Scorpions and life-history strategies: from evolutionary dynamics toward the scorpionism problem

This work aims to contribute to the general information on scorpion reproductive patterns in general including species that can be noxious to humans. Scorpions are unusual among terrestrial arthropods in several of their life-history traits since in many aspects their reproductive strategies are more similar to those of superior vertebrates than to those of arthropods in general. This communication focuses mainly on the aspects concerning embryonic and post-embryonic developments since these are quite peculiar in scorpions and can be directly connected to the scorpionism problem. As in previous similar contributions, the content of this communication is addressed mainly to non-specialists whose research embraces scorpions in several fields such as venom toxins and public health. A precise knowledge of reproductive strategies presented by several scorpion groups and, in particular, those of dangerous species may prove to be a useful tool in the interpretation of results dealing with scorpionism, and also lead to a better treatment of the problems caused by infamous scorpions.(AU)

Segmentation and tagmosis in Chelicerata.

Patterns of segmentation and tagmosis are reviewed for Chelicerata. Depending on the outgroup, chelicerate origins are either among taxa with an anterior tagma of six somites, or taxa in which the appendages of somite I became increasingly raptorial. All Chelicerata have appendage I as a chelate or clasp-knife chelicera. The basic trend has obviously been to consolidate food-gathering and walking limbs as a prosoma and respiratory appendages on the opisthosoma. However, the boundary of the prosoma is debatable in that some taxa have functionally incorporated somite VII and/or its appendages into the prosoma. Euchelicerata can be defined on having plate-like opisthosomal appendages, further modified within Arachnida. Total somite counts for Chelicerata range from a maximum of nineteen in groups like Scorpiones and the extinct Eurypterida down to seven in modern Pycnogonida. Mites may also show reduced somite counts, but reconstructing segmentation in these animals remains challenging. Several innovations relating to tagmosis or the appendages borne on particular somites are summarised here as putative apomorphies of individual higher taxa. We also present our observations within the concept of pseudotagma, whereby the true tagmata - the prosoma and opisthosoma - can be defined on a fundamental change in the limb series while pseudotagmata, such as the cephalosoma/proterosoma, are expressed as divisions in sclerites covering the body without an accompanying change in the appendages.

Scorpions and scorpionism in Iran's central desert.

Venomous scorpions have extreme importance in field of medicine and public health. This descriptive - analytic study was done to identify scorpion fauna, their ecological aspects as well as scorpionism for risk management and prevention of this health problem in Iran's central desert. Four urban and fifteen rural areas with various climates and topography locations were selected for monthly scorpion collection through a randomly cluster sampling in 2013. The clinical data was obtained from questionnaires provided in 2009-2014. Totally, 1481 scorpion sting cases were recorded. The majority were treated less than 6h after the sting. Statistical tests showed significant difference between season, scorpion's color, living place of patients and scorpionism cases. Plain areas had the most occurrence of scorpionism followed by foothills. Moreover, 311 scorpion samples belonged to 7 species of Buthidae were collected. Mesobuthus eupeus was the dominant species in both rural and urban areas. Most of the collected samples were from indoors, yards and around the houses. The most scorpion activity was recorded in the summer. The studied areas had rich scorpion fauna due to various climates and topography locations. Scorpion stings can be important and fatal in this area, particularly in the plain regions with semi-desert climate. An investigation for assessment of peoples' awareness on prevention methods of scorpionism and also the determination and the assessment of effective factors on reducing the elapsed time between scorpion stings and receiving medical care are here recommended.

Sex-Linked Chromosome Heterozygosity in Males of Tityus confluens (Buthidae): A Clue about the Presence of Sex Chromosomes in Scorpions.

Scorpions of the genus Tityus show holokinetic chromosomes, achiasmatic male meiosis and an absence of heteromorphic sex chromosomes, like all Buthidae. In this work, we analysed the meiotic behaviour and chromosome rearrangements of a population of the scorpion Tityus confluens, characterising the cytotypes of males, females and embryos with different cytogenetic techniques. This revealed that all the females were structural homozygotes, while all the males were structural heterozygotes for different chromosome rearrangements. Four different cytotypes were described in males, which differed in chromosome number (2n = 5 and 2n = 6) and meiotic multivalent configurations (chains of four, five and six chromosomes). Based on a detailed mitotic and meiotic analysis, we propose a sequence of chromosome rearrangements that could give rise to each cytotype and in which fusions have played a major role. Based on the comparison of males, females and a brood of embryos, we also propose that the presence of multivalents in males and homologous pairs in females could be associated with the presence of cryptic sex chromosomes, with the male being the heterogametic sex. We propose that the ancestral karyotype of this species could have had homomorphic XY/XX (male/female) sex chromosomes and a fusion could have occurred between the Y chromosome and an autosome.

Comprehensive analysis of venom from the scorpion Centruroides tecomanus reveals compounds with antimicrobial, cytotoxic, and insecticidal activities.

Centruroides tecomanus is a medically important scorpion of the state of Colima (Mexico). This communication reports the identification of venom components of this scorpion with biological activity over insects/crickets (Acheta domestica), crustaceans/fresh water shrimps (Cambarellus montezumae), and mammalians/mice (Mus musculus, strain CD1). It also describes the pharmacological effects on cell lines in culture (L5178Y cells, HeLa cells, HuTu cells and Jurkat E6-1 cells), as well as on several types of bacteria (see below). The soluble venom of this scorpion was fractionated by high-performance liquid chromatography (HPLC) and collected separately in twelve independent fractions collected over 60 min run (5 min time apart each other). The HPLC components of fraction VII were lethal to all three species used for assay. The IVth fraction had a toxic effect on freshwater shrimps. In this species, fractions VI, VII and VIII were all lethal. For crickets, fractions V and VI were toxic and fraction VII was lethal. In mouse, the lethal components were found in fraction VII, whereas fraction VIII was toxic, but not lethal, at the doses assayed. The molecular weight of peptides from the various group of fractions were identified by mass spectrometry determination. Components lethal to mice showed molecular weights from 7013 to 7487 Da. Two peptides were obtained in homogeneous form and shown to be lethal to the three species of animal used for assay. The soluble venom tested on L5178Y cell line survival was shown to be cytotoxic, at 10-100 µg/mL concentration, when compared to control murine splenocytes (p = 0.007). The soluble venom applied to Hela, Hutu and Jurkat cell lines did not show cytotoxic effects at these concentrations. On the contrary, it seems to have a proliferative effect. However the HPLC fractions I, III, VI and XII do have a cytotoxic effect on Jurkat E06-1 cells in culture at 200 µg/mL concentration. The antimicrobial activity of the venom fractions on Staphylococcus aureus (gram-positive), Escherichia coli, Pseudomonas aeruginosa y Salmonella spp (gram-negative) was measured, using the liquid inhibition growth system. The four strains of bacteria used were susceptible to fractions III and IV, affecting all four bacterial strains at concentrations below 5 µg/mL.

Insights into reproductive strategies of Tityus (Archaeotityus) pusillus Pocock, 1893 (Scorpiones, Buthidae).

A remarkable diversity of life history strategies has evolved among species for achieving reproductive success, including adaptive growth, protandry, iteroparity, and extra molting. Here, we report on the reproductive strategies of the litter-dwelling scorpion, Tityus (Archaeotityus) pusillus, the most abundant and widespread scorpion species in the Atlantic Forest of northeastern Brazil. We observed both iteroparity and protandry reproductive strategies in this species. Females were competent to produce up to three broods after a single insemination, and no correlation between female size and litter size was observed. Most males reached adulthood 1 month before females following four molts, characterizing protandry. Nevertheless, an extra molt was observed to occur in some males (n=4) and females (n=1). These findings highlight the life history traits of T. (A.) pusillus, which may imply in reproductive success and adaptation to changes in environmental conditions.

Scorpionism by Tityus silvestris in eastern Brazilian Amazon

Scorpionism is a serious public health problem in Brazil. Although cases of envenomation by scorpions are frequent in Brazil, Tityus silvestris found throughout the Amazon region is considered of minor medical significance and with only a few descriptions in the literature. This article aims to describe for the first time the epidemiological characteristics and clinical manifestations of scorpion stings by T. silvestris that occurred in eastern Brazilian Amazon. Methods A prospective and observational study was carried out on 13 confirmed cases of T. silvestris envenomation registered from 2007 to 2011 in the cities of Belém and Ananindeua, Pará state, Brazil. Results The stings occurred mainly during daytime, at domiciliary environment, and the scorpions were found in clothing, fruits or vegetables. Envenomation was more frequent in the age group between 21 and 30 years old, upper limbs were more affected and medical aid was usually provided within two hours. Men and women were equally affected. Regarding severity, ten patients were classified as Class I and three patients as Class II according to the Scorpion Consensus Expert Group. Local manifestations were present in all patients, being pain the most common symptom. Mild systemic manifestations including nausea, vomiting, somnolence, malaise and prostration were observed in three victims. Symptomatic treatment of pain was offered to all patients, and only one received specific antivenom. All victims had a favorable outcome. Conclusions To the best of our knowledge, this study is the first to report the systemic symptomatology of envenomation by T. silvestris in the Brazilian Amazon, highlighting the medical relevance of the species in this region. Further research on the venom and clinical manifestations of envenomation by T. silvestris should be conducted in order to verify the relevance of this species to public health.

Scorpion incidents, misidentification cases and possible implications for the final interpretation of results

The aim of this contribution is to bring general information on the classification and in particular on the specific identification of scorpion species dangerous to humans. Several generic groups are taken into consideration, but the Neotropical genus Tityus C. L. Koch, 1836 is used as a major example. The content of this paper is mostly addressed to non-specialists whose research embraces scorpions in several fields such as venom toxins and public health. Although efforts have been made in the last 20 years to create better links between true scorpion experts and non-specialists who use scorpions in their research, such exchanges had never led to a consensus among those different branches of biological and medical research. Consequently, many cases of species misidentification and even more serious errors concerning scorpion classification/identification are often present in the specialized literature. In conclusion, it is suggested here that the frequent cases of misidentification observed in several reports may induce mistakes in the final interpretation of results, leading only to more inefficacity in the treatment of problems caused by infamous scorpion species.

Two new species of Euscorpius (Scorpiones: Euscorpiidae) from Bulgaria, Serbia, and Greece.

Two new species of Euscorpius Thorell, 1876 (subgenus Euscorpius s.str.) (Scorpiones: Euscorpiidae) are described based on morphology and the COI DNA barcoding marker: E. deltshevi sp. n. from northern Bulgaria and neighbouring Serbia (formerly reported as E. carpathicus) and E. solegladi sp. n. from southwestern Bulgaria and neighbouring Greece (formerly reported as E. hadzii).

[Study on artificial compound feed for Buthus martensii].

OBJECTIVE: To solve the problem that the single feed causing malnutrition, extension of the life cycle and low survival rates of Buthus martensii. METHODS: By using Minitab (R) 15.1.1.0.0., 7 different kinds of compound feed were designed, including minced meat mud (pork,chicken and rabbit), bran (fried yellow), sugar, milk, vegetable paste and multivitamin as raw material. RESULTS: Different proportions of compound feed that taking the yellow mealworm as main component had a significant effect on the growth, mortality and birth number of Buthus martensii. Compound feed 5 significantly reduced mortality of youth scorpion. The compound feed 4 was the best in improving the weight of scorpion larvae and youth scorpion, and the farrowing number of mother scorpion. Other indicators were also good. Proportions in meatmud (pork, chicken and rabbit), bran (fried yellow), sugar, milk, vegetable paste and multivitamin was 30.00%, 25.00%, 20.08%, 15.58%, 8.08% and 1.25%, respectively. CONCLUSION: The growth of Buthus martensii is significantly influenced by the type of feed. In the production of compound feed, the yellow mealworm with compound feed 4 can be popularized.

Differentiation of somatic cells in the ovariuteri of the apoikogenic scorpion Euscorpius italicus (Chelicerata, Scorpiones, Euscorpiidae).

In apoikogenic scorpions, growing oocytes protrude from the gonad (ovariuterus) and develop in follicles exposed to the mesosomal (i.e. hemocoelic) cavity. During subsequent stages of oogenesis (previtellogenesis and vitellogenesis), the follicles are connected to the gonad surface by prominent somatic stalks. The aim of our study was to analyze the origin, structure and functioning of somatic cells accompanying protruding oocytes. We show that these cells differentiate into two morphologically distinct subpopulations: the follicular cells and stalk cells. The follicular cells gather on the hemocoelic (i.e. facing the hemocoel) surface of the oocyte, where they constitute a cuboidal epithelium. The arrangement of the follicular cells on the oocyte surface is not uniform; moreover, the actin cytoskeleton of these cells undergoes significant modifications during oocyte growth. During initial stages of the stalk formation the stalk cells elongate and form F-actin rich cytoplasmic processes by which the stalk cells are tightly connected to each other. Additionally, the stalk cells develop microvilli directed towards the growing oocyte. Our findings indicate that the follicular cells covering hemocoelic surfaces of the oocyte and the stalk cells represent two distinct subpopulations of epithelial cells, which differ in morphology, behavior and function.

[Study on spatial distribution pattern and seasonal distribution of Buthus martensii in yan'an].

OBJECTIVE: To study population dynamics of Buthus martensii and its wild spatial distribution pattern in Yan'an. METHODS: Every month, the 1st, 10th and 20th day were designated to collect Buthus martensii samples and count on the collection by using expansion pattern target, Taylor power law and Lwao m-x regression analysis. RESULTS: Taylor power law regression equation was Ig S2 = 1.6671 lg x + 0.1554; Lwao m-x regression equation was m = 1.3179 + 1.5133x. The spatial distribution pattern of Buthus martensii was negative binomial distribution and its population fluctuation curve in the wild presented a single peak. CONCLUSION: Spatial pattern of Buthus martensii in Yan'an is aggregated distribution which based on individual group, and the higher density,the greater aggregation degree.

Effects of in utero exposure to Tityus bahiensis scorpion venom in adult rats

The toxicity of Tityus bahiensis scorpion venom is well known, but there are little data about the damage in offspring of dams that were exposed to the venom during pregnancy. The objective of this work was to determine the toxic effects of venom in adult offspring of Wistar rats exposed to venom in utero. Dams were divided into a control group, subcutaneously injected with saline solution on the 10th (GD10) and 16th (GD16) days, and two experimental groups, subcutaneously injected with venom (2.5 mg/kg) on GD10 or GD16, respectively. Adult offspring were evaluated according to behavioral development and neuronal integrity in the hippocampus. Tests performed in the activity box and in the enriched environment demonstrated that males from GD10 had motor decrease. Females from GD10 showed a depressive-like state and were more anxious, as demonstrated by the forced swimming test and social interaction. The plus-maze discriminative avoidance task demonstrated that GD16 males had lower levels of anxiety. The number of neuronal cells was decreased in CA1, CA3 and CA4 hippocampal areas of males and females from GD10 group and in CA1 of females and CA4 of males from GD16 group. Thus, we conclude that venom exposure in pregnant dams causes subtle alteration in the behavioral and neuronal development of offspring in adult life in a gender-dependent manner.

Observations on the life history of Chaerilus philippinus Lourenço & Ythier, 2008 (Scorpiones, Chaerilidae) from the Philippines.

Biological observations on Chaerilus philippinus were based on specimens from the region of Appari, North of Luzon in the Philippines. The total duration of embryonic development was estimated as being between 110 to 136 days, while the moults between successive juvenile instars and adulthood took place at ages that averaged 7, 39, 73, 190 and 327 days. These developmental periods are shorter and different from those previously observed among species of non-buthid scorpions. They prove to be rather similar to those observed in buthid scorpions, however. Morphometric growth values of the different instars are similar or smaller than those of other species of scorpions that have been studied. Aspects of maternal care and social behaviour are also commented.

Development of respiratory structures in embryos and first and second instars of the bark scorpion, Centruroides gracilis (Scorpiones: Buthidae).

The SEM was used to study the development of respiratory structures in successive stages in relation to the overall changes occurring in the scorpions. Book lung development is a slow process, starting with spiracles and a sac-like atrium in the early embryo and continuing lamellar formation to 150 or more in the adult. In the embryo, the primordial epithelial cells become aligned in a planar pattern as they secrete granules of material that aggregate spontaneously to form the cuticular walls of the lamellae. A blade-like structure is formed consisting of cells sandwiched within the two cuticle walls they secreted. These cells are in the primordial air channel. The adjacent hemolymph channel is nearly devoid of cells, but cross-bridges develop and help stabilize the cuticle walls and maintain the width of the channel. The cells in the primordial air channel undergo cytolysis, leaving it open for air except for cuticular cross-bridges. Development continues in the newborn (first instars); the air channels of some lamellae still contain cells and are not yet functional for gas exchange. The first instars are weak and relatively inactive. They climb up on the mother's dorsum until the first molt (about 8 days). With the cuticular walls of the lamellae in place, cells adhering to the wall in the hemolymph channel produce a thin, new tissue layer (epithelium) on the lamellar wall facing the hemolymph channel. This layer has many discontinuities as though it is slowly developing. Formation of the tissue layer and cytolysis of the cells in the air channels continue through the first molt in which little book lung cuticle is shed as exuvium. The air channels of the second instars (foraging nymphs) are now cell free and open for air passage except for the cross-bridges. The tissue layer is still incomplete and continues to be formed. It may provide the hypodermal primordium for cuticle replacement in later molts, but development was not studied beyond the second instar except for comparison with book lungs in the adult. The blade-like lamellae in the adult are larger and more numerous than in the second instar, but in the anterior book lung the shape of the cuticle wall and cross-bridges and the widths of the air and hemolymph channels are about the same as in the second instar. The air channels in the posterior part of the lamellae have distinctive, vein-like space-holders. The similarity of the adult anterior lamellae with those in the second instar suggests retention of this part through the 4-5 molts to maturation, and/or cell processes like those in the embryo are repeated, but this needs to be examined in further studies of cell and cuticle changes before and during the molts.

Epidemiological and clinical aspects of scorpionism by Tityus trivittatus in Argentina.

This is a descriptive study of epidemiological and clinical aspects of stings caused by the scorpion Tityus trivittatus in Argentina. We analyzed 511 cases recorded from different health centers in 22 provinces. Most accidents took place during the period November-April (76%), in or nearby houses (86%). Over 50% of the accidents involved children and teenagers. Envenomation by T. trivittatus was mainly characterized by local symptoms: pain (85%), edema (26.6%), burning sensation (24.7%), erythema (20.7%), local hyperthermia (13.1%), paresthesia (9.8%) and general manifestations such as vomiting (25%), paleness (18.8%), headache (11.4%) and sweating (8.2%). Neurological, cardiovascular and respiratory disorders were uncommon. Almost 90% of the injured people got treated with specific antivenom within 2 h, 6% were treated 2-h after the accident and only 2 people were treated 12 or more hours after being stung. The global mortality recorded was 6 per 1000 cases. Scorpionism in Argentina is a public health problem under control due to the relatively low incidence and the accessibility of specific antivenom.