Spiracle structure in scolopendromorph centipedes (Chilopoda: Scolopendromorpha) and its contribution to phylogenetics

The spiracles of scolopendromorph centipedes have long been a source of systematic characters based on their segmental distribution and gross morphology, but microscopic investigations to date have documented only a small number of species. A scanning electron microscopic survey of 34 species that samples the major groups of Scolopendromorpha reveals variability in such features as the structure of the peritremal margin, specific kinds of sensilla and glandular pores on the peritrema, projections on the valves that subdivide the atrium (in Scolopendrinae), and the form of the trichomes around the tracheal openings. Adding new characters from the spiracles to recent morphological datasets for phylogenetic inference reinforces the monophyly of major groups of Scolopendridae and is particularly informative for relationships within Scolopendrini. A bowl-like atrium with the tracheae opening between humps in its floor is more widespread in Scolopendromorpha than previously reported. Shared presence of spiracle muscles in Cryptopidae and Scolopendrinae may reflect convergent evolution of a subatrial cavity in these groups rather than being an apomorphic character for Scolopendromorpha as a whole.     

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.     

The sternal pore areas of geophilomorph centipedes (Chilopoda: Geophilomorpha)

Most geophilomorph centipedes have segmental clusters of exocrine glands whose opening pores are arranged in more or less well-defined sternal pore areas. We describe here the cuticular structures forming and/or accompanying the gland openings on the sternites and the shape of the pore areas along the body axis in representatives of most geophilomorph families. The cuticular ring around the pore may exhibit either of two forms. In Himantariidae ( Himantarium ) and in Dignathodontidae ( Henia ) the ring looks like a continuous ribbon with a visible suture, whereas in the representatives of the remaining families no suture is seen. As to the distribution of the pores on ventral surface of the body, we record the presence of pores on the last leg-bearing segment of Clinopodes flavidus , whereas that segment was described as poreless in all geophilomorphs. We also provide a taxonomic survey of shape and distribution of pore areas in the individual families, where the pore areas may take very different shapes that we regard as transformational homologues. As for the segmental distribution of sternal pore areas, there is a considerable amount of complexity along the trunk of geophilomorph centipedes, in contrast to the apparently uniform trunk structure.     

Morphology and diversity of the forcipules in Strigamia centipedes (Chilopoda,Geophilomorpha)

The morphology of the venomous limbs (forcipules) of 13 species of Strigamia and of six other geophilomorphs was studied with light microscopy, scanning electron microscopy, and, for a subsample, with confocal laser scanning microscopy. In all Strigamia species a well-distinct denticle is present invariantly on the inner side of the terminal article (tarsungulum), in sub-basal position, just proximal to a faint transverse sulcus and a cuticular introflexion that corresponds to the insertion point of a tendon. Strigamia species differ mainly in size and shape of the denticle and thickness of the distal part of the tarsungulum, suggesting some functional diversity in piercing and handling prey. Anatomical evidence supports the hypothesis that the tarsungulum corresponds to two ancestral articles and a denticle at the basis of the tarsungulum originated multiple times within geophilomorphs, however in different positions corresponding to either the ancestral sub-terminal article (in Strigamia, other Geophiloidea and some Schendylidae) or the ancestral terminal article (in the himantariid Thracophilus).     

Type specimens of centipedes (Myriapoda,Chilopoda) in the National Museum,Prague (Czech Republic)

The centipede collection in the National Museum in Prague contains type material of 16 taxa (14 species and two subspecies), of which 15 were described by Luděk J. Dobroruka and one by Karl W. Verhoeff: Allothereua wilsonae Dobroruka, 1979; Chinobius alenae Dobroruka, 1980; Lithobius corrigendus Dobroruka, 1988; Lithobius creticus Dobroruka, 1977; Lithobius erythrocephalus mohelensis Dobroruka, 1959; Lithobius evae Dobroruka, 1958; Lithobius magurensis Dobroruka, 1971; Lithobius purkynei Dobroruka, 1957; Lithobius tatricus Dobroruka, 1958; Lithobius tatricus monounguis Dobroruka, 1958; Monotarsobius homolaci Dobroruka, 1971; Monotarsobius krali Dobroruka, 1979; Pachymerium dilottiae Dobroruka, 1976; Pachymerium hanzaki Dobroruka, 1976; Scolopendra aztecorum Verhoeff, 1934 and Strigamia olympica Dobroruka, 1977. Of these 16 taxa, five were described from the Czech Republic, three from Slovakia and eight from other countries (Greece, Iraq, Kyrgyzstan, Mexico, Nepal, Russia and Uzbekistan). The eight taxa described from the Czech and Slovak Republics are now considered as junior synonyms but the eight taxa described from the other countries are still valid.     

Water and temperature relations, and diurnal rhythms of scolopendromorph centipedes

Water is lost by transpiration at relative humidities (R.H.) as high as 98% at 27° C and there is no evidence of the uptake of moisture through the integument from unsaturated air or moist surfaces. Water-loss through transpiration is directly related to saturation deficiency and there is no ‘critical’ transitional temperature at which the cuticle becomes porous. Nevertheless, water is retained more efficiently in Scolopendra than it is in Lithobius, suggesting an adaptation to hot-dry environments. The mean supercooling point of large specimens conditioned in an insectary is -3.1±0.48° and a single supercooling can be injurious or fatal. This indicates that cold winter weather must be avoided by burrowing deeply. Aktograph experiments show that these centipedes are nocturnal. The rhythm persists in darkness at constant temperature, but is quickly lost in constant light.

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Résumé Les Myriapodes étudiés montrent une perte d'eau par évaporation lorsqu'ils sont placés dans une ambiance à 27° et dont l'humidité relative est de 98%; il n'y a aucune indication permettant de penser que ces arthropodes peuvent se rehydrater à partir des surfaces mouillées ou de l'humidité atmosphérique, par prélèvement à travers leur tégument. La perte d'eau par transpiration est directement en rapport avec la déficience de saturation de l'air et il n'y a aucune température critique de transition pour laquelle la cuticule deviendrait perméable à l'eau. Néanmoins l'eau est retenue plus efficacement par Scolopendra que par Lithobius, ce qui suggère chez le premier genre une adaptation aux milieux chauds et secs. Le point de surfusion mesuré pour de grands spécimens maintenus en insectarium, est en moyenne de -3,1±0.48° et un seul choc à cette température de surfusion peut être préjudiciable ou même fatal. Cela indique que ces arthropodes doivent éviter les froids de l'hiver en creusant plus profondément les galeries. Des expériences utilisant un actographe montrent que ces Myriapodes ont une activité nocturne. Le rythme naturel d'activité persiste à l'obscurité et à une température constante, mais il est rapidement perdu par exposition à un éclairement continu.

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This work was carried out during tenure of a National Science Foundation Senior Visiting Scientist Fellowship, for which appreciative acknowledgment is hereby given.     

Impact of windstorm on a community of centipedes (Chilopoda) in a beech forest in Western Poland

The study was carried out in the years 2016–2017, five years after a windstorm which destroyed 1/3 of the protected beech forest area in the west of Poland. The community of centipedes in the area affected by the windstorm was depleted in terms of the species richness, diversity, and population density. The dominance structures were shortened and the species composition was rebuilt. The areas that proved to be the richest in terms of species richness and diversity among the sites affected by the windstorm were the one where windfallen trees were left and the other where beech trees had been planted by humans. In total, the quantitative and qualitative samples collected four times throughout a year featured 608 specimens from 11 species of two centipede orders – Lithobiomorpha and Geophilomorpha. Lithobius curtipes and L. forficatus were found in all of the investigated areas. L. pelidnus and L. piceus were captured at control sites exclusively. Only one species –L. erythrocephalus was found solely at the damaged site. The most numerous and most frequently found species in the community were L. curtipes, L. mutabilis, and Strigamia acuminata respectively. Although windstorms are natural phenomena their consequences may lead to significant changes in the community of the investigated soil animals. The importance of coarse woody debris, significantly contributing to the improvement and maintenance of species richness and diversity of Chilopoda, has once again been confirmed.     

Pincer-like claws in centipedes (Chilopoda): multiple evolutionary origin of similar form and serial pattern

In most Chilopoda, the walking legs end in a single-tip claw usually accompanied by short accessory spines. Instead, in all species of three small and only distantly related geophilomorph taxa (Diphyonyx, Neogeophilidae, Eucratonyx), the claws of an anterior set of leg pairs are unusually pincer-like. By integrating different microscopic techniques, including confocal laser scanning microscopy, we found that these modified claws are very similar in form, internal structure, and pattern of variation in shape along the trunk in all three taxa: the claws are distinctly swollen and bent, provided with peculiar bulges, and flanked by a conspicuous additional branch, either cylindrical or flattened, which overreaches the tip of the claw; instead, the internal cuticular features are not modified with respect to the condition in the other centipedes, claiming against the possibility of controlled abduction/adduction between claw and branch. Irrespective of the total number of leg pairs (63–129), the claws change gradually from pincer-like to usual shape invariantly in the range spanning between the 34 and the 45% of the total number of leg pairs. Despite these similarities, pincer-like claws originated independently in the three taxa, and by way of fundamentally different changes, either by the dramatic modification of the already existent anterior accessory spine (Diphyonyx, Neogeophilidae) or by the production of a novel cuticular projection (Eucratonyx). Moreover, their shared pattern of variation along the body was most probably constrained by already operating developmental processes controlling the longitudinal patterning of the trunk.     

Tentorial mobility in centipedes (Chilopoda) revisited: 3D reconstruction of the mandibulo-tentorial musculature of Geophilomorpha

Mandibular mechanisms in Geophilomorpha are revised based on three-dimensional reconstructions of the mandibulo-tentorial complex and its muscular equipment in Dicellophilus carniolensis (Placodesmata) and Hydroschendyla submarina (Adesmata). Tentorial structure compares closely in the two species and homologies can be proposed for the 14/17 muscles that attach to the tentorium. Both species retain homologues of muscles that in other Pleurostigmophora are traditionally thought to cause swinging movements of the tentorium that complement the mobility of the mandibles. Although the original set of tentorial muscles is simplified in Geophilomorpha, the arrangement of the preserved homologues conforms to a system of six degrees of freedom of movement, as in non-geophilomorph Pleurostigmophora. A simplification of the mandibular muscles is confirmed for Geophilomorpha, but our results reject absence of muscles that in other Pleurostigmophora primarily support see-saw movements of the mandibles. In the construction of the tentorium, paralabial sclerites seem to be involved in neither Placodesmata nor Adesmata, and we propose their loss in Geophilomorpha as a whole. Current insights on the tentorial skeleton and its musculature permit two alternative conclusions on their transformation in Geophilomorpha: either tentorial mobility is primarily maintained in both Placodesmata and Adesmata (contrary to Manton’s arguments for immobility), or the traditional assumption of the tentorium as being mobile is a misinterpretation for Pleurostigmophora as a whole.     

ChiloKey,an interactive identification tool for the geophilomorph centipedes of Europe (Chilopoda,Geophilomorpha)

ChiloKey is a matrix-based, interactive key to all 179 species of Geophilomorpha (Chilopoda) recorded from Europe, including species of uncertain identity and those whose morphology is known partially only. The key is intended to assist in identification of subadult and adult specimens, by means of microscopy and simple dissection techniques whenever necessary. The key is freely available through the web at: http://www.biologia.unipd.it/chilokey/ and at http://www.interactive-keys.eu/chilokey/.     

Numbers and biomass of centipedes (Lithobiomorpha: Chilopoda) in a Betula-Alnus woodland in N.E. England

Summary Over a period of 1 year the mean monthly population densities of L. forficatus varied between 7.58 m^-2 (February) and 24.07 m^-2 (October). The equivalent values for L. crassipes were 17.05 m^-2 (February) and 64.17 m^-2 (October). The annual mean population density of the two species was estimated at 51 m^-2 with a biomass of 0.585 g live wt m^-2. Less than 4% of the total lithobiid population was found in decaying logs and tree stumps, most of the population inhabited the soil/litter layers.Seasonal fluctuation in numbers was shown to be unimodal in L. forficatus and bimodal in L. crassipes. A similar pattern was noted in the status of the ovaries, L. forficatus appears to have a single peak breeding period (September) while L. crassipes has two (June and September).In terms of biomass, the relative importance of centipedes among other woodland invertebrate predators is high.     

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.     

Phylogenetic implications of endosperm cell wall morphology in Douglasia, Androsace, and Vitaliana (Primulaceae)

We investigated differences in endosperm cell wall thickening in three genera of Primulaceae; seven species of Douglasia , 28 species of Androsace representing the four sections of the genus, and the single species of the genus Vitaliana . Endosperm cell walls have irregular wall thickenings and narrow constrictions and characterize all Douglasia species and Vitaliana , whereas evenly thickened cell walls without constrictions were found in the endosperm of Androsace and Primulu . The endosperm character supports the monophyly of Douglasia and Vitaliana , and we discuss the phylogenetic implications of this conclusion.     

Phylogenetic implications of endosperm cell wall morphology in Douglasia, Androsace, and Vitaliana (Primulaceae)

We investigated differences in endosperm cell wall thickening in three genera of Primulaceae; seven species of Douglasia , 28 species of Androsace representing the four sections of the genus, and the single species of the genus Vitaliana . Endosperm cell walls have irregular wall thickenings and narrow constrictions and characterize all Douglasia species and Vitaliana , whereas evenly thickened cell walls without constrictions were found in the endosperm of Androsace and Primula . The endosperm character supports the monophyly of Douglasia and Vitaliana , and we discuss the phylogenetic implications of this conclusion.     

Barcoding Fauna Bavarica: Myriapoda - a contribution to DNA sequence-based identifications of centipedes and millipedes (Chilopoda, Diplopoda)

We give a first account of our ongoing barcoding activities on Bavarian myriapods in the framework of the Barcoding Fauna Bavarica project and IBOL, the International Barcode of Life. Having analyzed 126 taxa (including 122 species) belonging to all major German chilopod and diplopod lineages, often using four or more specimens each, at the moment our species stock includes 82% of the diplopods and 65% of the chilopods found in Bavaria, southern Germany. The partial COI sequences allow correct identification of more than 95% of the current set of Bavarian species. Moreover, most of the myriapod orders and families appear as distinct clades in neighbour-joining trees, although the phylogenetic relationships between them are not always depicted correctly. We give examples of (1) high interspecific sequence variability among closely related species; (2) low interspecific variability in some chordeumatidan genera, indicating that recent speciations cannot be resolved with certainty using COI DNA barcodes; (3) high intraspecific variation in some genera, suggesting the existence of cryptic lineages; and (4) the possible polyphyly of some taxa, i.e. the chordeumatidan genus Ochogona. This shows that, in addition to species identification, our data may be useful in various ways in the context of species delimitations, taxonomic revisions and analyses of ongoing speciation processes.     

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.     

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.     

Variation and specialisation of the forcipular apparatus of centipedes (Arthropoda: Chilopoda): A comparative morphometric and microscopic investigation of an evolutionary novelty

The forcipules of centipedes are the only known example in the animal kingdom of an evolutionary transition from walking legs to venom-injecting appendages. They provide a classic case of an evolutionary novelty under most (but not all) definitions of that concept. Although there is a reasonable literature on forcipules, and on the forcipular segment more generally, it is fragmentary and scattered. Also, many previous studies have been based on a single species and hence have no comparative component. Here, we build on this earlier literature by providing detailed qualitative and quantitative information on the forcipular segments of representatives of the five extant orders of centipedes. Our results reveal notable differences between the orders – as well as considerable variation within some of them. The pattern of inter-group differences can be used to infer, albeit cautiously, a major evolutionary trend from a presumed scutigeromorph-like last common ancestor (LCA), in which the forcipules were probably leg-like (as in present-day scutigeromorphs) to a more specialized claw-like structure with movement restricted to the horizontal plane. This morphological trend may reflect an ecological trend from open-habitat ambush predation to leaf-litter and subterranean predatory opportunism.     

Phylogenetic reconstruction of Polymastiidae (Demospongiae: Hadromerida) based on morphology

Phylogeny of the sponge family Polymastiidae was reconstructed based on 25 morphological characters. Twenty-one polymastiid species and three suberitid species, Suberites domuncula as outgroup, Aaptos aaptos and A. papillata as sister groups, were included in the analyses. The reconstructions were done in PAUP* running heuristic search with the parsimony criterion. We analysed three possible evolutionary scenarios based on three alternative interpretations of the body plan of Quasillina brevis and Ridleia oviformis: first—Ridleia possesses aquiferous papillae whereas Quasillina lacks them, second—both genera lack papillae and third—the body in both genera is a single hyper-developed papilla. All three scenarios excluded the secondary loss of the papillae in the polymastiid evolution. Scenario 2 also excluded the secondary loss of the regular choanosomal skeleton, while scenario 1 assumed its loss in Ridleia and scenario 3 admitted its loss in both Ridleia and Quasillina. We prioritised scenario 2 due to its maximal parsimony and rescaled consistency index and subsequently favoured the clustering of Ridleia and Quasillina separately from the monophyletic polymastiid clade. In all three scenarios Pseudotrachya hystrix clustered separately from other polymastiids in agreement with the molecular evidence, and thus the exclusion of Pseudotrachya from Polymastiidae was proposed. The relationships between A. papillata, Tentorium semisuberites, Polymastia uberrima, the clade Weberella bursa + Polymastia boletiformis and the main polymastiid clade were ambiguous. Meanwhile, all scenarios showed the non-monophyly of Polymastia and Aaptos. Our hypotheses should be tested by reconstructions based on larger taxon sampling of hadromerid species and larger sets of morphological and molecular characters before any ultimate taxonomic decisions are taken.