Late Ordovician eurypterid preserves oldest euchelicerate musculature in pyrite.

Pyritization of soft tissues of invertebrates is rare in the fossil record. In New York State, it occurs in black shales of the Lorraine Group (Late Ordovician), the best-known example of which is Beecher's Trilobite Bed. Exceptional preservation at the quarry where this bed is exposed allowed detailed examination of trilobite and ostracod soft-tissue anatomy. Here, we present the first example of a eurypterid (sea scorpion) currently ascribed to Carcinosomatidae from this deposit that also preserves the first evidence for mesosomal musculature in eurypterids. This specimen demonstrates that eurypterid musculature can be preserved in pyrite and evidences the oldest example of euchelicerate muscles within the fossil record. Sulfur isotope data illustrate that pyrite rapidly replicated muscle tissue in the early burial environment, prior to the pyritization of biomineralized exoskeleton and cuticular trilobite limbs. This discovery therefore expands the limited fossil record of euchelicerate musculature, while extending the taphonomic scope for preservation of detailed internal structures, more broadly, within arthropods.

Scorpions trapped in amber: a remarkable window on their evolution over time from the Mesozoic period to present days

This synoptic review aims to bring some general information on fossil scorpions, namely those trapped in amber - fossilized resin - ranging from Lower Cretaceous through the Palaeocene and up to the Miocene. The question to be addressed is how the study of these fossils can be connected with possible present scorpionism problems. A precise knowledge of these ancient lineages provides information about the evolution of extant lineages, including the buthoids, which contain most known noxious species. Among the Arthropods found trapped in amber, scorpions are considered rare. A limited number of elements have been described from the Late Tertiary Dominican and Mexican amber, while the most ancient Tertiary amber from the Baltic region produced more consistent results in the last 30 years, primarily focusing on a single limited lineage. Contrarily, the Cretaceous amber from Myanmar, also called Burmite, has yielded and continues to yield a significant number of results represented by several distinct lineages, which attest to the considerable degree of diversity that existed in the Burmese amber-producing forests. As in my previous similar contributions to this journal, the content of this note is primarily addressed to non-specialists whose research embraces scorpions in various fields such as venom toxins and public health. An overview knowledge of at least some fossil lineages can eventually help to clarify why some extant elements associated with the buthoids represent dangerous species while others are not noxious.(AU)

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.

Air Breathing in an Exceptionally Preserved 340-Million-Year-Old Sea Scorpion.

Arachnids are the second most successful terrestrial animal group after insects [1] and were one of the first arthropod clades to successfully invade land [2]. Fossil evidence for this transition is limited, with the majority of arachnid clades first appearing in the terrestrial fossil record. Furthermore, molecular clock dating has suggested a Cambrian-Ordovician terrestrialization event for arachnids [3], some 60 Ma before their first fossils in the Silurian, although these estimates assume that arachnids evolved from a fully aquatic ancestor. Eurypterids, the sister clade to terrestrial arachnids [4-6], are known to have undergone major macroecological shifts in transitioning from marine to freshwater environments during the Devonian [7, 8]. Discoveries of apparently subaerial eurypterid trackways [9, 10] have led to the suggestion that eurypterids were even able to venture on land and possibly breathe air [11]. However, modern horseshoe crabs undertake amphibious excursions onto land to reproduce [12], rendering trace fossil evidence alone inconclusive. Here, we present details of the respiratory organs of Adelophthalmus pyrrhae sp. nov. from the Carboniferous of Montagne Noire, France [13], revealed through micro computed tomography (µ-CT) imaging. Pillar-like trabeculae on the dorsal surface of each gill lamella indicate eurypterids were capable of subaerial breathing, suggesting that book gills are the direct precursors to book lungs while vascular ancillary respiratory structures known as Kiemenplatten represent novel air-breathing structures. The discovery of air-breathing structures in eurypterids indicates that characters permitting terrestrialization accrued in the arachnid stem lineage and suggests the Cambrian-Ordovician ancestor of arachnids would also have been semi-terrestrial.

Insights into the 400 million-year-old eyes of giant sea scorpions (Eurypterida) suggest the structure of Palaeozoic compound eyes.

Sea scorpions (Eurypterida, Chelicerata) of the Lower Devonian (~400 Mya) lived as large, aquatic predators. The structure of modern chelicerate eyes is very different from that of mandibulate compound eyes [Mandibulata: Crustacea and Tracheata (Hexapoda, such as insects, and Myriapoda)]. Here we show that the visual system of Lower Devonian (~400 Mya) eurypterids closely matches that of xiphosurans (Xiphosura, Chelicerata). Modern representatives of this group, the horseshoe crabs (Limulidae), have cuticular lens cylinders and usually also an eccentric cell in their sensory apparatus. This strongly suggests that the xiphosuran/eurypterid compound eye is a plesiomorphic structure with respect to the Chelicerata, and probably ancestral to that of Euchelicerata, including Eurypterida, Arachnida and Xiphosura. This is supported by the fact that some Palaeozoic scorpions also possessed compound eyes similar to those of eurypterids. Accordingly, edge enhancement (lateral inhibition), organised by the eccentric cell, most useful in scattered light-conditions, may be a very old mechanism, while the single-lens system of arachnids is possibly an adaptation to a terrestrial life-style.

The evolution and distribution of noxious species of scorpions (Arachnida: Scorpiones)

This contribution attempts to bring some general information on the evolution and, in particular, on the geographic distribution of scorpion species noxious to humans. Since 95% of the scorpions incidents are generated by specimens of the family Buthidae C. L. Koch, the analysis will be limited to this familial group. As in previous similar contributions, the content of this work is mostly addressed to non-specialists whose research embraces scorpions in several fields such as venom toxins and public health. Only in recent years, efforts have been made to create better links between 'academic scorpion experts' and other academic non-specialists who use scorpions in their research. Even if a larger progress can yet be expected from such exchanges, crossed information proved to be useful in most fields of scorpion studies. Since the taxonomy of scorpions is complex, misidentifications and even more serious errors concerning scorpion classification/ identification are often present in the general literature. Consequently, a precise knowledge of the distribution patterns presented by many scorpion groups and, in particular, those of infamous species, proves to be a key point in the interpretation of final results, leading to a better treatment of the problems caused by infamous scorpion species.(AU)

Scorpionism by Hemiscorpius spp. in Iran: a review

Scorpions are distributed throughout Iran and the genus Hemiscorpius is particularly important in this region. Hemiscorpius lepturus is the most significant species within the genus in the country. Since scorpionism provoked by Hemiscorpius comprises a medical emergency, the present study is focused on this important issue. In order to perform the present work, a review of the medical and health-related literature was carried out in several databases. The current findings indicate that six species of Hemiscorpius are found in 15 states of Iran, mainly in the south and southwest. Deaths caused by stings were reported only for two species. The morphological characteristics and geographical distribution of H. lepturus in Iran, its venom and the toxic compounds, epidemiologic data and clinical manifestations of envenomation as well as treatment for affected people are herein reviewed and described. H. lepturus venom toxicity differs from other Iranian scorpions regarding duration and severity. Scorpionism is an important public health problem in Iran, especially in southwest and south regions and in urban areas. It is more prevalent in children and young people. H. lepturus venom is primarily a cytotoxic agent and has hemolytic, nephrotoxic and to some extent hepatotoxic activity. The use of polyvalent antivenom to prevent scorpion sting symptoms is recommended. A well-planned health education program might be useful in preventing scorpionism.(AU)

A checklist of the scorpions of Ecuador (Arachnida: Scorpiones), with notes on the distribution and medical significance of some species

Ecuador harbors one of the most diverse Neotropical scorpion faunas, hereby updated to 47 species contained within eight genera and five families, which inhabits the "Costa" (n= 17), "Sierra" (n= 34), "Oriente" (n= 16) and "Insular" (n= 2) biogeographical regions, corresponding to the western coastal, Andean, Amazonian, and the Galápagos archipelago regions, respectively. The genusTityus Koch, in the family Buthidae, responsible for severe/fatal accidents elsewhere in northern South America and the Amazonia, is represented in Ecuador by 16 species, including T. asthenes, which has caused fatalities in Colombia and Panama, and now in the Ecuadorian provinces of Morona Santiago and Sucumbíos. Underestimation of the medical significance of scorpion envenoming in Ecuador arises from the fact thatCentruroides margaritatus (Gervais) (family Buthidae) andTeuthraustes atramentarius Simon (family Chactidae), whose venoms show low toxicity towards vertebrates, frequently envenom humans in the highly populated Guayas and Pichincha provinces. This work also updates the local scorpion faunal endemicity (74.5 %) and its geographical distribution, and reviews available medical/biochemical information on each species in the light of the increasing problem of scorpionism in the country. A proposal is hereby put forward to classify the Ecuadorian scorpions based on their potential medical importance.(AU)

Second record of Tityus bahiensis (Scorpiones, Buthidae) from Venezuela: epidemiological implications

This work reports the second record of the scorpion Tityus bahiensis Perty from Venezuela. The specimen was found alive in a wardrobe at a hotel resort in Margarita Island, northeastern Venezuela. Morphological characterization allowed its assignment to the Tityus bahiensis population inhabiting the southernmost area of the species' geographic range, e.g. the state of São Paulo in Brazil, northern Argentina and Paraguay. The fact that the only available Venezuelan antiscorpion (anti-Tityus discrepans) serum does not neutralize the effects of alpha- and beta-toxin from Tityus serrulatus venom (which resembles in composition that of T. bahiensis) constitutes a warning to local clinicians confronted with envenomations by noxious species transported to Venezuela from Brazil by human agency.

The organic preservation of fossil arthropods: an experimental study.

Modern arthropod cuticles consist of chitin fibres in a protein matrix, but those of fossil arthropods with an organic exoskeleton, particularly older than Tertiary, contain a dominant aliphatic component. This apparent contradiction was examined by subjecting modern cockroach, scorpion and shrimp cuticle to artificial maturation (350 degrees C/700 bars/24 h) following various chemical treatments, and analysing the products with pyrolysis-gas chromatography/mass spectrometry (Py-GC/MS). Analysis of artificially matured untreated cuticle yielded moieties related to phenols and alkylated substituents, pyridines, pyrroles and possibly indenes (derived from chitin). n-Alkyl amides, C16 and C18 fatty acids and alkane/alk-1-ene homologues ranging from C9 to C19 were also generated, the last indicating the presence of an n-alkyl component, similar in composition to that encountered in fossil arthropods. Similar pyrolysates were obtained from matured pure C16 and C18 fatty acids. Py-GC/MS of cuticles matured after lipid extraction and hydrolysis did not yield any aliphatic polymer. This provides direct experimental evidence that lipids incorporated from the cuticle were the source of aliphatic polymer. This process of in situ polymerization appears to account for most of the fossil record of terrestrial arthropods as well as marine arthropods that lacked a biomineralized exoskeleton.

Nueva especie de Tityus (Scorpiones: Buthidae) del Turimiquire, Venezuela / A new species of Tityus (Scorpiones: Buthidae) from Turimiquire, Venezuela

We describe a new species of Buthid scorpion, Tityus quirogae, from the Turimiquire mountains, eastern side of the La Costa range, Monagas and Sucre States, Venezuela. It is morphologically similar to T. nematochirus and T. meridanus. It differs from these species in the distribution of the dorsal femoral trichobothria and in the following combination of characters: (1) I caudal segment with two parallel ventral keels. Caudal segments II to IV with ventral keels that are double in the extremes of the segments and single in the middle, (2) noticeable sexual dimorphism, especially in pedipalp shape; the male chelae are markedly thin and long, (3) number of pectineous teeth (male: 18 in each pecten; female: 18 in the right pecten and 19 in the left one), and (4) number of denticle rows in the movable finger of the right chela (16 in both sexes). T. quirogae is the first species of this genus, in the Northeastern region of Venezuela, with thin and elongated pedipalps.

Se describe una nueva especie de la familia Buthidae, Tityus quirogae, del Macizo del Turimiquire en el tramo oriental de la Cordillera de La Costa en los Estados Monagas y Sucre, Venezuela. Es similar, morfológicamente, a T. nematochirus y T. meridanus. Se distingue de estas especies por la disposición de las tricobotrias dorsales del fémur y por presentar la siguiente combinación de características: (1) I Segmento caudal con dos carenas ventrales paralelas. Segmentos caudales II al IV con carenas ventrales dobles hacia el área proximal que convergen y se hacen únicas, para finalmente ser divergentes hacia la región distal, (2) marcado dimorfismo sexual, especialmente en la forma de los pedipalpos; las quelas en los machos son marcadamente delgadas y largas, (3) número de dientes pectíneos (macho: 18 derechos y 18 izquierdos y hembra: 18 derechos y 19 izquierdos), (4) número de hileras de dentículos del dedo móvil de la mano derecha (macho 16 y hembra 16). T. quirogae es la primera especie del género con pedipalpos delgados y alargados para la región nororiental de Venezuela.

Characterization of a novel cDNA encoding a short venom peptide derived from venom gland of scorpion Buthus martensii Karsch: trans-splicing may play an important role in the diversification of scorpion venom peptides.

A novel cDNA clone (named BmKT-u) which is a hybrid molecule of the 5'-terminal region of BmKT' cDNA and the 3'-terminal region of an undocumented cDNA (named BmKu), was isolated from a cDNA library made from the venom gland of scorpion Buthus martensii Karsch. BmKT-u codes for a 30 amino acid residue precursor peptide composed of a 20-residue signal sequence, and a putative 10-residue novel mature peptide. Northern blot hybridization showed BmKT-u cDNA is generated from a transcript. RT-PCR experiments excluded the possibility that BmKT-u cDNA is an artifact generated during reverse transcription. Genomic amplifications performed with three pairs of BmKT-u gene-specific primers showed the BmKT-u gene does not exist in the genome of the scorpion as a single transcriptional unit. Genomic cloning for BmKT' showed that the BmKT' gene contains an intron of 509 bp inserted into the region encoding the C-terminal region of the signal peptide. A sequence alignment comparison of the cDNA of BmKT-u with genomic BmKT' revealed that the junction site of the hybrid molecule is located at the 5'-splicing site of the intron. The data suggest that the BmKT-u transcript is a naturally occurring mature mRNA that is generated by trans-splicing. Trans-splicing may contribute to the diversity of venom peptides from venomous animals.

Parametric values of Androctonus crassicauda (Oliver, 1807) (scorpiones: buthidae) from Turkey

The aim of the present study was to analyze the parametric values of Androctonus crassicauda (Oliver, 1807), the most significant scorpion species in Turkey. For this purpose, 11 male and 11 female A. crassicauda specimens were collected from Sanliurfa region. The body parts most important for species identification were measured by using sensitive calipers. Differences between sexes were statistically significant for pectinal organ length (female: 7.50±1.26mm;male: 8.63±1.05mm; p=0.033), pectinal tooth count (female: 25.36±0.86;male: 33.36±0.80; p=0.000), and carapace width (female: 10.27±1.19mm;male: 9.09±0.73mm; p=0.000). However, body length (female: 90.09±14.74mm;male: 82.95±5.40mm), abdomen width (female: 12.09±1.90mm;male: 11.09±0.88mm), and metasoma length (female: 43.09±3.08mm;male: 44.63±6.50mm) were not significantly different between sexes.(AU)

The first report of Tityus (Scorpiones: Buthidae) in anzoátegui state, Venezuela. A new species

Tityus gonzalespongai n. sp. is a species endemic to the high mountains of Anzoátegui State, Venezuela. It is found between 1,600 and 2,200 m in "La Laguna" mountain. Its habitat includes the area of "Bosque Húmedo Montano Bajo" with the type of vegetation of "Bosque Ombrófilo Montano Siempreverde" ("Bosques Nublados Costeros" that includes the "Subpáramos Arbustivos"). It is distinguished from other Tityus species (T. caripitensis, T. monaguensis, and T. nororientalis) by the following characteristics: 1- the disposition of the trichobothria in the pedipalps; 2- ventral keels of the caudal segments of the metasoma (segment II, double and parallel in the proximal two thirds, then convergent and finally divergent in the base; segments III and IV, double and parallel in the basal third, then convergent in a single keel that divides in the base); 3- the number of lines of denticles of the movable finger of the right pedipalp (male=14; female=14); 4- pectineous teeth (right/left: male=15/15: female=16/15); 5- color: movable and fixed fingers dark brown; prosoma and metasoma, ochre; caudal segment IV, slightly darker than the previous ones; V and the telson, dark brown. T. gonzalespongai belongs to the "androcottoides" group and presents a marked sexual dimorphism. It is the first species of the Tityus genus described and reported in Anzoátegui State, expanding this taxa distribution in Venezuela.

Complementos a la descripción de Tityus dedoslargos (Scorpiones: Buthidae) / Complements to the description of Tityus dedoslargos (Scorpiones: Buthidae)

The female of the Costarican scorpion Tityus dedoslargos Francke and Stockwell, 1987 is described for the first time and new data on its morphological variation and natural history are given. The female differs from the male mainly in having an enlarged basal median lamellae on the pectines, and lacking modified pedipalps. The species type locality relocated. This species is recorded from only two Costarican provinces: San Jose and Puntarenas.

Escorpiones (Arachnida) del ArchipiÚlago de Bocas del Toro, Panamá / Scorpions(Arachnida) from Bocas del Toro archipelago, Panamá

Four species of buthid scorpions (Ananteris platnicki Lourenço, 1993; Centruroides limbatus [Pocock, 1898]; Tityus pachyurus [Pocock, 1897]; and T. ocelote Francke and Stockwell, 1987) are recorded for the first time from some islands and cays of the Bocas del Toro Archipelago, Panama. Morphological variation and ecological data are given for all the species. This is the first Panamanian record for both C. limbatus and T. ocelote. The scorpion fauna of the Bocas de Toro Archipelago is related with the recent fauna of the Atlantic continental lowland region of Costa Rica and Panama and includes Amazonian-Guyanese (genera Ananteris and Tityus) as well as Mexican-North Central American (genus Centruroides) elements.

Venezuelan arachnids. Two new species of the Tityus Genus (Scorpionida: Buthidae) and the chromatographic profile of venom as a possible taxonomical tool

Two new species of the Tityus genus are described. T. isabelceciliae n. sp lives on the northern central slope of the Cordillera de la Costa. It belongs to the discrepans group and is dangerous to man due to its high number, aggressive behavior, domiciliary habits, and high toxicity of its venom. T. isabelceciliae venom is similar to other Tityus in relation to the molecular weight range and the biological activity of its components. However, the proportions of each fraction in the venom pooled from many T. isabelceciliae differ from the proportions in other Tityus, indicating that these proportions may have a taxonomical value. The venom LD50 is 38.1 (36.3, 39.9) µg/g mouse (Death in 30 min, Dixon and Mood (14) sequential method, median and 95 per cent confidence interval, n=7). Venom production was 916 (625, 1213) µg protein per animal (n=38): females [944 (750, 1150) mg protein per animal, n=24] and males [824 (550, 112) mg protein per animal, n=14] did not differ in venom production (P>0.05). There was no correlation between animal total weight and venom production. T. rusmelyae n. sp. from the androcottoides group lives near the town of Humocaro Alto in the Lara State, Venezuela. The male specimens have clearly defined keels and granules. It differs from other species of this genus in that the prominent characteristics are observed in male specimens.

Venezuelan arachnides: two new species of the Tityus genus (Scorpionida: Buthidae) in the State of Lara

Two new scorpion species of the Buthidae family, Tityus ivic-nancor and Tityus sanarensis, are described in this paper. Both species belong to the "androcottoides" group and live from 1300 to 1400 m. above sea level near the town of Sanare (69º42'17.8W, 9º45'3.9"N) in the southern part of the state of Lara in Venezuela. The number of Tityus species already knwon in Venezuela increases to 28 with the inclusion of these two new species. This paper includes a drescriptive diagnosis and also a differential diagnosis of the species of Tityus that are geographically related to T. ivic-nancor and T. sanarensis. Drawings of the significant morphological characteristics of male and female specimens and a morphometric table are also included.