Centipede Species Research Articles

The First Venomous Crustacean Revealed by Transcriptomics and Functional Morphology: Remipede Venom Glands Express a Unique Toxin Cocktail Dominated by Enzymes and a Neurotoxin

Animal venoms have evolved many times. Venomous species are especially common in three of the four main groups of arthropods (Chelicerata, Myriapoda, and Hexapoda), which together represent tens of thousands of species of venomous spiders, scorpions, centipedes, and hymenopterans. Surprisingly, despite their great diversity of body plans, there is no unambiguous evidence that any crustacean is venomous. We provide the first conclusive evidence that the aquatic, blind, and cave-dwelling remipede crustaceans are venomous and that venoms evolved in all four major arthropod groups. We produced a three-dimensional reconstruction of the venom delivery apparatus of the remipede Speleonectes tulumensis, showing that remipedes can inject venom in a controlled manner. A transcriptomic profile of its venom glands shows that they express a unique cocktail of transcripts coding for known venom toxins, including a diversity of enzymes and a probable paralytic neurotoxin very similar to one described from spider venom. We screened a transcriptomic library obtained from whole animals and identified a nontoxin paralog of the remipede neurotoxin that is not expressed in the venom glands. This allowed us to reconstruct its probable evolutionary origin and underlines the importance of incorporating data derived from nonvenom gland tissue to elucidate the evolution of candidate venom proteins. This first glimpse into the venom of a crustacean and primitively aquatic arthropod reveals conspicuous differences from the venoms of other predatory arthropods such as centipedes, scorpions, and spiders and contributes valuable information for ultimately disentangling the many factors shaping the biology and evolution of venoms and venomous species.

Venomic and Transcriptomic Analysis of Centipede Scolopendra subspinipes dehaani

Centipedes have venom glands in their first pair of limbs, and their venoms contain a large number of components with different biochemical and pharmacological properties. However, information about the compositions and functions of their venoms is largely unknown. In this study, Scolopendra subspinipes dehaani venoms were systematically investigated by transcriptomic and proteomic analysis coupled with biological function assays. After random screening approximately 1500 independent clones, 1122 full length cDNA sequences, which encode 543 different proteins, were cloned from a constructed cDNA library using a pair of venom glands from a single centipede species. Neurotoxins, ion channel acting components and venom allergens were the main fractions of the crude venom as revealed by transcriptomic analysis. Meanwhile, 40 proteins/peptides were purified and characterized from crude venom of S. subspinipes dehaani. The N-terminal amino acid sequencing and mass spectrum results of 29 out of these 40 proteins or peptides matched well with their corresponding cDNAs. The purified proteins/peptides showed different pharmacological properties, including the following: (1) platelet aggregating activity; (2) anticoagulant activity; (3) phospholipase A(2) activity; (4) trypsin inhibiting activity; (5) voltage-gated potassium channel activities; (6) voltage-gated sodium channel activities; (7) voltage-gated calcium channel activities. Most of them showed no significant similarity to other protein sequences deposited in the known public database. This work provides the largest number of protein or peptide candidates with medical-pharmaceutical significance and reveals the toxin nature of centipede S. subspinipes dehaani venom.

Effects of reforestation with Quercus species on selected arthropod assemblages (Isopoda Oniscidea, Chilopoda, Coleoptera Carabidae) in a Mediterranean area

Arthropod assemblages (Isopoda Oniscidea, Chilopoda, Coleoptera Carabidae) were studied in three 20-year-old reforestations with native oak species (holm-oak Quercus ilex, Turkey-oak Quercus cerris, and downy-oak Quercus pubescens) and in a natural mixed oak forest (Q. cerris and Quercus frainetto) in Latium (central Italy). The three reforested areas had been previously used for agricultural purposes. Samples were collected monthly by pitfall traps for a period of 1year (March 2009–February 2010). Structural parameters and the arthropod assemblage compositions of the four studied areas were compared. The effects of the different forest types and the influence of environmental variables on the activity density of each species were analyzed, with particular emphasis to forest species. Contrary to centipedes and ground beetles, woodlice showed lower values of richness, diversity and equitability in reforestations than in the natural forest. According to Canonical Correspondence Analysis, forest species of woodlice and centipedes resulted mainly associated with forests characterized by a high structural heterogeneity (natural forest and reforestation with Q. pubescens). In these two forests the activity density of centipede forest species is mainly influenced by the coverage of both shrub and leaf litter layer, and woodlice forest species only by the coverage of the latter. The ground beetle forest species were mainly associated with forests characterized by low structural heterogeneity and an almost total closure of the canopy throughout the year (reforestation with Q. ilex). However, some ground beetle forest species are present also in the natural forest. Our results suggest that reforestations with different native broadleaf species belonging to the local “potential” vegetation can contribute to the conservation of the diversity of forest arthropod assemblages in the extremely fragmented agricultural landscape of the middle Tyrrhenian area.

Discovery of the subgenus Lithobius (Sigibius) Chamberlin, 1913 (Chilopoda: Lithobiomorpha: Lithobiidae) in East Asia: A review the Chinese species

The present paper reviews the centipede species of the subgenus Lithobius (Sigibius) Chamberlin, 1913 (Lithobiomorpha: Lith-obiidae) of the Chinese fauna, including a new species Lithobius (Sigibius) trichinocaput sp. n. recently discovered from HebeiProvince, the only East Asian record of this otherwise Central and North America subgenus of Lithobiomorpha. Diagnoses ofthe subgenus and the species, detailed account on species distribution and a key to the Chinese Lithobius (Sigibius) species are presented.

Ultrastructure and regeneration of midgut epithelial cells in Lithobius forficatus (Chilopoda, Lithobiidae)

AbstractLithobius forficatus (Myriapoda, Chilopoda, Lithobiidae) is a widespread species of centipede that is common across Europe. Its midgut epithelial cells are an important line of defense against toxic substances that originate in food, such as pathogens and metals. Despite this important role, the biology of the midgut epithelium is not well known. Here we describe the ultrastructure of the midgut epithelium, as well as the replacement of degenerated midgut epithelial cells. The midgut epithelium of L. forficatus is composed of digestive, secretory, and regenerative cells. The cytoplasm of digestive cells shows regionalization in organelle distribution, which is consistent with the role of these cells in secretion of enzymes, absorption of nutrients, and accumulation of lipids and glycogen. Secretory cells, which do not reach the luminal surface of the midgut epithelium, possess numerous electron‐dense and electron‐lucent granules and may have an endocrine function. Hemidesmosomes anchor secretory cells to the basal lamina. Regenerative cells play the role of midgut stem cells, as they are able to proliferate and differentiate. Their proliferation occurs in a continuous manner, and their progeny differentiate only into digestive cells. The regeneration of secretory cells was not observed. Mitotic divisions of regenerative cells were confirmed using immunolabeling against BrdU and phosphohistone H3. Hemocytes associate with the midgut epithelium, accumulating between the visceral muscles and beneath the basal lamina of the midgut epithelium. Hemocytes also occur among the digestive cells of the midgut epithelium in animals infected with Rickettsia‐like microorganisms. These hemocytes presumably have an immunoprotective function in the midgut.

Trophic interactions in centipedes (Chilopoda, Myriapoda) as indicated by fatty acid patterns: Variations with life stage, forest age and season

Fatty acid (FA) analysis investigates changes in the relative contribution of prey from major energy channels in decomposer food webs for predator nutrition. Adopting this approach we investigated whether the trophic niche of centipedes, as major invertebrate predators in forest soil food webs, changes with maturation, season or forest age. Generally, each of the four centipede species studied differed significantly in their FA composition suggesting trophic niche differentiation. FA profiles differed more strongly in the two geophilomorph (Strigamia acuminata and Geophilus ribauti) than in the two lithobiomorph species (Lithobius crassipes and Lithobius mutabilis) suggesting that in particular the former feed on markedly different prey. FA profiles changed during post-embryonic development in each of the four centipede species. Differences were most pronounced in the two lithobiomorph species shifting to predominantly fungal feeding prey. Further, FA profiles varied with season indicating that centipedes exploit more prey out of the bacterial channel in autumn. FA profiles of centipedes varied little with forest age suggesting that soil food webs are remarkably invariant across different forest ecosystems. The results indicate that FA composition of second order consumers closely reflects changes in diet of prey species and composition of basal resources. The study proved FA profiles as powerful tool to gain insight into critical characteristics of soil food web stability, i.e., compartmentalisation and the relative importance and variability of energy channels.

A new species of centipede of the genus Cryptops Leach (Scolopendromorpha: Cryptopidae) from southern Western Ghats with a key to the species of Cryptops in India

A new species of blind cryptopid centipede of the genus Cryptops Leach belonging to the hortensis group viz. Cryptops (C.) malabarensis is described from the southern Western Ghats, Kerala, India and the family Cryptopidae (Scolopendromorpha) is reported for the first time from the area. Affinities of the new species with a Madagascar species are discussed and a key to separate the Indian species of Cryptops is also provided.

Book review: Gerlach J, Marusik Y (Eds) (2010) Arachnida and Myriapoda of the Seychelles islands. Siri Scientific Press, 435 pp.

The Seychelles islands are located in the Indian Ocean, some 1,500 kilometres east of Africa and northeast of the island of Madagascar. They are particularly interesting from a biogeographical perspective, with ancient affinities to Africa and Asia, recent colonizing species from the Indo-Pacific and modern introductions. Until recently, relatively little was known about the biodiversity of these islands. However, this has changed through the publication of a series of monographs on the Seychelles fauna, presenting the latest information on all the terrestrial and freshwater animals of the islands. In this volume on the Arachnida and Myriapoda of the Seychelles islands, 14 expert scientists from 11 different countries have provided contributions that cover all 433 species of spiders, scorpions, mites, centipedes, millipedes and related groups currently recorded from the islands, including systematic keys, diagnostic illustrations and descriptions for most species, in addition to distribution records. The volume opens with a brief history of arachnid and myriapod collecting in the Seychelles islands followed by summary data for the numbers of indigenous, endemic and introduced species for all the orders covered and a brief classification scheme for the Arachnida. The following chapters cover the various different orders as follows: Araneae (M.I. Saaristo [deceased], 300 pp., 216 species), Amblypygi (P. Weygoldt, 5 pp.), Opiliones (J. Gerlach, 8 pp.), Scorpiones (L. Prendini, 10 pp.), the smaller arachnid orders including Pseudoscorpiones, Schizomida and Palpigradi (M. Harvey, 10 pp.), Acari: Holothyroidea (J. Gerlach, P. Lehtinen & M. Madl, 7 pp.), Acari: Oribatida (W. Niedbala, 6 pp.), additional Acari (J. Gerlach, 5 pp.), Geophilomorpha (L. Bonato & A. Minelli, 16 pp.), Lithobiomorpha & Scutigerimorpha (P. Stoev & J. Gerlach, 3 pp.), Scolopendromorpha (J.G.E. Lewis, 8 pp.), Diplopoda (S. Golovatch & J. Gerlach, 16 pp.), Symphyla and Pauropoda (J. Gerlach, 6 pp.). Each chapter is fully referenced with its own reference list at the end. Thus, the majority of the book is devoted to arachnids and spiders in particular, although this is a true reflection of the relative diversity of these groups on the islands. In total, 362 arachnids and 71 myriapod species are documented. In terms of formal taxonomic/nomenclatural changes, one new synonymy is established in the spider family Filistatidae, two new genera of holothyrid mites are described, in addition to three new combinations in the latter group. The book is richly illustrated throughout with clear line drawings, which will permit identification of most of the species covered. The last chapter includes assessments of species conservation status as defined by the International Union for the Conservation of Nature (IUCN) for the indigenous and endemic taxa. There is a comprehensive index. On the downside, some chapters are more thorough than others, but this is presumably a reflection of the state of knowledge and/or the availability of taxonomic experts with experience in this geographical region. For example, many of the mite species are merely listed as known records without any associated figures, although this still represents a valuable systematic inventory for this group as locality data and citing references are provided. Furthermore, not all Diplopoda are figured and some of the figures provided for this group are rather small. Overall the book is well produced and the editing is tight, although some of the figures and and several of the photographs have not reproduced as well as they may have done, and some e.g. in the Diplopoda chapter are rather small, but they all still serve their intended purpose. Most of the figures are new, although several have been reproduced from other works. It is also the first book to provide complete coverage of the known spider fauna of any single country in the Southern Hemispere. In summary, it provides a comprehensive synthesis of current arachnological and myriapodological knowledge for this previously poorly known region. This review was written by an arachnologist, but I expect the book will be of use to invertebrate taxonomists and biogeographers and deserves a place on the shelves of any serious arachnological or myriapodological taxonomic library. Price: £60.00 (available directly from the publisher: http://www.siriscientificpress.co.uk)

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.

The species-area relationship in centipedes (Myriapoda: Chilopoda): a comparison between Mediterranean island groups

The present study article examines the shapes of centipede species–area relationships (SARs) in the Mediterranean islands, compares the results of the linear form of the power model between archipelagos, discusses biological significance of the power model parameters with other taxa on the Aegean archipelago, and tests for a significant small-island effect (SIE). We used 11 models to test the SARs and we compared the quality-of-fit of all candidate models. The power function ranked first and Z-values was in the range 0.106–0.334. We assessed the presence of SIEs by fitting both a continuous and discontinuous breakpoint regression model. The continuous breakpoint regression functions never performed much better than the closest discontinuous model as a predictor of centipede species richness. We suggest that the relatively low Z-values in our data partly reflect better dispersal abilities in centipedes than in other soil invertebrate taxa. Longer periods of isolation and more recent island formation may explain the somewhat lower constant c in the western Mediterranean islands compared to the Aegean islands. Higher breakpoint values in the western Mediterranean may also be a result of larger distance to the mainland and longer separation times. Despite the differences in the geological history and the idiosyncratic features of the main island groups considered, the overall results are quite similar and this could be assigned to the ability of centipedes to disperse across isolation barriers.

A new centipede species of Lithobius Leach (Lithobiomorpha: Lithobiidae) from China

A new centipede species of the genus Lithobius Leach (Lithobiomorpha: Lithobiidae), namely Lithobius zhui sp. nov., from Xinjiang Autonomous Region, China, is described. A key to the 15 species of the subgenus from China are provided.

Comparative phylogeography of the centipedes Cryptops pictus and C. niuensis (Chilopoda) in New Caledonia, Fiji and Vanuatu

The South Pacific is a biodiverse region of extreme evolutionary importance because it harbors ancient lineages and recent radiations. However, few population-level studies of genetic variation have been conducted in the land masses of this region. Likewise, the number of population-level studies using myriapods as models is extremely small. In this article, we compare the genetic structure of two species of centipedes in the genus Cryptops endemic to the South Pacific, one from a continental island, the other from oceanic islands. The level of genetic diversity and structure in C. pictus, a species endemic to New Caledonia, is much higher than in C. niuensis in Fiji and Vanuatu, despite the fact that C. niuensis is spread across two different archipelagos and several islands. The most likely explanation is the relatively young age of the remnants of the Vitiaz Arc (Fiji and Vanuatu) compared to New Caledonia. Using the emergence of Fiji-Vanuatu as a calibration point, C. pictus is estimated to have diverged by 23.4 Mya (upper 95% confidence interval) with a mean estimate of 11.7 Mya versus the 9.7 Mya of C. niuensis. Considering the absence of shared sequences between specimens from different sampling sites and the high genetic structuring within populations, C. pictus appears to be an ideal candidate to assess historical processes at a micro-evolutionary scale in New Caledonia.

A further contribution to the knowledge of Pectiniunguis minutus (Demange, 1968), a little known dwarf schendylid centipede from western equatorial Africa (Chilopoda: geophilomorpha)

The geophilomorph centipede Pectiniunguis minutus (Demange, 1968), a little known dwarf schendylid from Gabon (Western equatorial Africa), is redescribed and illustrated based on the type material and an additional non-type specimen preserved in the collections of the Muséum national d'Histoire naturelle, Paris. P. minutus can be easily differentiated from all the other species currently assigned to the genus Pectiniunguis, by the very low number of leg-bearing segments (35 in the males, 37 or 39 in the females) and very small body size (12-16 mm in length). P. minutus is also distinguished by having ventral pore-fields on the anterior region of the body only, this character being shared by a single species of the genus, i.e., P. ascendens Pereira, Minelli & Barbieri, 1994, from the Neotropics (Brazil: State of Amazonas) with which a morphological comparison is given. Comments about other dwarf centipede species belonging to several families of the order Geophilomorpha, are also added.

A review of the Chinese species of Hessebius Verhoeff, 1941 (Chilopoda: Lithobiomorpha: Lithobiidae)

The present paper reviews the centipede species of the genus Hessebius Verhoeff, 1941 (Chilopoda: Lithobiomorpha: Lithobiidae) of the Chinese fauna, H. jangtseanus (Verhoeff, 1942), H. longispinipes Ma, Pei & Zhu, 2009 and H. multiforaminis Pei, Ma, Zapparoli & Zhu, sp. nov. H. kosswigi Verhoeff, 1941, from Turkey, is selected by subsequent designation as type species of the genus, which was previously unavailable. Schizotergitius styliferus Loksa, 1978, from Mongolia, is transferred in Hessebius (n. comb.) and its relationships with H. longispinipes are briefly discussed. A comparative table of the diagnostic characters of the Sino–Mongolian species of the genus is given.

Nestedness in centipede (Chilopoda) assemblages on continental islands (Aegean, Greece)

In natural ecosystems, species assemblages among isolated ecological communities such as continental islands often show a nested pattern in which biotas of sites with low species richness are non-random subsets of biotas of richer sites. The distribution of centipede (Chilopoda) species in the central and south Aegean archipelago was tested for nestedness. To achieve this aim we used distribution data for 53 species collected on 24 continental Aegean islands (Kyklades and Dodekanisa). Based on the first-order jackknife estimator, most of islands were comprehensively surveyed. In order to quantify nestedness, we used the nestedness temperature calculator (NTC) as well as the nestedness metric based on overlap and decreasing Fill (NODF). NTC indicated that data exhibited a high degree of nestedness in the central and south Aegean island complexes. As far as the Kyklades and Dodekanisa are concerned, NTC showed less nested centipede structures than the 24 islands. Likewise, NODF revealed a significant degree of nestedness in central and south Aegean islands. It also showed that biotas matrices without singletons were more nested than the complete ones (Aegean, Kyklades and Dodekanisa). The two commonest centipede taxa (lithobiomorphs and geophilomorphs) contributed differently to centipede assemblages. In the Kyklades and Dodekanisa, geophilomorphs did not show a reliable nested arrangement unlike lithobiomorphs. In relation to the entire data set, nestedness was positively associated with the degree of isolation. In the Kyklades altitudinal range best explained nestedness patterns, while in Dodekanisa habitat heterogeneity proved to be more important for the centipede communities. Island area does not seem to be a significant explanatory variable. Some of our results from the Kyklades were critically compared with those for terrestrial isopod and land snail nested assemblages from the same geographical area. The complex geological and palaeogeographical history of the Aegean archipelago partly accounted for the pattern of centipede assemblages.

Demonstration of a heritable component of the variation in segment number in the centipede Strigamia maritima

Here we address the question of how arthropod segment number may evolve by reporting the results of further work on the model system Strigamia maritima. Recently, we showed that there was a plastic component of the variation in segment number within this species; now we demonstrate that there is also a heritable component. This is important because it enables a connection to be made between the known latitudinal trend among species of geophilomorph centipedes (more segments at lower latitudes) and the parallel trend within them. This latter trend is best documented in S. maritima but is also known in several other species. However, while a general connection between the inter- and intraspecific trends can now be made, deciding upon a specific hypothesis of the nature of the selection involved is still problematic. We provide two alternative hypotheses, one based on the temperature-related plasticity in segment number being adaptive, the other based on it being nonadaptive.

Comparative morphological study of the venom glands of the centipede Cryptops iheringi, Otostigmus pradoi and Scolopendra viridicornis

Centipedes are widely distributed over all the continents. As they are well adapted to urban areas they can often cause accidents to humans by injecting venom produced in the glands located inside their maxillipeds. The fine morphology of the centipede venom glands is practically unknown. This present study is the first comparative report on the histology, histochemistry and ultrastructure of the venom glands of the centipede species responsible for the majority of accidents to humans in Brazil: Scolopendra viridicornis, Cryptops iheringi and Otostigmus pradoi. In all species the glands are basically composed of columnar secretory cells radially disposed side by side, individually opening through pores in a central chitinous duct. Each secretory cell is covered by striated muscular fibres. The secretion has the form of small PAS positive granules and large hyaline secretory bromophenol blue positive vacuoles, indicating the presence of neutral polysaccharides and protein. The secretion is conducted through the secretory cell necks to the pores, which open into the central chitinous duct. The results indicate a great similarity both in morphology and primary chemical composition of the venom among the studied species, except for the size of the glands, which is proportional to the body dimensions of each species.

A review of the Chinese species of Bothropolys Wood, 1862 (Chilopoda: Lithobiomorpha: Lithobiidae)

The present paper reviews the centipede species of the genus Bothropolys Wood, 1862 (Lithobiomorpha: Lithobiidae) of the Chinese fauna, including two species recorded from China for the first time. Diagnoses of the genus and the species, detailed account on species distribution and a key to the Chinese Bothropolys species are included.

Ecology of the Southeastern Crowned Snake,Tantilla coronata

There are very few comprehensive studies of the ecology of small-bodied snakes. Here, we describe the ecology and demography of the Southeastern Crowned Snake (Tantilla coronata) based on 1,640 captures on the Savannah River Site in the Upper Coastal Plain of South Carolina, USA from 1951–2007. Female T. coronata were significantly longer, heavier, and heavier-bodied than males but had relatively shorter tails. Clutch size based on oviductal eggs was positively correlated to maternal body mass and length. Snakes exhibited a unimodal seasonal activity pattern that peaked in summer. Pitfall captures were significantly male-biased from July–October, corresponding to the suggested mating period for this species in this part of its range. We identified three classes of animals in the population: neonates, second year animals, and older animals that included both non-reproductive subadults and reproductive adults. Longevity was at least five years for two recaptured males originally captured as mature adults. Centipede species were the exclusive prey identified from T. coronata collected on the Savannah River Site. Our study demonstrates that research on underrepresented species is possible and can contribute to understanding of snake ecology.

Toxic activities of Brazilian centipede venoms

Centipedes have a venom gland connected to a pair of forceps, which are used to arrest preys. Human victims bitten by centipedes usually manifest burning pain, paresthesia and edema, which may develop into superficial necrosis. The aim of this work was to characterize and compare toxic activities found in venoms of three species of Brazilian centipedes— Otostigmus pradoi, Cryptops iheringi and Scolopendra viridicornis. By SDS–PAGE (4–20%), important differences were noticed among venoms (between 7 and 205 kDa). Few bands showed feeble caseinolytic, fibrinogenolytic and gelatinolytic activities by zymography, but strong hyaluronidase activity was observed in S. viridicornis and O. pradoi venoms. In addition, such activities could be inhibited by o-phenanthroline, indicating that these enzymes are metalloproteinases. All venoms induced nociception, edema and myotoxicity in mice, but only S. viridicornis induced mild hemorrhagic activity. No coagulant activity was detected in centipede venoms. Low phospholipase A 2 activity was observed exclusively in S. viridicornis and O. pradoi venoms, but these venoms had intense direct hemolytic activity on human erythrocytes. Cross-reactivity among venoms was observed using species-specific sera raised in rabbits. Differences were noticed among centipede venoms, but S. viridicornis is indeed the most toxic venom and thereby it could induce a more severe envenomation.