal tubule. Consequences of histological alterations in osmoregulatory organs after Nikkomycin application are discussed with special reference to the composition of salivary secretions.     

山羊蠕形螨扫描电镜观察

本文通过扫描电镜对寄生虫山羊体的山羊蠕形螨生活史各期颚体,足体和末体的超微结构观察,发现以下超微结构;一个圆锥状须乳突位于雌螨触须第２节背面;叶状背阳茎侧突和腹阳茎侧突竖立于阳茎同一基部,未受精雌螨船形阴门的一对瓣汇成纵线,雌螨交配后阴门半开,产卵后全部敞开,同时描述该螨的背基刺,锥状突,口下板,螯肢,基内叶,口孔,须爪和肛道等的超微结构,文中还讨论了锥状突,口下板,螯肢,基内叶,须爪和须乳突的功     

The Neotropical Genus Notodiaptomus Kiefer, 1936 (Calanoida: Diaptomidae): Redescription of the Type Species Notodiaptomus deitersi (Poppe, 1891) and Designation of a Neotype

Ribautia williamsi sp. nov., a new dwarf geophilomorph centipede from the Lower Urubamba Region, Peruvian Amazonia, is described and illustrated based on the holotype female. The new species is characterized by having the coxal organs grouped in clusters (three of these in each coxopleuron of the ultimate leg-bearing segment) and ventral pore-fields present along all the body; these two combined traits being shared by five other Neotropical species currently included in the genus Ribautia Brölemann, 1909, i.e. R. centralis (Silvestri, 1907) (from Colombia and Brazil), R. difficilis Pereira, Minelli &; Barbieri, 1995 (from Brazil), R. montana Kraus, 1954 (from Peru), R. peruana (Verhoeff, 1941) (from Peru), and R. titicacae (Turk, 1955) (from Peru). The new taxon is differentiated from the aforementioned species by the low number of leg-bearing segments and small body length; it is included in a key which will enable the identification of all known Neotropical members having coxal organs grouped in clusters. R. williamsi sp. nov. is the 14th species of Ribautia recorded from Peru.

http://zoobank.org/urn:lsid:zoobank.org:pub:D9FA5FAF-7652-4A5B-AC09-8783F05A694D