The fine structure of the midgut epithelium in a centipede,Scolopendra cingulata (Chilopoda,Scolopendridae), with the special emphasis on epithelial regeneration

Scolopendra cingulata has a tube-shaped digestive system that is divided into three distinct regions: fore-, mid- and hindgut. The midgut is lined with a pseudostratified columnar epithelium which is composed of digestive, secretory and regenerative cells. Hemocytes also appear between the digestive cells of the midgut epithelium. The ultrastructure of three types of epithelial cells and hemocytes of the midgut has been described with the special emphasis on the role of regenerative cells in the protection of midgut epithelium. The process of midgut epithelium regeneration proceeds due to the ability of regenerative cells to proliferate and differentiate according to a circadian rhythm. The regenerative cells serve as unipotent stem cells that divide in an asymmetric manner.Additionally, two types of hemocytes have been distinguished among midgut epithelial cells. They enter the midgut epithelium from the body cavity. Because of the fact that numerous microorganisms occur in the cytoplasm of midgut epithelial cells, we discuss the role of hemocytes in elimination of pathogens from the midgut epithelium. The studies were conducted with the use of transmission electron microscope and immunofluorescent methods.     

The evolutionary history of the Mediterranean centipede Scolopendra cingulata (Latreille, 1829) (Chilopoda: Scolopendridae) across the Aegean archipelago

In this study we investigate the evolutionary relationships of Scolopendra cingulata (Latreille, 1829) within insular Greece. Our main goal is to infer the time frame of the differentiation of the species in the study area. In this regard, sequence data originating from three mitochondrial genes are used to reconstruct the evolutionary history of 47 insular populations of S. cingulata from the Aegean archipelago. Within the phylogenetic framework and by implementing a relaxed molecular clock methodology, we infer the time estimates of separations of the S. cingulata lineages. The results of the phylogenetic analysis support the presence of three distinct S. cingulata groups in the region. The first group accommodates populations from the eastern Aegean islands, and is closely related to the second group that hosts mainly populations of northern and central Cyclades. The third group is composed of insular populations originating from southern Cyclades. Different temporal splitting scenarios have been evaluated. Based on the scenario strongly supported by the data, we propose a biogeographical scenario that could account for the contemporary distribution of the species' lineages. The splitting events of S. cingulata are estimated to have occurred within the late Miocene. The historical events of the last 13.77 Myr have shaped, through a series of mostly vicariant and dispersal incidents, the present‐day biogeographical pattern of the species. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 105 , 507–521.     

Venom apparatus and toxicity of the centipede Ethmostigmus rubripes (Chilopoda, Scolopendridae)

The venom apparatus of Ethmostigmus rubripes, a generalized predator, consists of the telopodites of the postcephalic segment, the basal article of w which contains the venom gland. Within the gland, venom granules are concentrated in intracellular secretory granules, from which they are discharged into vacuoles in the cytoplasm of the secretory cells and thereafter by exocytosis into the lumen of the gland. A venom duct carries venom to the venom claw, which introduces it into prey via a subterminal pore on the outer curvature of the claw. Pits containing pegs, presumed to be sensory, are concentrated near grooves leading to a cutting ridge proximal to the point of the claw. The venom is toxic both to mammals and insects.     

Effect of urbanization on centipede (Chilopoda) diversity in the Wielkopolska-Kujawy Lowlands of western Poland

Over fifty years of research data from various sources were compiled and analyzed in order to determine the effect of urbanization on centipede diversity in the Wielkopolska-Kujawy Lowlands of western Poland. Urbanization has had a significant effect on species composition and community structures. However, it has not reduced overall species richness and diversity in the centipede fauna. The centipede fauna from built-up areas was found to be rich and varied. The habitats with the highest levels of species richness were the city of Poznań and the beech forests in the surrounding region.     

Venom regeneration in the centipede Scolopendra polymorpha: evidence for asynchronous venom component synthesis

Venom regeneration comprises a vital process in animals that rely on venom for prey capture and defense. Venom regeneration in scolopendromorph centipedes likely influences their ability to subdue prey and defend themselves, and may influence the quantity and quality of venom extracted by researchers investigating the venom's biochemistry. We investigated venom volume and total protein regeneration during the 14-day period subsequent to venom extraction in the North American centipede Scolopendra polymorpha. We further tested the hypothesis that venom protein components, separated by reversed-phase fast protein liquid chromatography (RP-FPLC), undergo asynchronous (non-parallel) synthesis. During the first 48 h, volume and protein mass increased linearly. Protein regeneration lagged behind volume regeneration, with 65–86% of venom volume and 29–47% of protein mass regenerated during the first 2 days. No additional regeneration occurred over the subsequent 12 days, and neither volume nor protein mass reached initial levels 7 months later (93% and 76%, respectively). Centipede body length was negatively associated with rate of venom regeneration. Analysis of chromatograms of individual venom samples revealed that 5 of 10 chromatographic regions and 12 of 28 peaks demonstrated changes in percent of total peak area (i.e., percent of total protein) among milking intervals, indicating that venom proteins are regenerated asynchronously. Moreover, specimens from Arizona and California differed in relative amounts of some venom components. The considerable regeneration of venom occurring within the first 48 h, despite the reduced protein content, suggests that predatory and defensive capacities are minimally constrained by the timing of venom replacement.     

Hemocytes of the centipede Scutigera coleoptrata (Chilopoda,Notostigmophora) with notes on their interactions with the tracheae

The hemocytes of Scutigera coleoptrata were investigated by light and electron microscopy. Four types of hemocytes were identified: prohemocytes, plasmatocytes, granulocytes, and spherulocytes. Only granulocytes could be distinguished from the three other types by May-Grünwald staining, as this is the only hemocyte type demonstrating an eosinophilic reaction. Shape and size give further indications for distinguishing the cell types. In addition, differentiation is possible on the basis of their ultrastructure. However, only a combination of all three methods (staining and light and electron microscopy) allows clear separation of the cell types. As transitional stages between the cell types occur in S. coleoptrata, it is likely that prohemocytes, plasmatocytes, and granulocytes are ontogenetic stages of a single cell lineage. Special cell components and their possible functions are described. Plasmatocytes exocytose tubular structures that probably play a role in coagulation processes. These tubular structures develop in the grana of plasmatocytes. Also, a special arrangement of microtubules and microfilaments was demonstrated. For the first time interactions between hemocytes and tracheae are documented within the Chilopoda. It is assumed that the hemocytes meet their oxygen requirements directly from the tracheae. Phylogenetic implications of the results are discussed.     

Centipede (Scolopendra subspinipes mutilans) venom toxin peptide exhibits cytotoxic and cell growth effects in a concentration-dependent manner

Centipede venom contains a variety of proteins, peptides, and enzymes. However, the biological actions of toxin peptides in centipede venom remain largely unknown. In this study, we identified a centipede (Scolopendra subspinipes mutilans) venom toxin peptide (SsmTP) that was shown to act as a cell growth factor at low concentrations-in vitro. SsmTP was found to consist of 66 amino acids that display seven cysteine residues, which exhibited high similarity to the predicted neurotoxin. SsmTP was expressed in the venom gland of S. s. mutilans. A recombinant SsmTP peptide of approximately 5.2 kDa was produced in baculovirus-infected insect cells. Interestingly, SsmTP exhibited cytotoxicity in murine cells in a concentration-dependent manner but displayed a cell growth effect at low concentrations. SsmTP at a low concentration also protected murine cells against oxidative damage through the inhibition of caspase-1 and apoptosis. Our data indicate that SsmTP acts as a cell growth factor and a toxin peptide in a concentration-dependent manner. Consequently, our results provide evidence that the identification of SsmTP, a centipede toxin peptide, may not only elucidate the biological action of toxin peptides but also offer additional insight into the pharmacological applications of centipede venoms.     

A redescription of the large geophilid centipede Chilenophilus corralinus (Attems, 1903) (Myriapoda: Chilopoda: Geophilomorpha), with new distribution records from southern Chile

The little known geophilomorph centipede Chilenophilus corralinus (Attems, 1903) (Myriapoda: Chilopoda, Geophilidae), a large geophilid species from South America is herein redescribed and illustrated based on new specimens collected in the Andes of Southern Chile. New morphological features of specific value are also given for the taxon. Chilenophilus corralinus is reported for the first time from the following Chilean localities: Region XIV (de Los Ríos region): Valdivia province: Pirihueico; camping “La Herradura”; Mafil. Region X (de Los Lagos region): Chiloé province: Chiloé Island. Llanquihue province: Parque Nacional Alerce Andino. Palena province: Hualaihué; Chaitén; Palena; Futaleufú.     

Prey orientation and the role of venom availability in the predatory behaviour of the centipede Scolopendra subspinipes mutilans (Arthropoda: Chilopoda)

Many animal phyla contain clades in which most or all species are venom-injecting predators. An example, in the arthropods, is the class Chilopoda, containing the approximately 3500 species of centipedes. Very little ecological or behavioural work yielding quantitative data has been conducted on centipede predation. Here, we describe a study of this kind. Our experiments employed one centipede species - a large tropical one, Scolopendra subspinipes mutilans - and two species of prey - a cricket, Gryllus assimilis, and a locust, Schistocerca gregaria. We conducted two experiments. The first was aimed at investigating the extent to which the centipedes attacked prey in particular tagmata as opposed to at random over the whole body surface. The results showed that the centipedes were highly selective, preferring to attack the head or thorax rather than the abdomen; indeed, they often reoriented the prey in order to achieve this. A possible explanation of this behaviour is to maximize the speed with which the neurotoxins in the venom reach either the brain or the thoracic ganglia that control limb movement. The second experiment was aimed at investigating the effect of venom-extraction on the attack rate, and specifically at testing if the magnitude of any such effect differed between the two types of prey, which differ considerably in size. The results showed a major effect of venom extraction in relation to both types of prey, but with the time taken to return to a 'normal' attack rate being longer in the case of the larger prey-type, namely the locust. We discuss these results in relation to the 'venom optimization hypothesis' and, more generally, to the principle of minimizing the production/use of venom, which is an energetically expensive resource.     

A common terminology for the external anatomy of centipedes (Chilopoda)

A common terminology for the external morphological characters of centipedes (Chilopoda) is proposed. Terms are selected from the alternatives used in the English literature, preferring those most frequently used or those that have been introduced explicitly. A total of 330 terms are defined and illustrated, and another ca. 500 alternatives are listed.     

Die Ultrastruktur der Photorezeptoren von Lithobius forficatus L. (Chilopoda: Lithobiidae)

Zusammenfassung Die Untersuchung der feinstrukturellen Organisation der Ocellen von Lithobius forficatus L. ergab, daß der dioptrische Apparat aus einer lamellenartig gebauten, ungleichseitig bikonvexen Cornealinse besteht. Ein Glaskörper und spezielle Pigmentzellen fehlen. Der Augenbecher wird von 35–110 Sinneszellen gebildet, unter denen 2 morphologisch distinkte Typen unterschieden werden können. Die großen Sehzellen des distalen Bereiches besitzen einseitig inserierende Rhabdomere, die in radiärsymmetrischer Anordnung ein umfangreiches geschlossenes Rhabdom bilden. Der proximale Teil des Augenbechers wird von kleineren, konischen Basalzellen in Form einer undeutlich abgesetzten Retinula eingenommen. Durch enge Verzahnung ihrer zirkumapikal oder zweiseitig angeordneten Mikrovilli entstehen stelzenförmige Doppel- und Mehrfachrhabdomere, die mit dem zentralen Rhabdom in Verbindung stehen. Alle Sehzellen sind durch eine Gliederung in verschiedene Zonen gekennzeichnet. Sie sind bei distalen Rezeptoren senkrecht zur optischen Achse, bei Basalzellen transversal zur Längsachse der Zelle angeordnet. Auf die rhabdomerischen Mikrovilli des Augenzentrums folgt nach außen eine Schaltzone aus Elementen des ER und anderen vesilukären Bildungen. Diese Schaltzone stellt wahrscheinlich eine mit dem Adaptationszustand des Auges korrelierte Funktionsstruktur dar. In der cytoplasmatischen Zone fällt die Zahl verschiedenartiger multivesikulärer Korpuskel neben wenigen großen multilamellären Körpern auf. Die funktionelle Bedeutung des Ocellusaufbaus bei Lithobius wird diskutiert.

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Ultrastructure of the Photoreceptors of Lithobius forficatus L. (Chilopoda: Lithobiidae)

Summary The ultrastructure of the ocelli of Lithobius forficatus L. was investigated by means of conventional electron microscopy. The dioptric apparatus consists of an unequal biconvex corneal lens which has a lamella-like fine structure. Crystalline cones and special pigment cells are lacking. The eye cup is composed of 35 to 110 sense cells of two different morphological types. The large visual cells of the distal region are characterized by unilaterally inserted rhabdomeres which form in a radial symmetrical arrangement the extended closed rhabdome. The proximal part of the eye cup is occupied by somewhat smaller basal cells of conical shape, showing an indistinct retinula. These cells have numerous microvilli either in the apex region or laterally which interdigitate to form stiltlike double or multiple rhabdomeres in close communication to the central rhabdome. In both types of sense cells particular zones appear because of the characteristic distribution of certain cell elements. They are arranged perpendicularly to the optical axis in the large receptors and vertical in the main axis of the basal cells. Elements of ER and other vesicles constitute a Schaltzone which borders the microvilli of the rhabdomer. This Schaltzone probably is a functional structure correlated to the adaptional state of the eye. The cytoplasmatic zone contains numerous different multivesicular and few large multilamellar bodies. The functional meaning of the organization of the Lithobius ocellus is discussed.

Mit dankenswerter Unterstützung durch die Deutsche Forschungsgemeinschaft.     

The peristomatic structures of Lithobiomorpha (Myriapoda, Chilopoda): comparative morphology and phylogenetic significance

A comparative survey of the epipharynx and hypopharynx of lithobiomorph centipedes by light and scanning electron microscopy examines 18 species that sample the major groups of both families, the Lithobiidae and Henicopidae. Cladistic analysis of 11 characters of the peristomatic structures together with 29 additional morphological characters serves as a basis for interpreting the evolution of the lithobiomorph peristomatic structures. Scutigeromorpha is used for outgroup comparison in the framework of a homology scheme for the basic components of the epi- and hypopharynx. Compared to other chilopods, the monophyly of Lithobiomorpha is supported by a row of distinctive bottle-shaped gland openings at the border between the labral and clypeal parts of the epipharynx, as well as by a distinctive shape of the hypopharynx. Paired rows of elongate spines on the clypeal part of the epipharynx are an apomorphic character of Lithobiidae. The transformation of these spine rows into a few groups of branching spines is characteristic for the Monotarsobius group sensu Verhoeff. Similar groups of branching clypeal spines characterize the Anopsobiinae within Henicopidae, whereas Henicopinae possess a dense cluster of short, simple spines instead. The recently described genus Dzhungaria is resolved closer to Henicopinae than to Anopsobiinae, a hypothesis supported by a field of grooves on the medial labral part of the epipharynx. Monophyly of Henicopidae does not receive unique support from the peristomatic structures although two homoplastic characters contribute to this node; among these, the reduction of a median spine field between clypeal and labral parts of the epipharynx to a narrow transverse band also supports a close relationship between the Ezembius group and Hessebius within Lithobiidae. An Ezembius+Hessebius clade is additionally supported by the absence of a transverse bulge between the clypeal and labral parts of the epipharynx, a character otherwise present in all lithobiomorph species studied so far. Lithobius is resolved as polyphyletic, with different species being most closely related to such genera as Australobius, Hessebius and Pleurolithobius.     

Morphometric and meristic diversity of the species Scolopendra cingulata Latreille, 1829 (Chilopoda: Scolopendridae) in the Mediterranean region

Scolopendra cingulata Latreille, 1829 is the commonest scolopendromorph species in the Mediterranean area. However, its morphological and geographic variation has not been examined so far, and therefore robust hypotheses about the factors that have shaped that variability are lacking. We examined, using multivariate methods for 19 morphometric and meristic characters, the morphological variation of that species using a comprehensive sample of 503 adult specimens from 130 restricted geographic localities. The localities were distributed in three major geographic areas (Balkans, Asia Minor-Middle East, and Italy) in order to discern biological entities and to estimate the morphological relationships between populations and geographic regions. Results showed significant differences between the three geographic groups. Characters such as the distance between the paramedian sulci of the 7th and 8th tergite, the number of antennal segments, the number of spines on the dorsal side of 21st prefemur and number of spines on the 21st prefemoral process significantly discriminated populations of S. cingulata along a west-east geographic gradient. Both eastern (Balkan) and easternmost populations (Asia Minor-Middle East) showed higher mean values of antennal articles and spines of 21st prefemur than the Italian populations. No significant morphological variation was discovered between the sexes of S. cingulata apart from cephalic width and distance between the two paramedian sulci of the 7th and 8th tergite. Based on certain morphometric differences among the three major geographic regions, we suggest that S. cingulata originated in the east and colonized south European area from two different geographic routes. It is suggested that west-east morphological trend of S. cingulata is related either to the prominent palaeogeographic events of the area or to the competition with S. oraniensis Lucas, 1846 in the western Mediterranean.     

The current knowledge on centipedes (Chilopoda) in Slovenia: faunistic and ecological records from a national database

In spite of Slovenia’s very high biodiversity, it has only a few animal groups that have been significantly investigated and are well known in this area. Slovenian researchers have studied only about half of the species known to be living in the country (Mršić 1997), but among well investigated species are centipedes. All available data about centipedes in Slovenia collected from 1921 to 2014 have been consolidated and constitute a general electronic database called “CHILOBIO”, which was created to provide an easy overview of the Slovenian centipede fauna and to allow entry and interpretation of new data collected in future research. The level of investigation has been studied with this database, in conjunction with a geographic information system (GIS). In the study period, 109 species were identified from 350 localities in 109 of the 236 UTM 10 × 10 km quadrants which cover the study area. The south-central part of the country has been the subject of the best investigations, whereas there is an absence of data from the south-eastern, eastern and north-eastern regions The highest number of species (52) has been recorded near the Iška valley (Central Slovenia, quadrant VL68). In 48% of the UTM quadrants investigated fewer than 10 species were recorded and just 5 species were found in one locality. Seventeen species were reported only in the Dinaric region, 4 in the Prealpine-subpannonian region and 7 in the Primorska-submediterranean region.     

The importance of looking at small-scale patterns when inferring Gondwanan biogeography: a case study of the centipede Paralamyctes (Chilopoda, Lithobiomorpha, Henicopidae)

Gondwanan biogeography has fascinated zoologists and botanists for over a century, but most biogeographical work has used continent-scale areas as analytical units. More finely resolved patterns, as can be obtained from small invertebrates with limited dispersal abilities, will be obscured in those studies. A common case is treating Australia as a single biogeographical region. In the present study, the necessity of splitting Australia into multiple microareas is demonstrated using centipedes as an example. The lithobiomorph centipede Paralamyctes is distributed on fragments of Gondwana, with species in southern Africa, Madagascar, southern India, Patagonia, eastern Australia, and New Zealand. A cladogram for Paralamyctes is based on morphology and sequences for four molecular markers for 30 terminals that sample 20 of 26 known ingroup species and four outgroups. Analysis with direct optimization across a range of indel costs and transversion : transition cost ratios identifies two main clades: Paralamyctes ( Paralamyctes ) unites species from southern Africa, Madagascar, tropical and warm temperate Australia, and New Zealand. The other group includes the temperate Australian/New Zealand Paralamyctes ( Haasiella ) and Paralamyctes ( Thingathinga ) and a Chilean clade. Subtree analysis finds that different parts of Australia have closest affinities to other Gondwanan fragments, and some of these relationships (such as that between north Queensland and New Zealand) are based on taxonomically stable clades. Area delimitation for large continental fragments should use sufficiently fine resolution to test the 'monophyly' of those fragments and attempt to eliminate spurious geographical paralogy.  © 2006 The Linnean Society of London, Biological Journal of the Linnean Society , 2006, 89 , 65–78.     

On the function of the ultimate legs of some Scolopendridae (Chilopoda,Scolopendromorpha)

The function of the variously shaped ultimate legs of Scolopendridae is briefly reviewed. Their function in Scolopendra heros Girard, 1853, Scolopendra subspinipes Leach, 1815, Scolopendra morsitans (Linnaeus, 1758), Scolopendra galapagoensis Bollman, 1889, Scolopendra hainanum Kronmüller, 2012, Scolopendra spinosissima Kraepelin, 1903 Cormocephalus aurantiipes (Newport, 1844) and Ethmostigmus trigonopodus (Leach, 1817), in which they are least specialised has been investigated. Specimens were tapped with forceps on different parts of the trunk to simulate the attack of a predator. When tapped on the first third of the trunk (near the head), the centipedes attacked the forceps with their forcipules. When tapped on the last third or the ultimate legs, they adopted a warning position, raising the ultimate legs to display the ventral and medial prefemoral spines as well as the spined coxopleural processes. In some cases the centipedes attacked the forceps with the claws of the ultimate legs by chopping down on them after lifting the legs high into the warning position. When tapped in the mid part of the trunk, the centipedes curled sideways to reach the forceps with their forcipules and ultimate legs simultaneously. Scolopendra galapagoensis not only lifted the ultimate legs into the warning position but also the last 3-4 pairs of locomotory legs, presenting their distodorsal prefemoral spines. This resembles the warning posture of some spiders. In addition to their function in warning behaviour, defensive stabbing, ritualised meeting reactions and during courtship behaviour, the ultimate legs may in addition act as hooks and perhaps be involved in species recognition. No evidence was found that the ultimate legs are used to catch prey, nor of prey or predators being held between the prefemora.     

An exotic species spreads through Europe: Tygarrup javanicus (Chilopoda: Geophilomorpha: Mecistocephalidae) is reported from the Slovakia and the Czech Republic

Geophilomorph centipedes, due to their soil life style, are frequently transported with plants. This report tries to underline some adaptations that facilitate them being introduced and becoming alien species. Tygarrup javanicus (Attems, 1907), tropical species reported recently from different greenhouses in Central Europe, is used as an example of such a spreading species.     

Distribution and movements of Buller's albatross (Diomedea bulleri) in Australasian seas

Abstract

The distribution and movements of Buller's albatross in Australasian seas are analysed using results of shipborne surveys (13 238 10‐min counts), counts from trawlers, banding data, recoveries on beaches and fishing vessels, and records from the literature. Patterns of marine distribution are documented by monthly accounts and maps. During the breeding season, highest abundances are recorded over shelves and slopes off southern New Zealand (The Snares shelf to 41–43°S off the South Island, D. b. bulleri), around the Chatham Islands and over oceanic subtropical waters east of New Zealand (probably D. b. platei), with marked seasonal variations observed off southern New Zealand. Both subspecies disperse mostly outside Australasian waters during the non‐breeding season. Birds banded on The Snares were recovered off south‐eastern New Zealand (Stewart Island to Cook Strait) and in the eastern tropical Pacific. Immatures accounted for only 0.25% of birds censused during the ship‐borne surveys; they are recorded around the New Zealand mainland in August‐October and February‐May, off south‐eastern Australia and in the Tasman Sea in November‐December, February, and June‐July. Around New Zealand, males predominate among birds recovered along the eastern seaboard, whereas the sex ratio in south‐western waters tends to vary according to water depth and season. Distribution patterns and movements in New Zealand and Australian seas are discussed in relation to breeding events and breeding status.