A Comparative Morphometric Study on Odontobuthus bidentatus and Odontobuthus doriae (Scorpionida: Buthidae) in Iran.

Scorpions are one of the most venomous animals which cause serious public health problems. The sting of scorpions can sometimes be fatal depending on the scorpion species involved. So far, sixty-six (66) scorpion species have been identified in Iran. Annually, about 40-50000 cases of scorpionism are reported in Iran. Odontobuthus doriae and O. bidentatus are among the most medically important scorpion species in Iran, and they are very similar to each other in coloration, carination, and trichobotrial patterns. This morphometric study aimed to compare some of the important morphological characteristics in order to identify the key differences between these two species. A total of 45 morphological characters were measured using calipers and stereomicroscope, and 55 morphological characters and ratios (relative of length to width ratio of morphological characters of scorpions) were analyzed. The independent sample t-test in SPSS software (version 24) was used for the statistical analyses in this study. The mean total length, carapace width, length of fixed and moveable fingers, and chelicerae length of O. doriae were greater than those of O. bidentatus in our study area. The morphological measurements displayed a clear distinction between O. doriae and O. bidentatus in our study area; therefore, they can be used as morphological identification keys for distinguishing between these two species.

Back to Tityus serrulatus Lutz & Mello, 1922 (Scorpiones: Buthidae): new comments about an old species

A synopsis on the historical, geographical and ecological aspects related to the most conspicuous scorpion species of the genus Tityus known from Brazil is proposed. Tityus serrulatus Lutz & Mello, 1922 was described precisely one century ago, nevertheless many questions related to its ecological adaptations and geographical expansion remain without a precise response. This species, well known for its infamous reputation of noxious species, is also known for its capacity to reproduce asexually, by parthenogenesis. Although the individuals of a given population are considered clones, a new hypothesis could suggest the occurrence of mutations within isolated individuals, leading to distinct subpopulations that could present better phenotypic performances in ecological habitats distinct from those of the original area of distribution of the species.(AU)

Burrowing into the forest: Phylogeny of the Asian forest scorpions (Scorpionidae: Heterometrinae) and the evolution of ecomorphotypes.

Asian forest scorpions (Scorpionidae Latreille, 1802: Heterometrinae Simon, 1879) are distributed across South and Southeast Asia. All are fossorial, constructing burrows under stones or in open ground, in habitats differing in precipitation and vegetation cover, from rainforests and tropical deciduous forests to savanna and scrubland. The systematics of these scorpions has long been confused due to bad taxonomy and the absence of a phylogenetic framework. Although the monophyly of the group was previously confirmed as part of broader phylogenetic analyses based on exemplar species, the only quantitative analysis of species-level variation to date was based on overall similarity. This contribution presents the first species-level phylogenetic analysis of Asian Scorpionidae, based on 186 morphological characters and 4188 aligned base-pairs of DNA sequence data from two nuclear and three mitochondrial loci for 132 terminals including all 41 ingroup species and four outgroup species. Simultaneous analyses of the morphological and molecular datasets with parsimony, Maximum Likelihood and Bayesian Inference provided the framework for a revised classification presented elsewhere. In order to understand how adaptation following dispersal into new habitats has driven the morphological diversification of Asian forest scorpions, species were scored for 10 characters concerning morphology and burrow architecture, which contributed to an ensemble index of adaptation to habitat aridity. Species were classified into three ecomorphotypes based on the index, and ancestral state reconstruction of ecomorphotypes performed on the phylogeny. A pattern was recovered in which lineages and species occurring in different habitats on a continuum from wet (evergreen forest) to dry (savanna, scrubland) exhibited characters presumed to be adaptive and hence responsible for driving scorpion diversification.

Transcriptomic and proteomic analyses of venom glands from scorpions Liocheles australasiae, Mesobuthus martensii, and Scorpio maurus palmatus.

Scorpion venom contains a cocktail of differing peptides and proteins. Previous studies focused on the identification of species-specific components in scorpion venoms, and whether there could be peptides and/or proteins conserved in the venom gland of a scorpion ancestor has been rarely investigated. Here, using a combination of transcriptomic and proteomic approaches, putative conserved toxins from the venom glands of scorpions Liocheles australasiae, Mesobuthus martensii, and Scorpio maurus palmatus were identified and compared. Similar to other studies, more than half of the conserved toxins are predominantly proteins including proteases. On the other hand, unique venom peptides, including ion channel toxins were revealed specifically in the M. martensii. The sodium channel toxin peptides revealed in M. martensii consolidated that scorpions in the Buthidae are able to envenomate their prey wih highly neurotoxic venom. This study suggested that these conserved proteins had already formed part of the arsenal in the venom gland of the common ancestor of scorpions, and likely perform important functional roles in envenomation during scorpion evolution.

Some additional comments on the Ananteridae or 'Ananteris group' and description of a new species of Ananteris Thorell from Central Brazil (Scorpiones: Buthidae).

Some additional comments are proposed on the historical aspects of the subfamily Ananterinae (sensu Pocock 1900). The worldwide pattern of distribution of the elements associated with this subfamily is briefly discussed. The biogeographic patterns presented by extant and fossil elements of this subfamily confirm not only the characteristics of a group presenting a typical Gondwanian distribution, but may also correspond to older Pangaean patterns. One unexpected new species is described for the genus Ananteris Thorell from the State of Goiás in Central Brazil, Ananteris martensi sp. n., suggesting that the diversity levels of the Cerrados formations are most certainly higher than what is normally admitted.

Phylogeographic Patterns in a Semi-Lithophilous Burrowing Scorpion, Opistophthalmus pallipes, from South Africa.

Southern Africa has a diverse endemic scorpion fauna, but a paucity of information currently confounds conservation of the group. Phylogeographic approaches represent a useful tool to identify the patterns and processes which underpin scorpion diversity, but these studies are lacking for southern African species. Among southern African scorpions, the semi-lithophilous Opistophthalmus pallipes has strict habitat requirements, and a distribution historically subjected to profound environmental turnover. As such, the species offers a model system to investigate the interplay between intrinsic and extrinsic factors as drivers of diversity and endemism. To investigate spatial genetic patterns within O. pallipes and the possible drivers thereof, the current study combines mitochondrial DNA and ecological information under a phylogeographic approach. The species is characterized by several genetically discrete and divergent populations. The factors which shape these genetic patterns appear to be both intrinsically (ecological specificity) and extrinsically (landscape structure and ecogeographic conditions) influenced, with major divergences corresponding to periods of profound environmental changes. Taken together, the findings of this study provide evidence of spatial genetic isolation and genetic diversity within a stenotopic southern African scorpion species. These findings partly explain the staggering diversity and endemism in southern African scorpions, but further phylogeographic studies are necessary to propose conservation scenarios for this group.

Epidemiology of scorpionism in France: nationwide scorpion exposure.

INTRODUCTION: In France, 57 species of scorpions are described with a limited number of clinical studies. In this article, we report the epidemiology of scorpion sting events in mainland France and its overseas territories based on cases reported to the French poison-control centres (FPCC). MATERIAL AND METHOD: This retrospective multicentre study was conducted with data from FPCC's files about scorpion stings between January 1, 2011 and December 31, 2020. RESULT: Among 975 recorded files, 624 patients were included because they were stung by scorpions native to French territories. Most stings occurred along the Mediterranean coast in summer and indoors (in persons' homes) during the daytime. The scorpions were identified in 50% of cases. According to signs of envenoming, patients were divided into class III (2 cases; 1%), class II (51 cases; 8%), class I (444 cases; 71%) and asymptomatic stings (127 cases; 20%). Twelve pregnant women were stung and two of them had contractions, which triggered childbirth in one woman. Ten patients had local infections in the first week after the sting. One patient had venous thrombosis 2 days after the sting. Life-threatening scorpions, i.e., Tityus obscurus, Tityus sylvestris and Centruroides pococki, in French territories are limited to French Guiana and Lesser Antilles. Class II envenoming cases are recorded for Buthus occitanus, Euscorpius spp. in mainland France, and Isometrus maculatus in French Guiana, the Lesser Antilles (Guadeloupe and Martinique) and territories in the Indian Ocean (Mayotte and Réunion Island) and Pacific Ocean (French Polynesia). Only cases of local manifestation was reported for Belisarius xambeui in mainland France. CONCLUSION: Scorpion stings in French territories are frequently on the Mediterranean coast and French Guiana. Life-threatening cases are limited to T. obscurus, T. sylvestris and Centruroides pococki stings in French Guiana and Lesser Antilles.

Taxonomic Sampling and Rare Genomic Changes Overcome Long-Branch Attraction in the Phylogenetic Placement of Pseudoscorpions.

Long-branch attraction is a systematic artifact that results in erroneous groupings of fast-evolving taxa. The combination of short, deep internodes in tandem with long-branch attraction artifacts has produced empirically intractable parts of the Tree of Life. One such group is the arthropod subphylum Chelicerata, whose backbone phylogeny has remained unstable despite improvements in phylogenetic methods and genome-scale data sets. Pseudoscorpion placement is particularly variable across data sets and analytical frameworks, with this group either clustering with other long-branch orders or with Arachnopulmonata (scorpions and tetrapulmonates). To surmount long-branch attraction, we investigated the effect of taxonomic sampling via sequential deletion of basally branching pseudoscorpion superfamilies, as well as varying gene occupancy thresholds in supermatrices. We show that concatenated supermatrices and coalescent-based summary species tree approaches support a sister group relationship of pseudoscorpions and scorpions, when more of the basally branching taxa are sampled. Matrix completeness had demonstrably less influence on tree topology. As an external arbiter of phylogenetic placement, we leveraged the recent discovery of an ancient genome duplication in the common ancestor of Arachnopulmonata as a litmus test for competing hypotheses of pseudoscorpion relationships. We generated a high-quality developmental transcriptome and the first genome for pseudoscorpions to assess the incidence of arachnopulmonate-specific duplications (e.g., homeobox genes and miRNAs). Our results support the inclusion of pseudoscorpions in Arachnopulmonata (new definition), as the sister group of scorpions. Panscorpiones (new name) is proposed for the clade uniting Scorpiones and Pseudoscorpiones.

Karyotype evolution and preliminary molecular assessment of genera in the family Scorpiopidae (Arachnida: Scorpiones).

The scorpions represent an ancient and morphologically conserved order of arachnids. Despite that, their karyotypes may differ considerably even among closely related species. In this study, we identify the trends of the karyotype evolution in the family Scorpiopidae based on integrating cytogenetic data and multi-locus molecular phylogenetic approaches. We detected considerable variability in diploid numbers of chromosomes (from 48 to 147), 18S rRNA gene cluster positions (from terminal to pericentromeric) at the interspecific level. Moreover, we identified independent fusions, fissions and inversions in the evolution of the family Scorpiopidae, leading to a remarkable diversification of the karyotypes. The dynamic system of the karyotype changes in this group is further documented by the presence of interstitial telomeric sequences (ITS) in two species. The cytogenetic differences observed among the analyzed species highlight the potential of this type of data for species-level taxonomy in scorpion lineages with monocentric chromosomes. Additionally, the results of our phylogenetic analyses support the monophyly of the family Scorpiopidae, but rendered several genera para- or polyphyletic.

Scorpion species of medical importance in the Brazilian Amazon: a review to identify knowledge gaps

Scorpionism is a relevant medical condition in Brazil. It is responsible for most accidents involving venomous animals in the country, which leads to severe symptoms that can evolve to death. In recent years, an increase of almost 50% in the incidence of scorpionism has been observed in the Northern Region, where the highest severity of envenoming has been notified since the beginning of the 21st century. This review aims to provide an in-depth assessment of public data and reports on symptoms and epidemiology of envenoming, ecological aspects of scorpions, and characterization of venoms and toxins to access the gaps that need to be filled in the knowledge of the scorpion species of medical importance from the Brazilian Amazon. A systematic search using the string words "Amazon" and "scorpion" was performed on 11 databases. No restriction on date, language or status of the publication was applied. Reports not related to the Brazilian Amazon were excluded. Therefore, 88 studies remained. It is shown that populations of scorpions of medical importance, even of the same species, may present significant toxic variations peculiar to some regions in the Brazilian Amazon, and commercial scorpion antivenoms were not able to shorten the intensity and duration of neurological manifestations in patients stung by T. silvestris, T. apiacas or T. obscurus. It is also highlighted that the toxins responsible for triggering these alterations have not been elucidated yet and this is a fruitful field for the development of more efficient antivenoms. Furthermore, the geographic distribution of scorpions of the genus Tityus in the Brazilian Amazon was revised and updated. The cumulative and detailed information provided in this review may help physicians and scientists interested in scorpionism in the Brazilian Amazon.(AU)

Phylogenetic and Morphological Analyses of Androctonus crassicuda from Khuzestan Province, Iran (Scorpiones: Buthidae).

The Androctonus crassicuda is the most diverse scorpion species in the family of Buthidae, which is endemic to Khuzestan province, Iran. Investigation of the relationship of species by means of a molecular study of specimens is one of the new approaches due to the limitations of the morphological approaches. In the current study, the analysis was based on 32 morphological characteristics of A. crassicuda native to southwest Iran. Moreover, the DNA sequencing of two mitochondrial markers, namely cytochrome oxidase subunit I and 12sRNA loci was performed, and the phylogenetic tree was constructed using maximum likelihood method with 1000 replications using MEGA software (version 7). Based on the results of the phylogenetic tree, A. crassicuda was classified into a monophyletic group. However, the genetic diversity of this species populations was not significant (0.001). The highest and lowest genetic distance of A. crassicuda was compared with the reports obtained in Urmia and west Azerbaijan, Iran. There was a clear divergence between the A. crassicuda isolated from northern and southern areas of Iran. This study showed the importance of geographical and climate features of the region and genetic distance among the populations. The phylogenetic analysis of Androctonus species from other regions showed the highest and lowest genetic distance with A. gonneti (Morocco) and A. amoreuxi (Portugal), respectively. The comparison of the morphological characteristics and morphometric results revealed that metasoma characteristics are important in the identification of A. crassicuda. The results of the analysis of the morphometric values of A. crassicuda were mainly compatible with the phylogenetic trees and supported the traditional morphological classification, thereby presenting a clearly definition of the genera of Androctonus species.

Characterization of the fluorescence intensity and color tonality in the exoskeleton of scorpions.

Scorpions are a mesodiverse and nocturnal group of arachnids inhabiting most biomes worldwide. Different species of scorpions have divergent adaptations to the substrate they live in, but most of them share an intriguing characteristic: their exoskeleton contains fluorophores that emit blue-greenish fluorescence under ultraviolet radiation. Although there are some reports in the literature on the study of fluorescence in scorpions, the biological functionality of this light emission is currently unknown and is under debate. In this work, the properties of emission from the scorpion's exoskeleton are studied by means of digitally processed photographs taken of living specimens under ultraviolet illumination and complemented with standard spectroscopic measurements of emission and excitation spectra. With the aim of identifying possible correlations between the fluorescence, the characteristics color of the exoskeleton and the biology of the scorpions, 4 families, 9 genera and 24 species were studied. Our results suggest that the intensity of fluorescence is heterogeneous throughout the scorpion's exoskeleton studied here in such a way that pedipalps and metasomal segments fluoresce more intensely than the mesosomal segments. The spectrum of fluorescence across species is practically identical, suggesting that the same fluorophores are present in their exoskeletons. However, the fluorescence intensity emitted by each species varies according to their characteristic color (associated with the exoskeleton optical reflectance). Since the coloration of the exoskeleton is determined by the concentration of melanin and other pigments according to the substrate where scorpions live in, we conclude by suggesting that fluorescence may correlate directly to the ecomorphotype of the scorpions.

Mouthpart homologies and life habits of Mesozoic long-proboscid scorpionflies.

Mesozoic long-proboscid scorpionflies (Mesopsychoidea) provide important clues to ancient plant-pollinator interactions. Among them, the family Aneuretopsychidae is especially important because its mouthparts are vital to deciphering the early evolution of Mesopsychoidea and putatively the origin of fleas (Siphonaptera). However, the identification of mouthpart homologs among Aneuretopsychidae remains controversial because of the lack of three-dimensional anatomical data. Here, we report the first Aneuretopsychidae from Late Cretaceous Burmese amber, which have short maxillary palpi and elongate mouthpart elements consisting of one pair of galeae and one hypopharynx. Their mouthparts are identical to those of Pseudopolycentropodidae (= Dualulidae, new synonym) but are not homologous to those of Siphonaptera. Our phylogenetic analysis provides robust evidence for the debated monophyly of Mesopsychoidea. Our results suggest that the long-proboscid condition has most likely evolved once in Mesopsychoidea, independently from fleas, and further reveal the variety and complexity of mid-Cretaceous pollinating insects.

First report of scorpionism caused by Tityus serrulatus, described by Lutz and Mello, 1922 (Scorpiones, Buthidae), a species non-native to the state of Pará, Brazilian Amazon.

This reports a case of scorpionism caused by Tityus serrulatus. A male adult was stung while unloading bananas at the supply center in Belém, Pará, Brazil. The bananas originated in another state (Bahia) and were brought to Belém by truck. The patient presented with pain, edema, and erythema at the sting site, and was classified as low-risk. The specimen was identified as T. serrulatus and symptomatic treatment and clinical observation were advised. The patient was discharged later without further complications. This is the first known envenomation caused by T. serrulatus, a non-native species to Pará, in the Brazilian Amazon.

Venom diversity in the Neotropical scorpion genus Tityus: Implications for antivenom design emerging from molecular and immunochemical analyses across endemic areas of scorpionism.

Scorpions of the Neotropical genus Tityus are responsible for most severe envenomations in the Caribbean, South America, and Lower Central America (LCA). Although Tityus is taxonomically complex, contains high toxin polymorphism, and produces variable clinical manifestations, treatment is limited to antivenoms produced against species with restricted distributions. In this study, we explored the compositional and antigenic diversity of Tityus venoms to provide improved guidelines for the use of available antivenoms at a broader geographic scale. We used immunoblotting, competitive ELISA, and in vivo studies to compare reactivity against commercial antivenoms from Brazil, Venezuela, and Mexico, as well as MALDI-TOF mass spectrometry, cDNA sequencing, and phylogenetic analyses to assess venom sodium channel-active toxin (NaTx) content from medically important Tityus populations inhabiting Brazil, Colombia, Costa Rica, Ecuador, Panama, Trinidad and Tobago, and Venezuela. Additionally, we raised rabbit antibodies against Tityus venoms from LCA to test for cross-reactivity with congeneric species. The results suggest that Tityus spp. possess high venom antigenic diversity, underlying the existence of four toxinological regions in Tropical America, based on venom composition and immunochemical criteria: LCA/Colombia/Amazonia (Region I), Venezuela (Region II), southeast South America (Region III), and a fourth region encompassing species related to toxinologically divergent Tityus cerroazul. Importantly, our molecular and cross-reactivity results highlight the need for new antivenoms against species inhabiting Region I, where scorpions may produce venoms that are not significantly reactive against available antivenoms.

First report of scorpionism caused by Tityus serrulatus, described by Lutz and Mello, 1922 (Scorpiones, Buthidae), a species non-native to the state of Pará, Brazilian Amazon

Abstract This reports a case of scorpionism caused by Tityus serrulatus. A male adult was stung while unloading bananas at the supply center in Belém, Pará, Brazil. The bananas originated in another state (Bahia) and were brought to Belém by truck. The patient presented with pain, edema, and erythema at the sting site, and was classified as low-risk. The specimen was identified as T. serrulatus and symptomatic treatment and clinical observation were advised. The patient was discharged later without further complications. This is the first known envenomation caused by T. serrulatus, a non-native species to Pará, in the Brazilian Amazon.

Expansión de la distribución de escorpiones del género Tityus C. L. Koch 1836 en Argentina: Implicancias sanitarias / Expansion of the distribution of scorpions of the genus Tityus C. L. Koch 1836 in Argentina: Implications in public health

Se ha observado la presencia de especies de Tityus en diferentes regiones del país, en las cuales su presencia no había sido comunicada previamente: 1- Tityus bahiensis en las provincias de Entre Ríos y Buenos Aires, en esta última en la localidad de Lanús y en San Clemente del Tuyú, y 2- Tityus confluens en la Ciudad Autónoma de Buenos Aires (CABA) y en la provincia de Buenos Aires en las localidades de Pilar, La Plata, Mar del Plata y Bahía Blanca. Estos hallazgos modifican el mapa de la distribución de escorpiones de importancia sanitaria en Argentina por lo que ante la picadura de escorpiones deben considerarse estos nuevos hallazgos. Esto es especialmente importante en el ámbito de la CABA y la provincia de Buenos Aires, en donde la enorme mayoría de los accidentes por escorpiones han sido causados por T. trivittatus y en donde ahora, al menos en algunas de sus regiones se pueden encontrar T. confluens y T. bahiensis. Se discuten posibles razones de esta nueva distribución así como la ocurrencia de accidentes graves en zonas donde no ocurrían históricamente y de sus posibles causas. En base a los casos graves producidos en los últimos tiempos y a este nuevo mapa de distribución, se hace énfasis en la necesidad de capacitación al personal de salud en general y de los médicos de guardia y terapistas en particular, para tratar adecuadamente los accidentes por escorpiones.

Several species of Tityus have been described in regions of Argentina where their presence had not been previously described. These are: 1- Tityus bahiensis in the provinces of Entre Ríos and Buenos Aires (in the localities of Lanús and San Clemente del Tuyú), and 2- Tityus confluens in the city of Buenos Aires and in the province of Buenos Aires in the localities of Pilar, La Plata, Mar del Plata and Bahía Blanca. These findings modify the distribution map of scorpions of sanitary importance in Argentina, reason for which this new distribution must be considered when facing a scorpion sting. This is especially important in the city of Buenos Aires and the province of Buenos Aires, where most of the accidents by scorpions are caused by Tityus trivittatus, and where at least in some of their regions, T. confluens or T. bahiensis can be found at present. The possible reasons of this new distribution, as well as the possible causes for the occurrence of severe envenomations in regions where these were not observed historically, are discussed. Based on the severe envenomations observed and on this new distribution map, emphasis is placed on the need to capacitate health personnel in general and intensivists or critical care physicians in particular to adequately treat scorpion accidents.

A new species of Centruroides (Scorpiones, Buthidae) from Colima, Mexico.

As part of an ongoing survey of scorpion diversity in Colima, Mexico, the isolated mountain Cerro Grande, part of the Biosphere Reserve Sierra de Manantlán, was investigated. Centruroides possanii sp. nov., the fifth species of the genus from the state, was discovered during fieldwork in the massif and is described in the present paper. Physiographical and climatic features of Cerro Grande may restrict the range of this new species; thus, we hypothesized that it may be a microendemic species that requires priority conservation. The new species is not assigned to any Centruroides species group recognized because some of its morphological features do not fit the current diagnosis of any of these groups, and these different groups are non-monophyletic and consequently ill-diagnosed. The new species is profusely illustrated, particularly the hemispermatophore. A distribution map is presented along with the other two more common species distributed in Colima. Because only indirect data on the potency of its venom is available, the medical importance of this new species described here is yet to be known.

Hadrurid Scorpion Toxins: Evolutionary Conservation and Selective Pressures.

Scorpion toxins are thought to have originated from ancestral housekeeping genes that underwent diversification and neofunctionalization, as a result of positive selection. Our understanding of the evolutionary origin of these peptides is hindered by the patchiness of existing taxonomic sampling. While recent studies have shown phylogenetic inertia in some scorpion toxins at higher systematic levels, evolutionary dynamics of toxins among closely related taxa remain unexplored. In this study, we used new and previously published transcriptomic resources to assess evolutionary relationships of closely related scorpions from the family Hadruridae and their toxins. In addition, we surveyed the incidence of scorpine-like peptides (SLP, a type of potassium channel toxin), which were previously known from 21 scorpion species. We demonstrate that scorpine-like peptides exhibit gene duplications. Our molecular analyses demonstrate that only eight sites of two SLP copies found in scorpions are evolving under positive selection, with more sites evolving under negative selection, in contrast to previous findings. These results show evolutionary conservation in toxin diversity at shallow taxonomic scale.

Perspectives and recommendations towards evidence-based health care for scorpion sting envenoming in the Brazilian Amazon: A comprehensive review.

Although underreported across the Amazon region, scorpion stings are very prevalent in some areas and can be potentially life-threatening, especially in children. The most vulnerable populations are those living in locations far from the capitals, hence having limited access to the health system where the appropriate structure for the treatment of severe cases is found. An abundant and diverse fauna of scorpions is found in the region, but few studies have been conducted to decipher the clinical characteristics and therapeutic response of the available antivenoms in envenomings caused by the various species. Antivenom underdosage as well as delayed medical assistance are common among indigenous populations, resulting in poor outcome rates. An in depth understanding of the epidemiological, clinical and therapeutic aspects of scorpion sting envenomings in the Amazon is necessary to improve the outcome of these cases.