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The Genus Hadruroides Pocock, 1893 (Scorpiones: Iuridae), in Peru: New Records and Descriptions of Six New Species
Abstract
We review the taxonomy of the Hadruroides Pocock, 1893 (Iuridae: Caraboctoninae), scorpions of Peru, describe six new species from the north of the country, and report new records of other poorly known species. The description of these species raises to 16 the number of described species in the genus, 13 of which occur in Peru. Four species inhabit dry forest in northern Peru: H. charcasus (Karsch, 1879); H. chinchaysuyu, n. sp.; H. geckoi, n. sp.; H. leopardus Pocock, 1900. Three species occur in inter-Andean valleys along the Cordillera: H. bustamantei Ochoa and Chaparro, 2008; H. carinatus Pocock, 1900; H. mauryi Francke and Soleglad, 1980. Six species inhabit desert along the Pacific coast: H. aguilari Francke and Soleglad, 1980; H. graceae, n. sp.; H. juanchaparroi, n. sp.; H. lunatus (L. Koch, 1867); H. tishqu, n. sp.; H. vichayitos, n. sp. Most species of Hadruroides have restricted distributions, except H. charcasus and H. lunatus, which are apparently more widely distributed. We consider it necessary to reassess all previous records of the latter two species, because we suspect several are based on misidentifications.
... Species of Hadruroides (n = 22) are restricted to Ecuador, Peru, northern Chile, and several offshore islands (including the Galápagos), where they inhabit inter-Andean valleys, Pacific deserts, and dry forest habitats [32,81]. In Ecuador the genus is represented by seven species, recently split into two subgenera by Rossi [32]. ...
... Ecuadorians records of Hadruroides (Lourencoides) lunatus (Koch) [82] are suggested to be probable misidentifications of other Hadruroides species, which is also the case in reports of this species from northern Chile [81]. Records of Hadruroides (Lourencoides) leopardus Pocock from the city of Loja, province of Loja [18], are probably referencing Hadruroides (Lourencoides) udvardyi Lourenço [81]. ...
... Ecuadorians records of Hadruroides (Lourencoides) lunatus (Koch) [82] are suggested to be probable misidentifications of other Hadruroides species, which is also the case in reports of this species from northern Chile [81]. Records of Hadruroides (Lourencoides) leopardus Pocock from the city of Loja, province of Loja [18], are probably referencing Hadruroides (Lourencoides) udvardyi Lourenço [81]. ...
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 genus Tityus 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 that Centruroides margaritatus (Gervais) (family Buthidae) and Teuthraustes 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.
... Hadruroides (Pocock, 1893) is a scorpion genus included in the family Iuridae, subfamily Charaboctoninae. This genus comprises sixteen species and there members appear brown in color with darker stains and have median size of 80 mm (Ochoa and Prendini, 2010;Maury, 1975). Hadruroides scorpions have been reported in Bolivia, Chile, Colombia, Ecuador, Peru, and Venezuela (Mello-Leitão, 1945;Esquivel de Verde, 1968;Kinzelbach, 1973;Maury, 1975;Cekalovic, 1983;Sissom and Fet, 2000), but are actually restricted to Ecuador, Peru, northern Chile, and several offshore islands, including the Galápagos (Cekalovic, 1966;Maury, 1975;Francke and Soleglad, 1981). ...
... Species of Hadruroides inhabit inter-Andean valleys, Pacific desert, and dry forest habitats (Ochoa and Prendini, 2010). ...
This communication describes the general biochemical properties and some immunological characteristics of the venom from the Peruvian scorpion Hadruroides lunatus, which is the most medically relevant species in Peru. The soluble venom of this scorpion is toxic to mice, the LD₅₀ determined was 0.1 mg/kg and 21.55 mg/kg when the venom was injected intracranial or intraperitoneally, respectively. The soluble venom displayed proteolytic, hyaluronidasic, phospholipasic and cardiotoxic activities. High performance liquid chromatography of the soluble venom resulted in the separation of 20 fractions. Two peptides with phospholipasic activity were isolated to homogeneity and their molecular masses determined by mass spectrometry (MALDI TOF). Anti-H. lunatus venom sera were produced in rabbits. Western blotting analysis showed that most of the protein content of this venom is immunogenic. H. lunatus anti-venom displayed consistent cross-reactivity with venom antigens from the new World-scorpions Tityus serrulatus and Centruroides sculpturatus venoms; however, a weaker reactivity was observed against the venom antigens from the old World-scorpion Androctonus australis Hector.
... The hemispermatophores of Caraboctonus and Hadruroides were first illustrated in Francke & Soleglad [29] and Stockwell [4]. Ochoa & Prendini [118] later presented a revision of the Peruvian Hadruroides species, with illustrations of spermatophores for six of these taxa showing that the structure is extremely conservative within the genus. ...
Background
Insemination in scorpions is carried out by means of a partly sclerotized structure, the spermatophore, which is composed of two separate halves, the hemispermatophores. In most genera these reproductive structures can be used to differentiate species. However, many taxa such as the genus Euscorpius and the family Diplocentridae lack the morphological diversity observed in the copulatory organs of many other arthropods, rendering them useless for species level taxonomy. Such structural stasis, however, suggests that hemispermatophores have evolved relatively slowly and may thus provide a stronger phylogenetic signal for recognizing supra-generic ranks than previously thought. Based on the postulate that the phenotypic stability observed in some groups is the consequence of functional constraint, the most comprehensive comparative study of the male sexual apparatus to date was conducted for a complete reassessment of the morphology, phylogenetic value and hypotheses of homology of these structures.
Results
Hemispermatophores, pre- and post-insemination spermatophores, as well as the inherent mechanisms of insemination, were studied across the whole order, allowing the recognition and description of a series of five basic bauplans for the capsular region. For the most part, these patterns appear to be consistent within each major taxonomic group, but several cases of incongruence between spermatophore morphology and taxonomy raises questions about the monophyly of some clades. The Bothriuridae are traditionally regarded as a basal scorpionoid family. However, except for the genus Lisposoma, bothriurid hemispermatophores and spermatophores are morphologically more similar to those of the Chactoidea than to those of scorpionoids. On the other hand, the male copulatory structures of the hormurid clade (Hormiops (Hormurus + Liocheles)) are more akin to those of Diplocentridae and Heteroscorpionidae than to those of other hormurids.
Conclusions
Spermatophore capsular patterns appears to be congruent with a recent phylogeny of the order Scorpiones based on phylogenomic data that placed Bothriuridae outside of Scorpionoidea and Liocheles outside of Hormuridae, in contradicton with earlier phylogenetic reconstructions based on morphology. This raises questions about the potential use of functionally constrained traits to assess the reliability of contradicting phylogenetic hypotheses and emphasizes the need for a thorough reassessment of the scorpion phylogenetic relationships.
Electronic supplementary material
The online version of this article (10.1186/s12983-017-0231-z) contains supplementary material, which is available to authorized users.
... This species comprises small to medium-sized scorpions that have brownish coloration, with dorsal lighter spots (Fig. 1). H. lunatus differs from other Hadruroides scorpions by the curved morphology of the pedipalp fixed finger, creating a gap when fingers are closed, and by the rectangular shape of the spots on the tergites [2]. ...
Background:
Hadruroides lunatus is the most abundant scorpion species in the Peruvian central coast, where most of the accidents involving humans are registered. In spite of its prevalence, there are only very few studies on H. lunatus envenomation. The aim of the present study was to analyze the cardiorespiratory alterations caused by H. lunatus envenomation in rodents.
Methods:
Wistar rats injected with H. lunatus scorpion venom were submitted to electrocardiography. After euthanasia, rat lungs were collected and histopathologically analyzed. Mouse cardiomyocytes were used to perform immunofluorescence and calcium transient assays. Data were analyzed by ANOVA or Student's t-test. The significance level was set at p < 0.05.
Results:
It was observed that H. lunatus venom increased heart rate and caused arrhythmia, thereby impairing the heart functioning. Lungs of envenomed animals showed significant alterations, such as diffuse hemorrhage. In addition, immunofluorescence showed that H. lunatus venom was capable of binding to cardiomyocytes. Furthermore, mouse ventricular cardiomyocytes incubated with H. lunatus venom showed a significant decrease in calcium transient, confirming that H. lunatus venom exerts a toxic effect on heart.
Conclusion:
Our results showed that H. lunatus venom is capable of inducing cardiorespiratory alterations, a typical systemic effect of scorpionism, stressing the importance of medical monitoring in envenomation cases.
... Most of these symptoms 53 are caused by the action of neurotoxins on the victim's ion channels (Chippaux, 2012). Verde, 1968;Kinzelbach, 1973;Maury, 1975;Cekalovic, 1983;Sissom and Fet, 2000), 63 however they are currently restricted to Peru, Ecuador, northern Chile and several 64 offshore islands (Ochoa and Prendini, 2010). Hadruroides lunatus ("escorpion de los Hematological parameters were measured using the same blood samples collected for 115 biochemical analysis. ...
Toxic effects of Peruvian Hadruroides lunatus scorpion venom on different biochemical and enzymatic parameters in blood serum of Wistar rats and Swiss mice were determined after experimental envenomation. An increase in enzymatic activities of Aspartate Aminotransferase (AST), Lactate Dehydrogenase (LDH) and levels of serum protein and albumin were observed while a decrease in creatinine level in serum was perceived after 30 minutes of envenomation. No alterations in urea levels and in kidney histology were detected in the envenomed rats. The global leukocytes count was diminished, with decrease in lymphocytes, eosinophils and neutrophils levels in the bloodstream, while no alterations were found in hematological parameters of red series in rats injected with H. lunatus venom. IL-2, IL-4, IL-6, INF-γ, TNF, IL-17A and IL-10 levels were evaluated 0.5, 3 and 6 h after experimental envenomation of mice with H. lunatus venom. From all the analyzed cytokines, only IL-6 showed an increase in serum levels. Taken together, these results point out that envenomation by H. lunatus can impair hematological and immunological parameters and therefore might be monitored in accidents involving this species.
Copyright © 2015. Published by Elsevier Ltd.
Body size has always been the focus of several ecological studies due to its undeniable influence on other life-history traits. The conventional representation of body size in arthropods typically relies on linear measures, such as total body length, or the length of specific body parts that can be used to represent body size. While these measures offer simplicity over more complicated alternatives ( e.g ., dry mass), technical problems persist for arthropods with complex body structures, as is the case for scorpions. In these animals, accurate measurements often require extensive handling, including the stretching of body parts. In light of the difficulties associated with directly measuring total length and carapace length in scorpions (two prevalent proxies for body size in the group), this study evaluates the ability of seven simple linear measurements in predicting length measures of boy size in scorpions under a phylogenetic framework. Predictive equations derived from phylogenetic mixed linear models fitted under Bayesian framework were implemented in custom R functions that can be applied for size prediction in a wide range of scorpions. Overall, accurate predictions of total length and carapace length could be achieved using any of the studied traits as single predictors. However, the most accurate predictions for total length were obtained using the length of metasomal segment V, while the best predictions for carapace length were achieved using telson length. The addition of a secondary predictor had low impact on the quality of the size predictions, indicating that increasing model complexity by incorporating additional predictors is not necessary to achieve accurate size estimates. Technical advantages and limitations associated with each linear measurement are discussed. In conclusion, this study broadens the repertoire of methods available for accurately estimating body size in scorpions, particularly in instances where body size information can only be obtained indirectly through allometric relationships.
The true identity of Brachistosternus peruvianus Piza, 1974, a controversial Andean scorpion previously confused with Brachistosternus andinus Chamberlin, 1916, is established. Both species inhabit inter Andean valleys of southern Peru and belong to the subgenus Brachistosternus (Ministernus) Francke. A detailed redescription of both species based on external morphology, pigmentation pattern and hemispermatophore features are provided. New records for these species, along with brief discussions of the ecology and distribution are also reported.
The scorpion Diplocentrus zacatecanus Hoffmann (1931) was originally described as a subspecies of Diplocentrus keyserlingi Karsch 1880 on the basis of six syntypes and was later elevated to species level. We designate a male lectotype and redescribe the species, including illustrations of the hemispermatophore of a male collected near the type locality. In this genus, the hemispermatophore is poorly sclerotized and lacks elaborate capsular structures, which are taxonomically useful in other genera. We review the variability in the hemispermatophores of males from one population, including five comparisons of the right and left hemispermatophores of the same males. Our results showed asymmetry in the length of the right and left hemispermatophores of the same individual. We also observed the presence of ''crenulations'' or ''spines'' in two different hemispermatophores (not complementary ones). We conclude that caution should be used when describing the hemispermatophore of only one male and considering it as diagnostic for the species, because of the high levels of intraspecific variation.
Rumikiru, n. gen., a new bothriurid scorpion genus from the coastal Atacama Desert, Chile, is described. This is the first scorpion genus endemic to northern Chile. It is most closely related to Pachakutej Ochoa, 2004, from the inter-Andean valleys of Peru. Orobothriurus lou-rencoi Ojanguren-Affilastro, 2003, is transferred to the new genus and redescribed, creating Rumikiru lourencoi (Ojanguren-Affilastro, 2003), n. comb., and a second species of the genus, Rumikiru atacama, n. sp., is described.
The systematics of the Andean scorpion genus, Orobothriurus Maury, 1976 (Bothriuridae Simon, 1880), is revised. New locality records, obtained during recent field expeditions, distribution maps, and a key to identification of the 15 known species, are provided. Six new species are described: Orobothriurus calchaqui, n. sp., from northwestern Argentina; Orobothriurus compagnuccii, n. sp., from the central Andes of Argentina; Orobothriurus huascaran, n. sp., from central Peru; Orobothriurus quewerukana, n. sp., from southern Peru and northern Chile; Orobothriurus ramirezi, n. sp., from central Chile; and Orobothriurus tamarugal, n. sp., from northern Chile. The known distribution of Orobothriurus and the altitude record for scorpions are discussed. The world's altitude record for a scorpion, previously reported as 5550 m, is demonstrated to be 4910 m.
The fine structure of the book lungs of scorpions is diverse and phylogenetically informative,
but has not been comprehensively investigated across the major lineages of the order. In this
contribution, we present a fully illustrated atlas of the variation in book lung fine structure
among 200 exemplars from 100 genera and 18 families of extant scorpions. We document
variation in the surface sculpturing of the respiratory lamellae, the edges of the lamellae in the
atrial chamber, and the posterior valvelike edges of the spiracles. These data provide insights
into the phylogenetic relationships among Recent scorpions at several branches of the tree.
Se presenta la distribución geográfica de 24 especies de escorpiones (cinco géneros y tres familias) para el corredor andino en el sur del Perú. Se discuten algunos aspectos zoogeográficos de las especies involucradas (19 exclusivas del área de estudio), las que pueden reunirse en siete áreas escorpiológicas: Desierto costero, Lomas, Serranía esteparia, Puna, Queswa, Valles interandinos cálidos y Yungas. Las Lomas, Serranía esteparia y Queswa, presentan la mayor riqueza específica. Se analizan los patrones de distribución de las especies, que sugieren la presencia de tres componentes faunísticos en el sur del Perú: andino patagónico, amazónico y caribeño.
Hl3 es una toxina que, en un trabajo anterior, fue aislada por nuestro grupo de investigación a partir del veneno del escorpión Hadruroides lunatus mediante cromatografía de intercambio iónico en CMSephadex C-25 a pH 7, y que se caracteriza por producir contracción y parálisis en la extremidad inoculada de ratones albinos. En este trabajo se ha determinado que la inoculación de Hl3 (26 μg) en el músculo gastrocnemius produce la liberación, en el plasma, de creatina kinasa (CK) y lactato deshidrogenasa (LDH). La actividad de CK se incrementa desde 210 UI/L hasta 8015 UI/L luego de 60 minutos, mientras que la actividad de LDH aumenta desde 174 UI/L hasta 1697 UI/L luego de 90 minutos. Por PAGE-SDS, se ha demostrado que la incubación de Hl3 (26 μg) con el músculo gastrocnemius aislado de ratones albinos, en medio fisiológico a pH 7, provoca también la liberación de otras proteínas musculares. Además, Hl3 es capaz de incrementar mas de 3,5 veces los niveles normales de calcio intramuscular, luego de 45 minutos de acción. Nuestros resultados sugieren que Hl3 actúa específicamente a nivel muscular, afectando la permeabilidad del sarcolema y provocando la fuga de proteínas musculares. Además el incremento de los niveles de calcio podría generar hipercontracción y la destrucción de fibras musculares.
The proteins from the venom of the scorpion Hadruroides lunatus were separated by ion- exchange chromatography on CM-Sephadex C-25 with 0,05M ammonium acetate buffer pH 7, getting six protein peaks in the process. The toxicity tests allowed the identification of three toxins that were denominated Hl1, Hl2 y Hl3; which immobilize, respectively, insects (Grillus sp.), crustaceans (Porcellio laevis) and the inoculated limb of white mice. The three toxins are basic proteins and have no proteolitic or phospholipase activity. In addition, Hl3 increases the plasmatic level of creatine kinase after its inoculation in the gastrocnemius muscle of mice. By PAGE-SDS, Hl3 shows only one protein band of 12,5 KDa, while the other toxins have protein contaminants.
Venezuela se ubica en el primer lugar de los países suramericanos en cuanto a diversidad de escorpiones. La descripción de un número importante de nuevas especies y los recientes cambios taxonómicos han alterado drásticamente la composición de este grupo de arácnidos. Se presenta la lista actualizada de la escorpiofauna de Venezuela, y para cada especie se ofrece información detallada sobre material tipo y distribución geográfica, con comentarios taxonómicos y una selección de referencias bibliográficas. Se añade una lista de exclusión de especies citadas para el país por autores previos. Actualmente se reconocen para Venezuela cinco familias (Buthidae, Chactidae, Euscorpiidae, Hemiscorpiidae y Scorpionidae), 17 géneros y 184 especies. De estas últimas, 55 han sido descritas en el último quinquenio, elevando a 168 (91,3%) el número de elementos endémicos del país. Por último, se discuten algunos aspectos de la escorpiofauna de Venezuela.
A unique orthobothriotaxic pattern of family Iuridae is examined and illustrated. Significant positional differences in this pattern between two monotypic iurid genera, Calchas and Iurus, are presented. With the analysis of over 100 specimens of genus Iurus, representing many localities in Greece and Turkey, several unique occurrences of neobothriotaxy involving both the chela and patella are discussed and illustrated, representing the first report of neobothriotaxy in family Iuridae.
Hl3 is a toxin isolated by our investigation group from Hadruroides lunatus scorpion venom by ionexchange chromatography on CM-Sephadex C-25 to pH 7, and which produce contraction and paralysis in the inoculated limb of white mice. In this work we have determined that inoculation of Hl3 (26 μg) in gastrocnemius muscle from mice produces the release in plasma of creatine kinase and lactate deshidrogenase. The creatine kinase activity increases from 210 UI/L to 8015 UI/L in 60 minutes, whrereas the LDH activity increases from 174 UI/L to 1697 UI/L in 90 minutes. By PAGE-SDS it, was demonstrated that incubation of Hl3 with gastrocnemius muscle isolated from white mice, in physiological solution to pH 7, produces the release of other different muscular proteins as well. In addition Hl3 can increase the calcium intramuscular 3,5 fold, after 45 minutes of action. Our results suggest that Hl3 has specific action on the muscle, affecting the sarcolema permeability and producing the release of muscle proteins. In addition the increase of calcium could produce hipercontraction and destruction of muscular fibers.
In this paper are presented the results of the study of an interesting collection of neotropical scorpions deposited in the Geneva Museum. The collection is composed of 6 families, 14 genera and 42 species. Two new species, Tityus dinizi n. sp. (Buthidae) and Broteochactas kelleri n. sp. (Chactidae) are described. Some comments on the species which are able to climb the vegetation and on those living in high mountain habitats are also included.
S. A. Stockwell (1989, “Revision of the Phylogeny and Higher Classification of Scorpions (Chelicerata).” Univ. of California, Berkeley) proposed a cladogram and revised classification for the superfamily Scorpionoidea Latreille 1802 (comprising the families Bothriuridae, Diplocentridae, Heteroscorpionidae, Ischnuridae, and Scorpionidae), based on 47 morphological characters and 35 supraspecific terminal taxa, representing genera whose monophyly was implicitly assumed. Given the widespread practice of defining scorpion genera on the basis of plesiomorphic character states, the assumption of monophyly implicit in supraspecific terminal taxa reduces confidence in Stockwell’s cladistic findings and, consequently, his revised suprageneric classification. A re-investigation of scorpionoid phylogeny is presented here, based on 115 morphological characters (including the characters used by Stockwell) and 71 exemplar species. The criterion of “maximal morphological diversity” was employed for exemplar selection. This approach provides a stronger test of monophyly than random exemplar selection. Sixteen cladistic analyses were performed on the scorpionoid data matrix, which varied in the use of equal, successive, or implied weights and in the additive or nonadditive treatment of multistate characters. The preferred hypothesis, a single most parsimonious tree obtained by analysis with equal weights and 13 ordered multistate characters, yielded the scheme of relationships: (Bothriuridae ((Heteroscorpionidae Urodacinae) ((Hemiscorpiinae Ischnuridae) (Diplocentridae Scorpioninae)))). On the basis of these results, revisions are proposed to the existing suprageneric classification of the Scorpionoidea, including new diagnoses, new descriptions, and an illustrated key to the families and subfamilies. Familial status is provided for the scorpionid subfamilies Hemiscorpiinae and Urodacinae.
The fine structure of the book lungs of scorpions is diverse and phylogenetically informative, but has not been comprehensively investigated across the major lineages of the order. In this contribution, we present a fully illustrated atlas of the variation in book lung fine structure among 200 exemplars from 100 genera and 18 families of extant scorpions. We document variation in the surface sculpturing of the respiratory lamellae, the edges of the lamellae in the atrial chamber, and the posterior valvelike edges of the spiracles. These data provide insights into the phylogenetic relationships among Recent scorpions at several branches of the tree.
SUMARIO En la biorregión desértico-costera del Perú, los Tilland-siales constituyen un biotopo característico que se presen-ta a todo lo largo de ella, y está conformado por asocia-ciones de Bromeliáceas del género Tillandsia, cuya adapta-ción a zonas desérticas es muy notable y por ello reciben el nombre de "tillandsias grises". La zona de Cajamarquilla constituye una quebrada de orientación S -N, en la margen derecha del río Rímac, a unos 50 km. del mar (Distrito Vitarte)., Incluye también un "cauce seco" del mismo nombre. Las Tillandsia se encuentran principalmente sobre la-deras que reciben la influencia del viento húmedo. Crecen allí las especies T. purpurea, T. latifolia, T. paleacea y T. recurvata, entre los 300-700 m.s.n.h. (Türkowsky, 1976). Se informan los resultados de observaciones prelimina-res sobre la fauna, que en el caso de vertebrados fueron esporádicas (15 especies: 3 reptiles, 7 aves y 5 mamíferos) y en el caso de los artrópodos, se llevó a cabo a base de trampas líquidas enterradas a ras del suelo, en áreas abiertas, debajo el follaje y también a media altura, entre el follaje de las plantas. Las observaciones se hicieron durante 1974-1975. Se hallaron 5 órdenes de arácnidos con 12 familias y 13 órdenes de insectos con más de 38 fa-milias.
The southern African species of Parabuthus Pocock 1890 are reviewed. Twenty valid species are recognized and an illustrated key is provided for their identification. The diagnosis of each species is revised, its known distributional range summarized, and notes on its ecology and conservation provided. Three species are redescribed and their distributions mapped: P. calvus Purcell 1898; P. capensis (Ehrenberg 1831); P. planicauda (Pocock 1896). Six new synonyms are proposed: Buthus brevimanus var. β segnis Thorell 1876 = P. granulatus (Ehrenberg 1831); Buthus villosus var. β dilutus Thorell 1876 = P. raudus (Simon 1888); P. granulatus strenuus Hewitt 1918 = P. granulatus (Ehrenberg 1831); P. capensis frenchi Purcell 1901 = P. planicauda (Pocock 1889); Scorpio teter Müller 1828 = P. transvaalicus Purcell 1899; P. brachystylus Lawrence 1928 = P. villosus (Peters 1862). Two species, synonymized by previous authors, but subsequently resurrected, are returned to synonymy: P. neglectus Purcell 1899 = P. capensis (Ehrenberg 1831), first synonymized by Kraepelin (1908); P. flavidus Pocock 1899 = P. mossambicensis (Peters 1861), first synonymized by Kraepelin (1914).
The higher categories of the Vejovidae have been revised and keys to these categories are pre- sented . As part of the revision a new subfamily, the Hadrurinae, has been recognized . In the sub - family Syntropinae a new genus, Vejovoidus, has been introduced . In the subfamily Vejovinae ne w genera recognized are Serradigitus and Pseudouroctonus . Thirteen species previously placed in th e genus Uroctonus have been shown to belong to the genus Vejovis. An extensive study of the value o f trichobothria in scorpion systematics is presented . The systematic status of Uroctonus fractus i s doubtful . It has been eliminated from the Vejovidae and apparently should be placed in the Chactida e where it will undoubtedly be synonymized .
The ‘giant hairy scorpions’, genus Hadrurus Thorell, are the largest and most conspicuous scorpions in North America, but their systematics has long been confused. A new genus, Hoffmannihadrurus Fet et al. was recently created to accommodate the two species endemic to mainland Mexico, Hadrurus aztecus Pocock and Hadrurus gertschi Soleglad. In the present contribution, we review the taxonomic history of Hadrurus, re-evaluate the phylogenetic relationships among its component species with an analysis based on morphological characters and a taxon sample representing the known morphological variation, and assess the validity of the new genus. Seven independent analyses of the morphological character matrix, under weighting regimes that minimised length as well as those that maximised fit, each located a single most parsimonious tree with the following scheme of relationships: (Iurus ((Caraboctonus + Hadruroides) (H. aztecus ((H. gertschi + H. pinteri) (H. concolorous + H. hirsutus)) ((H. obscurus + H. spadix) (H. a. arizonensis (H. a. austrinus + H. a. pallidus)))))). This topology supports the monophyly of Caraboctoninae, Caraboctonini, Hadrurini, the ‘arizonensis’ subgroup, H. arizonensis and H. concolorous. It does not support the monophyly of Hoffmannihadrurus, Hadrurus, the ‘aztecus’ group, the ‘hirsutus’ group, and the ‘hirsutus’ subgroup, all of which were rendered paraphyletic by the grouping of H. gertschi with H. pinteri, rather than with H. aztecus. The results unequivocally demonstrate that the creation of a new genus for the mainland Mexican species was unfounded. We therefore propose the following new synonymy: Hoffmannihadrurus Fet et al., 2004 = Hadrurus Thorell, 1876, syn. nov.
We present a review and reassessment of anatomical variation in the ovariuterus (and associated follicles), digestive gland, and lateral lymphoid organs of scorpions, and discuss the contribution of these character systems to the understanding of scorpion phylogeny. New data, obtained using light microscopy, are presented from an examination of 55 scorpion species, representing most scorpion families, and are collated with observations from the literature. Six distinct types of ovariuterine anatomy are identified: five in the family Buthidae and one in the remaining (nonbuthid) families. The buthid genera Lychas C.L. Koch, 1845 and Rhopalurus Thorell, 1876 are exceptional in possessing multiple types of ovariuterine anatomy among the congeneric species studied. The presence or absence of lateral lymphoid organs appears to be phylogenetically informative: the organ is absent in buthids, chaerilids and Pseudochactas Gromov, 1998. Embryo follicle morphology appears to be phylogenetically informative within the superfamily Scorpionoidea Latreille, 1802, where it supports the controversial sister-group relationship between Urodacus Peters, 1861 and Heteroscorpion Birula, 1903. The mesosomal anatomy of Microcharmus Lourenço, 1995 (Microcharmidae Lourenço, 1996) is consistent with that of Buthidae C.L. Koch, 1837, and we therefore propose the following new synonymy: Microcharmidae Lourenço, 1996 = Buthidae C.L. Koch, 1837. © 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 154, 651–675.
S. A. Stockwell (1989, “Revision of the Phylogeny and Higher Classification of Scorpions (Chelicerata).” Univ. of California, Berkeley) proposed a cladogram and revised classification for the superfamily Scorpionoidea Latreille 1802 (comprising the families Bothriuridae, Diplocentridae, Heteroscorpionidae, Ischnuridae, and Scorpionidae), based on 47 morphological characters and 35 supraspecific terminal taxa, representing genera whose monophyly was implicitly assumed. Given the widespread practice of defining scorpion genera on the basis of plesiomorphic character states, the assumption of monophyly implicit in supraspecific terminal taxa reduces confidence in Stockwell's cladistic findings and, consequently, his revised suprageneric classification. A re-investigation of scorpionoid phylogeny is presented here, based on 115 morphological characters (including the characters used by Stockwell) and 71 exemplar species. The criterion of “maximal morphological diversity” was employed for exemplar selection. This approach provides a stronger test of monophyly than random exemplar selection. Sixteen cladistic analyses were performed on the scorpionoid data matrix, which varied in the use of equal, successive, or implied weights and in the additive or nonadditive treatment of multistate characters. The preferred hypothesis, a single most parsimonious tree obtained by analysis with equal weights and 13 ordered multistate characters, yielded the scheme of relationships: (Bothriuridae ((Heteroscorpionidae Urodacinae) ((Hemiscorpiinae Ischnuridae) (Diplocentridae Scorpioninae)))). On the basis of these results, revisions are proposed to the existing suprageneric classification of the Scorpionoidea, including new diagnoses, new descriptions, and an illustrated key to the families and subfamilies. Familial status is provided for the scorpionid subfamilies Hemiscorpiinae and Urodacinae.
Soleglad and Fet's (2003a) attempt to reconstruct the phylogeny of Recent (including extant) scorpions, the revised classification derived from it, and recent emendations, mostly published in their self-edited online journal, Euscorpius, are deficient. Separate analyses of three independent matrices (morphology, 16S rDNA, 18S rDNA) were presented. In the morphological matrix, 52 binary and 10 tristate trichobothrial characters were replaced with one character comprising six ordered states representing trichobothrial “types”. The remaining matrix of 105 characters was further reduced to 33 “fundamental” characters (20% of the morphological dataset), the analysis of which appears to be the basis for the revised classification presented. The taxon sample for the morphological analysis included 14 supraspecific terminal taxa representing genera, the monophyly of only 7 (12.5%) of which has been confirmed. A composite terminal, assembled from the fragments of fossils that may not be confamilial let alone monophyletic, was created for the Palaeopisthacanthidae, employed as the primary outgroup for the analysis. Other important outgroup taxa, notably eurypterids, xiphosurans and other arachnids, were omitted entirely. The morphological characters presented contained numerous unjustifiable assumptions of character polarity and phylogenetic relationship. An approach to character coding, deliberately adopted to reduce “homoplasy”, biased the analysis towards a preconceived result. Structurally and topographically similar features in different taxa were explicitly assigned separate (often autapomorphic) states according to presumed phylogenetic relationships among the taxa in which they were observed. Putative “reversals” were coded as separate characters or states. Character transformation was forced by ordering, additive coding or Sankoff optimization through allegedly intermediate states for which there is no empirical evidence. Many characters were defined in a manner that demonstrates either a lack of understanding of, or disregard for, established methods and standards of morphological character coding. Some states display overlapping variation whereas others subsume variation that is not structurally or topographically similar. Polymorphic “states” were created for terminals with interspecific variation and unknown “states” for terminals that should have been scored unknown. Many characters were not evaluated for particular terminal taxa, but merely scored inapplicable although the structures and, consequently, the characters in question are present and therefore applicable to them. In view of the significant theoretical and empirical problems with the approach to cladistics taken by Soleglad and Fet, we find no justification for accepting either the results of their analyses or the revised classification derived from them. Pending the outcome of a rigorous phylogenetic analysis, published according to acceptable standards of scholarship in a peer-reviewed journal, we revert to the suprageneric classification of Scorpiones reflected by the most recent peer-reviewed, published treatments and reject all changes to the classification proposed by Soleglad, Fet and colleagues since 2001. We argue that an analysis and revised classification of the kind presented in various papers by these authors could not survive the peer-review process of a mainstream scientific journal. The poor scholarship exemplified by these and other papers published in Euscorpius emphasize the importance of quality control associated with the emergent infrastructure of online publishing. A centralized register of taxa may be the only solution for ensuring quality control in the taxonomy of the future.© The Willi Hennig Society 2005.
The author gives an account of the flora and vegetation of the Peruvian coast, which is a belt located along the Pacific coast of South America from 3º 23' to 18° 21' South. This belt is 2.500 km long and 10 to 150 km wide. Within this territory there are two ecosystems: the algarrobal (carob wood) formation and the lomas (hills) formation. As a whole these ecosystems are characterized by the following types of vegetation: mangrove, chaparral, ceibal, perennial wood, algarrobal (carob wood), sapotal, gramadal (grassland), lomas (hills), tilansial and coastal woodland. El autor hace un análisis de las comunidades vegetales de la costa peruana que tiene una longitud aproximada de 2.500 km por 10 a 150 km de anchura. Se encuentra en la costa pacífica de Sudamérica desde los 3º 23' al norte hasta los 18° 21' al sur. En este territorio angosto habitan dos grandes formaciones que están estrechamente correlacionadas con los factores bioclimáticos: el algarrobal y las lomas. Dentro de estos ecosistemas hemos considerado los siguientes tipos de vegetación: manglar, chaparral, ceibal, monte perennifolio, algarrobal, sapotal, gramadal, lomas, tilansial y monte ribereño.