Catalogue
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Catalogue
Updated catalogue and taxonomic notes on the Old-World scorpion genus Buthus Leach, 1815 (Scorpiones, Buthidae)
expand article infoPedro Sousa§|, Miquel Arnedo|, D. James Harris§
‡ Universidade do Porto, Vairão, Portugal
§ Universidade do Porto, Porto, Portugal
| Universitat de Barcelona, Barcelona, Spain
Open Access

Abstract

Since the publication of the ground-breaking “Catalogue of the scorpions of the world (1758–1998)” (Fet et al. 2000) the number of species in the scorpion genus Buthus Leach, 1815 has increased 10-fold, and this genus is now the fourth largest within the Buthidae, with 52 valid named species. Here we revise and update the available information regarding Buthus. A new combination is proposed: Buthus halius (C. L. Koch, 1839), comb. n. from Portugal and Spain. B. halius is removed from junior synonymy with Buthus occitanus (Amoreux, 1789), and proposed as a senior synonym of B. ibericus Lourenço & Vachon, 2004, syn. n. Moreover, following I.C.Z.N. article 23.9.2 we propose to maintain as valid B. ibericus (nomen protectum) and to consider the disued B. halius as a nomen oblitum. Buthus europaeus tridentatus Franganillo, 1918 is proposed as a junior synonym of B. occitanus (Amoreux, 1789), syn. n. Buthus sabulicola Touloun, 2012 is proposed as a junior synonym of Buthus bonito Lourenço & Geniez, 2005, syn. n. Buthus occitanus tunetanus neeli Gysin, 1969 is proposed as an informal senior synonym of Buthus tassili Lourenço, 2002, informal syn. n. Two taxa are rised to species rank, Buthus nigrovesiculosus Hirst, 1925, stat. n. and Buthus parroti Vachon, 1949, stat. n.. We further confirm the restricted distribution of B. occitanus that is confined to southeastern France and northwestern Iberian Peninsula and does not occur in North Africa. Additionally, Androctonus barbouri (Werner, 1932), comb. n. from the Agadir region of Morocco, is hereby transferred to the genus Androctonus. We summarize and provide a critical appraisal of the diagnostic characters currently in use for the genus. The catalogue section considers the names for species, subspecies and varieties that have been used for Buthus scorpions. Information about types, including collection numbers and localities are included when available. Finally, an annotated listing of synonymies and an updated bibliography are given.

Keywords

Taxonomy, new synonymy, new combination, new status, Geographic distribution, Africa, Asia, Europe, diagnostic characters

Introduction

Members of the genus Buthus Leach, 1815 are medium-sized scorpions, usually yellowish in colour, with a robust metasoma that ends in a telson with a globular vesicle and a curved aculeus (Fig. 1). Buthus rest during the day in burrows under stones or shrubs and are active from dusk till dawn, although their activity typically peaks at the beginning of the night (Cloudsley-Thompson 1956). They are successful scorpions that, when present, tend to be the most abundant scorpion in their habitat, as for example in the Iberian Peninsula and Morocco. Buthus are usually described as sit-and-wait predators, although they can also actively search for prey (Skutelsky 1995, Piñero et al. 2013). Buthus is among the most venomous of all scorpion genera (Chippaux and Goyffon 2008). Buthus venom toxicity is considered much lower in Europe than in North Africa, which can be empirically corroborated by the few severe cases of scorpionism reported for Western Europe when compared to the North African countries (Chippaux and Goyffon 2008).

Figure 1. 

Buthus mariefranceae, from south of Morocco. Photo by Arie van der Meijden.

Buthus exhibits a wide distribution range, spanning over two biogeographic realms, the Palearctic (Western) and the Afrotropical (Udvardy 1975, Olson et al. 2001). Interestingly, none of the chorotypes proposed by Vigna Taglianti et al. (1999) satisfactorily describes Buthus distribution. The genus extends from the temperate Mediterranean areas of south-western Europe to the tropical and sub-tropical grasslands south of the Sahel and into the Horn of Africa, including the semi-arid and arid regions of North Africa and the Middle East (Fig. 2). Although first considered of European origin (Vachon 1952a), current data support the hypothesis that the centre of origin of the genus is North Africa (Lourenço 2002). North Africa harbours a disproportionate number of species (Fig. 2) as well as four of the five main genetic clades found in Buthus, as defined by mitochondrial DNA sequence variation (Sousa et al. 2012, Pedroso et al. 2013) (Fig. 7).

Buthus species are known from 17 countries in Africa: Algeria, Cameroon, Central African Republic, Chad, Egypt, Eritrea, Ethiopia, Guinea, Libya, Mauritania, Morocco, Niger, Senegal, Somalia, South Sudan, Sudan, and Tunisia; five countries in Asia: Cyprus, Egypt (Sinai), Israel, Jordan, and Yemen; and four European countries: France, Italy (Sicily), Portugal, and Spain (Fig. 2). Unidentified Buthus species have also been reported from Burkina Faso, Djibouti, Gambia, Ghana, Guinea-Bissau, Ivory Coast, Nigeria, Iraq, and Lebanon (Fig. 2). No records exist for Saudi Arabia or Syria, however the first might have been confused in the past with citations for the Arabian Peninsula (e.g. Vachon 1952a), although the existence of Buthus in either of these countries cannot be excluded. The frequently cited occurrence of Buthus in Iraq is based on a single specimen, deposited in the Czech National Museum of Natural History (Táborský 1934, Kovařík 1992). As such the actual distribution of the genus remains poorly delimited. Old records from mainland Greece and Turkey are highly doubtful, as these are well-studied areas with no recent Buthus collections (Ersen Yağmur pers. comm. for Turkey) (Fig. 2). The former records most likely refer to the genus Mesobuthus Vachon, 1949. As for the record for Malta, it was considered dubious by Fet and Lowe (2000), although other Buthus reported on islands that were previously regarded as doubtful have turned out to be correct, namely B. kunti Yağmur, Koc & Lourenço, 2011, from Cyprus, described from freshly collected material and B. trinacrius Lourenço & Rossi, 2013, from Sicily, based on 130-year-old material. Other island records include B. occitanus in the Columbrete islands (Castilla and Pons 2007) and B. tunetanus (Herbst, 1800) in the Tunisian islands of Djerba, Kerkena, and Zembra (Vachon 1952a).

Figure 2. 

Map of Buthus species distribution, and the known number of species by country. Also depicted are the species’ type localities (numbers according to the species’ Catalogue and Table 2) where known or the best possible approximation. Actual distribution within each country can be much smaller, but detailed distribution information is unknown for the majority of species.

Buthus is the type genus of the Buthidae C. L. Koch, 1837 (Koch 1837, 1850), the most diverse family within Scorpiones, with almost half of all known extant scorpion species (1101 of the 2311 known species) (Rein 2016). The Buthidae also includes most of the species venomous to humans (Chippaux and Goyffon 2008). The genus Buthus is the second oldest valid genus of the order Scorpiones C. L. Koch, 1837, only surpassed by the single genus created by Linnaeus in 1758, Scorpio, to accommodate all the scorpion species he described.

Buthus was first proposed by Leach (1815), with Scorpio occitanus Amoreux, 1789 as its type species (Fig. 3). The only diagnostic character for the genus proposed by Leach was the presence of eight eyes “Oculi octo.” (Leach 1815, page 391), apparently referring to the pair of central eyes and three pairs of lateral eyes. This character was mentioned as diagnostic for different scorpions early during Scorpiones taxonomy (De Geer 1778, Fabricius 1781). The same diagnostic character was used by Ehrenberg (in Hemprich and Ehrenberg 1828, 1829) for defining several genera and subgenera of scorpions with a varying number of eyes, ranging from six to 12. A more detailed explanation on the usage of the number of eyes in the classification of scorpions is given in Thorell (1876).

Figure 3. 

Original illustration of Scorpio occitanus (Amoreux 1789a). The work was retrieved from the Biodiversity Heritage Library and images were rearranged for compactness without re-scaling.

The poor description of Leach (1815) led to a rapid increase in the number of species included in the genus, which lacked any internal coherence. This taxonomic conundrum arose through the misidentification of the number of lateral eyes of B. occitanus (Amoreux, 1789), originally stated by Amoreux as three pairs. Several taxonomists of that century realised that there were actually four pairs of lateral eyes (e.g. Gervais 1844b; Simon 1879), but this information was not appreciated by some later authors. It has recently been shown that most Buthidae species (including Buthus) have five pairs of lateral eyes, although in many species two pairs of lateral eyes are much smaller in size and require extreme care and the help of UV light to be recognised (Yang et al. 2013; Loria and Prendini 2014). Ehrenberg (Hemprich and Ehrenberg 1828) modified the original meaning of the genus to include the species that are now part of Heterometrus Ehrenberg in Hemprich and Ehrenberg 1828 (Family Scorpionidae Latreille, 1802), all with five pairs of lateral eyes. Ehrenberg described several other genera that were soon synonymized with Buthus, at least by some taxonomists, which further exacerbated the taxonomic confusion within Buthus.

Because of the poor definition of the genus, many members (≈100 species) of the Buthidae family with no close relationship to the type species, were included in the genus Buthus up to the mid-20th century (Vachon 1952a, Levy and Amitai 1980, Lourenço 2002). Unfortunately, this obsolete taxonomy is still in use, for example in many toxicology and venom related papers on scorpions (e.g. Gopalakrishnakone et al. 2015). From 1948 to 1951, Vachon conducted a major taxonomic revision of the genus (compiled in Vachon 1952a), providing a more informative and exclusive definition, retaining only the species that were morphologically similar to the type species and hence restricting also the distribution range of the genus. He proposed two main morphological characters that in combination separate Buthus from all other known Buthidae genera: the central-lateral and posterior-median prosomal keels fused in a lyra shape (character shared with Cicileiurus Teruel, 2007, Leiurus Ehrenberg in Hemprich and Ehrenberg 1828, Mesobuthus Vachon, 1950, and Odontobuthus Vachon, 1950, Fig. 4A), and the presence of only three granules on the tip of the movable finger (or tarsus) of the pedipalp chela (character shared with Androctonus Ehrenberg in Hemprich and Ehrenberg 1828, Fig. 4B).

Figure 4. 

The two key morphological diagnostic characters of the genus Buthus. A- Prosoma carapace with lyra-shaped keels (Hjelle 1990); B - Tip of pedipalp movable finger highlighting the three distal granules (distal denticle not included) (Lourenço 2002).

While studying specimens from northwest Africa, Vachon recognized that the genus included a large amount of undescribed diversity. Vachon took a very conservative approach to Buthus taxonomy, recognizing only four species, further split in 12 subspecies, ten of which under B. occitanus, and naming four different varieties, along with other forms with no formal rank, all within B. occitanus. This was partly justified by Vachon’s view that Buthus species exhibited a large morphological plasticity, at least in the characters he used to diagnose the different taxa (Vachon 1952a). Stahnke (1972), in his key to Buthidae genera, recognized 21 species and subspecies in Buthus, without further explanation, although this is probably an error resulting from an outdated interpretation of the genus taxonomy. In accordance with the ICZN article 45, none of Vachon’s infra- subspecific varieties were included in the Catalogue of the Scorpions of the World (Fet et al. 2000). The Buthus Catalogue recognized as good five species and 12 subspecies, although the authors recognized that some taxa were probably not taxonomically good (Fet and Lowe 2000) Subsequently, Rossi (2015) transferred Buthus insolitus Borelli, 1925 to the recently erected genus Gint Kovařík et al., 2013. Lourenço (2003) marked a renewed interest in the taxonomy and diversity of the genus, describing six new species, some of which corresponding to Vachon’s infra-subspecific varieties.

During the last 15 years, the rate of description of new Buthus species has increased exponentially (Fig. 5). At present, the genus is composed of 52 species, three of which were described in 2016, making it the fourth most diverse genus of Buthidae, only surpassed by the megadiverse scorpion genera Tityus C. L. Koch, 1836, Centruroides Marx, 1890 and Ananteris Thorell, 1891 (Rein 2016). Thirty authors have been involved in the description of recent Buthus species, and most species (21) have been described in collaborative studies. Wilson Lourenço is by far the most prolific author, having authored or co-authored 29 Buthus species, 55% of the total.

Figure 5. 

Cumulative number of valid named Buthus species. Only current valid species' names were plotted; in the year they were first described.

Materials and methods

Nomenclature and measurements follow Stahnke (1970), except for trichobothriotaxy (Vachon 1974, Fet et al. 2005). All diagnostic morphological characters mentioned in the text refer to adults (or large sub adults) of both sexes, unless otherwise noted.

Most references prior to 1998 cited by Fet and Lowe (2000) are confirmed, but not all original literature could be obtained, and we made some corrections following comparisons with additional sources (Vachon 1952a, Lamoral 1979, Polis 1990, Hendrixson 2006, Dupré 2013). The criteria applied by Fet and Lowe (2000) citing both taxonomic and faunistic works are broadly followed. To the best of our knowledge we cited all works that follow these criteria up to November 2016. Fet and Lowe (2000) cited approximately 180 articles pertaining to the genus Buthus, we added approximately 80 new articles, ten of which were published before 1998.

Whenever possible, coordinates for the type localities are provided, using information available in articles or, if not available, finding approximate coordinates with the help of Google Maps (maps.google.com) and the GEOnet Names Server (geonames.nga.mil/gns/html). All coordinates are in WGS 1984 datum, in Latitude/Longitude format, in decimal degrees.

Collections abbreviation codes are listed below. Abbreviation codes follow Sabaj (2016), except for those marked with an asterisk that are not presented there.

ARPC * Andrea Rossi Private Collection, Massa, Italy

CBGP * Centre de Biologie pour la Gestion des Populations (UMR INRA, Cirad, IRD, Montpellier SupAgro), Montferrier-sur-Lez France, France

FKPC * František Kovařík Private Collection, Prague, Czech Republic

MCSNB Museo Civico di Scienze Naturali “Enrico Caffi”, Bergamo, Italy (formerly MSNB)

CRBA Centre de Recursos de Biodiversitat Animal of the Universitat de Barcelona, Barcelona, Spain

MCVR Museo Civico di Storia Naturale di Verona, Verona, Italy

MCZ Museum of Comparative Zoology, Harvard University, Cambridge, U.S.A.

MHNG Muséum d'histoire naturelle de Genève, Geneva, Switzerland

MNCN Museo Nacional de Ciencias Naturales, Madrid, Spain

MNHN Muséum national d’Histoire naturelle, Paris, France

MRSN Museo Regionale di Scienze Naturali di Torino, Turin, Italy

MTAS * Museum of the Turkish Arachnology Society, Ankara, Turkey

MZUF Museo di Storia naturale dell’Università di Firenze, sezione di Zoologia “La Specola”, Florence, Italy

NHMUK Natural History Museum, London; England, UK (formerly BMNH, British Museum of Natural History)

UCAM * Université Cadi Ayyad, Faculte des Sciences Semlalia, “Laboratoire Ecologie et Environnement”, Marrakech, Morocco (formerly Universite Cadi Ayyad, Faculte des Sciences, Semlalia, Depart. BioI., Lab. Ecol. Anim. Terrestre, Marrakech, Marocco)

UGA * University of Ghardaïa, Ghardaïa, Algeria

ZIN Zoological Institute, Russian Academy of Sciences, St. Petersburg, Russia (formerly ZISP)

ZMB Museum für Naturkunde - Leibniz-Institut für Evolutions und Biodiversitätsforschung, Berlin, Germany (formerly ZMBH)

ZMH Biozentrum Grindel und Zoologisches Museum, Hamburg, Germany

Additional abbreviations used in the text:

a.s.l. above sea level

ICZN International Code of Zoological Nomenclature

IOS incorrect original spelling

ISS incorrect subsequent spelling

juv. juvenile or juveniles

M male

F female

MIS misidentification

The type species of the genus Buthus

It is worth mentioning here the taxonomic confusion that surrounded the first Buthus species. Leach first named the genus with Scorpio occitanus Amoreux, 1789 as the type species (Figs 3; 6B). Amoreux (Amoreux 1789b) described this species to accommodate a scorpion from Souvignargues, Occitanie, France. Amoreux also called the same species Scorpio rufus, although he, as the first reviewer, chose the name S. occitanus to be the correct name for the newly described species (Amoreux 1789a). Amoreux also referred to this species as Malpertius’ scorpion and, in his second paper where he gave a full description of the species, included drawings from this author (Maupertuis 1731) (Fig. 6B). It is clear from observing Fig. 6A that Amoreux was well aware of the differences between Scorpio occitanus and S. europaeus Linnaeus, 1758, which he considered a member of the genus Euscorpius Thorell, 1876. Unfortunately, the name S. europaeus Linnaeus, 1758 was subsequently used to refer to three different taxa: (1) S. maculatus De Geer, 1778 (now part of the genus Isometrus Ehrenberg, 1828) (Lönnberg 1898), (2) S. occitanus Amoreux, 1789 (Thorell 1876b), and (3) a Euscorpius species (Fet & Sissom, 2000). This taxonomical confusion was solved by ICZN decision 60 (ICZN 1957), article 1b that suppressed the name europaeus, Linnaeus, 1758 when used in combination with Scorpio, and article 4, that placed Scorpio europaeus Linnaeus, 1758 on the Official Index of Rejected and Invalid Specific Names in Zoology with the number 381. As such, the first author to use the name Buthus europaeus was Thorell in 1876, now a junior synonym of B. occitanus (Amoreux, 1789) (for further details see Braunwalder 1997, Fet et al. 2002).

Figure 6. 

Original drawings of the habits of Scorpio europaeus Linnaeus, 1758 (A) and S. occitanus (B), according to Amoreux (1789b) reproduced from plate I of that work. These images are unfortunately rarely cited, as they are very informative regarding the reasoning of Amoreux while describing the new species.

Finally, although Buthus is considered the nominal genus of Buthidae, Koch (1837) used for the typification of the family, the species Buthus spinnifer Ehrenberg, 1828, which is currently the type species of the genus Heterometrus Ehrenberg, 1828 (Scorpionidae Latreille, 1802), and as such according to ICZN Article 65.2.1. [“type genus was misidentified (that is, interpreted in a sense other than that defined by its type species)] when the family-group name was established”) we will submit to the ICZN a petition to fix the type species of the Buthidae. This was not done by Fet et al. (2000) as the authors probably interpret it as ICZN Article 65.2.3. (“type genus was, when established, based on a type species then misidentified”), which does not necessarily require a ruling by the Commission.

Buthus taxonomy

The revised classification of Sharma et al. (2015) is followed, based on the first phylogenomics study on extant scorpions, which resolved most relationships between scorpion families.

Class Arachnida Lamarck, 1801

Order Scorpiones C. L. Koch, 1850

Suborder Neoscorpiones Thorell & Lindström, 1885

Infraorder Orthosterni Pocock, 1911

Parvoder Buthida Soleglad & Fet, 2003

Superfamily Buthoidea C. L. Koch, 1837

Family Buthidae C. L. Koch, 1837

There are no subfamilies in use within the Buthidae, although many have been proposed and rejected (Fet et al. 2000, 2005). Fet et al. (2005) defined six groups within the Buthidae, and placed Buthus in the Buthus group along with 38 additional genera. The phylogenomic study of Sharma et al. (2015) provided strong support for most groups (although generic level sampling was limited), including the sister group relationship of the Buthus group with the remaining members of the familiy.

Vachon (1952a) considered Androctonus to be the sister taxa to Buthus. However, the only molecular phylogenetic study addressing the relationships between Buthidae genera that includes both genera (Fet et al. 2003), recovered Buthus as the sister taxa to a clade formed by Androctonus and Leiurus, albeit with low support.

There are no taxonomically distinct groups within the genus Buthus, although two “species complexes” are generally recognised. Vachon (1952a, p. 251) suggested that “la «lignée» atlantis se sépare avec facilité de l’ensemble des autres Buthus par la forme élancée des appendices de la queue, de la vésicule et divers autres caractères que nos tableaux de détermination préciseront.” This distinction was retained by Lourenço (2002, 2003) who referred to B. occitanus as a “«complexe de forms»”, and subsequently also adopted it in most subsequent taxonomic works describing new Buthus species. According to Lourenço and Geniez (2005), the two complexes are distinguished by the level of keel development, weaker in the atlantis group. However, Lourenço (2005a) subsequently wrote that B. occitanus from Europe has a weak keel development in contradiction with the previous morphological definition. This statement has been used by subsequent authors. For example, Rossi (2012) described B. elongatus Rossi, 2012 as belonging to the occitanus complex, but if the author had applied Vachon’s definition it should have included it in the atlantis complex because of the slender metasoma (at least its fifth segment) when compared to B. occitanus. None of the published molecular phylogenies of Buthus supports the existence of the atlantis complex. The results of Gantenbein and Largiadèr (2003) grouped B. atlantis within the species included in the occitanus complex. Although Lourenço and Vachon (2004) acknowledged the work of Gantenbein and Largiadèr (2003), they rejected the phylogenetic position of B. atlantis presented in this study.

Recently, based on the information provided by a cox1 mtDNA tree, Sousa et al. (2012) and Pedroso et al. (2013) have defined a series of groups, (see Table 1). These groups have been renamed to facilitate communication and have been expanded to include all available molecular data with reliable species identifications.

Current composition of the groups proposed by Sousa et al. (2012) and Pedroso et al. (2013) based on cox1. To date, only 19 out of the 52 valid named Buthus species (37%) have been analysed. *, assignment based on Sousa (2017).

Group Species Group Species Group Species
boumalenii B. boumalenii occitanus B. atlantis rochati B. bonito
B. elongatus B. draa
mardochei B. elmoutaouakili B. ibericus B. mariefranceae
B. lienhardi B. malhommei B. rochati*
B. mardochei B. maroccanus tunetanus B. chambiensis*
B. parroti B. montanus B. pusillus*
B. occitanus B. tunetatus

In Fig. 7 the current distribution of these five groups in the Maghreb, the Iberian Peninsula, and southern France is presented, based exclusively on specimens with available molecular data. The group assignment does not necessarily correspond to the species assignment in the original publications.

Figure 7. 

Map representing the five phylogenetic Buthus cox1 groups in the Maghreb. Groups as defined by Sousa et al. (2012) and Pedroso et al. (2013), also including cox1 sequences from Gantenbein and Largiadèr (2003), Habel et al. (2012) and Husemann et al. (2012) (redrawing of Figure 1 from Pedroso et al. 2013).

Diagnostic characters used in Buthus taxonomy

Several morphological traits have been used by recent authors as diagnostic characters (in the sense of Winston 1999) for Buthus species.

Colour is of limited utility in Buthus taxonomy, as the underlying colour varies in tones of yellow, orange, reddish or light brown within and between species. Only one species has a fully dark body, Buthus maroccanus Birula, 1903, in some cases even black. Other species also have the mesosoma of a darker colour than the rest of the body. Of greater taxonomic use are colour patterns, such as darker marks, over a lighter background colour, that can be present on the carapace, the mesosoma or the metasoma; the latter being the more informative.

Adult size may also be diagnostic (Fig. 8). Buthus adult body sizes range from 38 to 90 mm (telson included) (the maximum size of 110 mm reported by Vachon (1952a) is presumably a mistake). Most species have a maximum size between 60 and 70 mm in females, and 55 to 70 mm for males (Fig. 8). On the 5% percentiles we have the smaller species of Buthus (less than 45 mm long), and the larger species of Buthus (more than 85 mm for females and 80 mm long for males).

Figure 8. 

Chart and boxplot summary of Buthus species maximum sizes. Only the known maximum size per species is represented. Size information is only available for males from 44 species and females from 43 species. Some individual data might correspond to subadult specimens since this information is not always explicit in species descriptions.

Two additional meristic traits have been used as diagnostic characters, namely the number of rows of granules on the cutting edge of the movable finger of the pedipalp chela, and the number of pectinal teeth, a sexual dimorphic trait. Variation in the number of rows of granules is not very informative because species show an incremental overlap in the numbers of rows, which range from 8 to 14 (Fig. 9). Pectinal teeth number, although carrying a potentially greater amount of information as they have a wider range to vary from, is actually of limited usefulness because of the interspecific overlap (Fig. 10); female counts range from 18–34 and males from 24–37. Pectinal teeth number is also of limited use due to the lack of any information for several species and the unknown range of variability for many other Buthus species (Fig. 10). It should be noted, however, that Buthus elizabethae Lourenço, 2005 is unique in having male pectines that do not overlap in their proximal portion (Lourenço 2005a).

Figure 9. 

Graphical representation of the variation in the number of rows in the movable finger of all Buthus species. ª Species for which the bibliographic ranges are conflicting. * Number of rows in species identified by us from images of the type specimens may be underestimated.

Figure 10. 

Graphical representation of the variation in pectinal teeth number of Buthus species arranged by geographical areas to facilitate comparison. All known Buthus species are represented, although female and male are ordered independently, from smaller to largest. ª Species for which the bibliographic ranges are conflicting.

Trichobothria number and position are not useful for Buthus species diagnosis, as their location shows little variation and have as much intraspecific as interspecific variability (P. Sousa pers. obs.). Conversely, body chaetotaxy (other than trichobothria) is very useful for taxonomy. Vachon (1952a) defined three, albeit diffuse, states in Buthus body chaetotaxy: low (“oligotriche”), high (“polytriche”) and medium (“mésotriche”), and used the number of setae on the fifth segment of the metasoma as example of the ranges: low has three or fewer setae, high more than 5–6 setae and medium four setae, although this latter category was fluid. Confusion can further arise from the fact that these categories apply to the metasoma and the pedipalp, and in the same species these two body parts can have different ranges of chaetotaxy. Nevertheless, this is a useful trait, and one that needs to be explicitly stated in species descriptions to avoid misinterpretations. The chaetotaxy of the leg tarsi and mesosoma terguites is also useful.

Most other diagnostic traits in use for Buthus species are found in the metasoma and the pedipalp chela.

The length/width ratio of the first metasomal segment, which is typically square in most species but can be elongated or sturdy in certain species, is informative. This ratio is also applied to the fifth metasomal segment, and Vachon (1952a) further compared the ratios of the first and second segment, and sometimes even the third segment. The number of keel rows in the metasoma segments is also useful, with special attention paid to the presence, and in some cases the relative length (Vachon 1952a), of the median lateral keel in the second, third and fourth segments. Both the degree of development of the inferior median keels of the five segments (except perhaps the forth), and the existence of larger granules may also be used for taxonomic purposes. The number of lateral lobes in the anal arch, either two or three (the latter only in B. atlantis Pocock, 1889 and B. lourencoi Rossi, Tropea & Yagmur, 2013) may be misleading because in some species, or even specimens (Vachon 1952a), a third smaller lobe may be present between the two larger lobes, which has been interpreted as a third state (e.g. Sadine et al. 2015). For instance, Lourenço and Qi (2006) state that in Buthus mariefranceae Lourenço, 2003 the anal arch may sometimes have 3 lobes but this is mentioned neither in the original description nor in Vachon's descriptions. The relationship between the length of the aculeus and the length of the vesicle that form the Telson are also used in Buthus taxonomy. In most species, the aculeus is shorter than the vesicle, or as long as the vesicle at most. For a few species the aculeus is clearly shorter than the vesicle and for another handful of species, the aculeus is clearly longer. The states are defined here using a 10% difference threshold, but other authors have used a 5% difference. This ratio is correlated with the shape of the aculeus, also in use, which can be more or less curved.

The shape of the pedipalp chelae in Buthus taxonomy has gained increased usage in recent years. The shape can be approximated by using the length to width ratio of the chela, which reflects its specific robustness or slenderness. However, in many Buthus species the chela shape is sexually dimorphic, a trait that was first used in a species key by Kovařík (2006), although its use goes back at least to Vachon (1952a). As a measure of sexual dimorphism the pedipalp chelae has three possible states: 1) no sexual dimorphism (male = female); 2) slender chela in male (male > female); 3) chela of male more robust (male < female). There is data available for 29 species (56% of the known species), and from these we can estimate that 38% do not have sexual dimorphism (+/- 10% threshold as a cut-off point), 52% of species have males with slender chelae and only 3 species (10%) show males with more robust chelae (Fig. 11). Interestingly, although chelae play a role in mating, defense, and as a sensory organ (van der Meijden et al. 2012), their prime importance in prey capture and handling (Polis 1990) may limit the slendering of female chelae, as these may be more prone to breakage (van der Meijden et al. 2012). For three of the 39 species, the available chelae data was contradictory, and they are further discussed below. Another useful pedipalp trait is the interrupted dorso-median keel of the patella in Buthus rochati Lourenço, 2003 (Vachon 1952a).

Figure 11. 

Graphical representation of the relation between female and male pedipalp chela aspect ratio in Buthus species. Only those species with available data from both sexes were plotted. The grey area represents species without perceived sexual dimorphism. Species with males above that area have slender pedipalp chela than females, while species with males plotted below have more robust pedipalp chela than females.

Several partial keys have been published over the years to assist Buthus species identification. However, due to the high rate of new species description (Fig. 5), they have become incomplete and even outdated in their taxonomy (Birula 1903, Vachon 1952a, Lourenço 2003, Lourenço and Vachon 2004, Kovařík 2006, Rossi 2012, Rossi et al. 2013, Teruel and Melic 2015). There are keys available for Morocco and the Maghreb by Birula (1903, only four species), Vachon (1952a, ten species in the Moroccan Buthus key), Lourenço (2003, 10+1 species), and Kovařík (2006, Tunisia, four species); for North Africa (excluding Morocco) by Rossi et al. (2013, 13 species), and for the Iberian Peninsula by Lourenço and Vachon (2004), Rossi (2012), and Teruel and Melic (2015).

Unfortunately, the identification of the majority of Buthus species remains difficult, in part because of the limited number of diagnostic characters and the incomplete knowledge regarding their intraspecific variation. Sexual dimorphism of pedipalp chelae is a promising trait, but for many species the male or female is still undescribed, which limits its applicability. Authors are urged to mention the variation on all the traits mentioned here (see Rossi et al. 2013 for a nice example) in future Buthus species descriptions.

Confirming the fast pace of new Buthus species descriptions, a new species, Buthus danyii Rossi, 2017 was published from Ghana (Rossi 2017) while this study was under revision. As such we were unable to include this species in the present update.

It is hoped that the present catalogue will facilitate a more precise, informative and comparative description of future species. Buthus are an important component of the scorpions’ fauna of North Africa and Western Europe, but it is only now becoming apparent that they are also diverse in the southern Sahara Desert, an area that should be prioritized in future surveys of Buthus scorpions.

Catalogue

Buthus Leach, 1815

Buthus: Leach 1815: 391; Latreille 1817: 310; Gervais 1844b: 203; Peters 1861 (part): 513; Thorell 1876a (part): 82; Thorell 1876b: 7; Simon 1879: 95–96; Karsch 1886 (part): 77; Pocock 1890 (part): 122; Karsch 1891: 18; Kraepelin 1891 (part): 35–42; Pocock 1893 (part): 312; Kraepelin 1895 (part): 79–80; Laurie 1896b: 131; Lönnberg 1897b (part): 194; Kraepelin 1899 (part): 9; Pocock 1900a (part): 13; Simon 1910: 67–68; Birula 1917a (part): 20–24, 164; Birula 1917b (part): 55; Pavlovsky 1924 (part): 77; Kastner 1941 (part): 230; Vachon 1948a: 206–208; Vachon 1949a: 155–162; Vachon 1952a: 155, 241–246, fig. 579; Vachon 1963b: 164, fig. 10; Bücherl 1964: 57; Stahnke 1972: 132, fig. 20; Vachon 1974: 906; Levy and Amitai 1980: 14–15; Francke 1985: 6, 15; Sissom 1990: 101; Nenilin and Fet 1992: 17; Kovařík 1998: 106; Fet and Lowe 2000: 91; Lourenço 2016b: 3–4.

Type species

(by original designation). Scorpio occitanus Amoreux, 1789 [=Buthus occitanus (Amoreux, 1789)].

Etymology

Leach did not provide an explanation for his selection of the genus name. A search on the original usage of the word may shed some light on the intended meaning. Buthus is the Latin form of the Greek name βοῦθος (Bouthos), an unusual name of a winning athlete of the ancient Pythian Games, mentioned by Hesychius and Aristotle (Müller 1848, Christesen 2007). The name was more familiar in antiquity when used in the adage “Βοῦθος περιφοιτᾷ”, translated to the Latin as “Buthus obambulat”, which translates into “Buthus who wanders”, which apparently was applied to stupid and simple people (Müller 1848, Christesen 2007). In Hofmann et al. (1698) the entry for Buthus reads “athleta nobilis, qui bovem integrum unô die devorare solebat; unde natum proverbium in edaces, Buthus obambulat” which roughly translates to “a noble athlete, who used to devour a great ox in a day, and who gave rise to the proverb, Buthus obambulat”. Noël (1824) entry for Buthus also refers to an athlete that devoured an ox in a single day, and that this voracity was the origin of the proverb “Buthus obambulat”, which according to the author refers to gluttony. Interestingly Noël also states that Buthus, in combination with “βῦς, θúιεν (thuein)”, also refers to sacrifice. This opinion shares roots with the meaning of two other words with similar etymology, būthysĭa (used by Nero) that translates to “sacrifice of an ox” and būthytēs (used by Pliny the Elder), that translates to a “sacrificed ox”, according to the Gaffiot Latin-French dictionary (Various 2016). Recently Dupré (2016) reached a similar conclusion, although he states that Buthus originates from the composition of the Greek word “Gr. bous, ox; - thouéin [Greek suffix?], killer”. Potentially therefore, Buthus refers to a stupid or voracious animal, an ox killer or to a sacrifice of an ox, from the latter two we can interpret it as a powerful and dangerous animal. In our opinion the later makes more sense and agrees well with what was known at the time about the potent venom of Buthus scorpions. As such, it is our opinion that Buthus is a singular masculine Latin word (of Greek origin), which Leach intended as homage to an ancient hero (a trend at that time), and that refers to an animal so venomous that it could kill an ox.

Distribution

AFRICA: Algeria, Cameroon, Chad, Egypt, Eritrea, Ethiopia, Guinea, Ivory Coast, Libya, Mali, Mauritania, Morocco (including Western Sahara), Niger, Senegal, Somalia, South Sudan, Sudan, Tunisia. ?Guinea-Bissau, Nigeria, Burkina Faso, ?Gambia, Ghana, ?Djibouti. ASIA: Cyprus, Israel, Jordan, Yemen. ?Iraq, ?Lebanon, ?Saudi Arabia, ?Turkey. EUROPE: France, Italy (Sicily), Spain, Portugal. ?Malta, ?Greece (Corfu, Thessaly). All currently valid records of Buthus species per country are presented in Table 2. Figure 10 offers an additional zoom to the most diverse region of Buthus species diversity, the Maghreb.

List of the countries for which there are valid records of the occurrence of Buthus species. The ID corresponds to the numbers used in Figures 1 and 10, and on the Catalogue. C.A.R. is the abbreviation of the Central African Republic.

ID Taxa France Italy Portugal Spain Algeria Cameroon C.A.R. Chad Djibouti Egypt Eritrea Ethiopia Guinea Lybia Mauritania Morocco Niger Senegal Somalia S. Sudan Sudan Tunisia Cyprus Israel Jordan Yemen Total (by taxa)
01 B. adrianae X 1
02 B. albengai X 1
03 B. amri X 1
04 B. atlantis X 1
05 B. aures X 1
06 B. awashensis X 1
07 B. barcaeus X 1
08 B. berberensis X X X X 4
09 B. bonito X 1
10 B. boumalenii X 1
11 B. brignolii X 1
12 B. centroafricanus X 1
13 B. chambiensis X 1
14 B. confluens X 1
15 B. draa X 1
16 B. dunlopi X 1
17 B. duprei X 1
18 B. egyptiensis X 1
19 B. elhennawyi X X 2
20 B. elizabethae X X 2
21 B. elmoutaouakili 1 1
22 B. elongatus X 1
23 B. hassanini X 1
24 B. ibericus X X 2
25 B. intermedius X 1
26 B. intumescens X 1
27 B. israelis X1 X 2
28 B. jianxinae X 1
29 B. karoraensis X 1
30 B. kunti X 1
31 B. labuschagnei X 1
32 B. lienhardi X 1
33 B. lourencoi X 1
34 B. malhommei X 1
35 B. mardochei X 1
36 B. mariefranceae X 1
37 B. maroccanus X 1
38 B. montanus X 1
39 B. nigrovesiculosus X2 1
40 B. occidentalis X 1
41 B. occitanus X X 2
42 B. orientalis X 1
43 B. paris X X X 3
44 B. parroti X 1
45 B. prudenti X 1
46 B. pusillus X 1
47 B. rochati X 1
48 B. saharicus X 1
49 B. tassili X X 2
50 B. trinacrius X3 1
51 B. tunetatus X X X X 4
52 B. yemenensis X 1
Total (by country) 1 1 1 4 6 1 1 2 1 5 1 2 1 4 1 17 1 2 1 1 2 4 1 1 1 2

Remarks

There are several old records of Buthus, marked with a question mark above, which have never been found again (independently of the material in which they were based being lost or not). As such, many have not been checked since the genus was reduced in scope by Vachon (1949), or those localities remain doubtful because no Buthus has been collected there since. This is of special significance in countries like Greece and Turkey that have been in recent years reasonably well prospected. Type specimens for several Buthus species described early on were not designated or have since become lost, but this does not necessarily represent a taxonomic problem. For example B. occitanus has no type specimen (Fet and Lowe 2000), but its type locality is well established and no other Buthus species occurs nearby. In this case the designation of a neotype is not justified under the ICZN (article 75.2). However, this is not the case for other species that have neither type specimens nor localities, and that we will further discuss below.

Buthus adrianae Rossi, 2013

Buthus adrianae: Rossi 2013: 188–191, fig. 1–2; Rossi, Tropea and Yağmur 2013: 3; 5, 8.

Type material

1 adult M holotype (MCSNB N° 14011), El-Hamam (30.8300°, 29.3150°), Alexandria, Egypt. Paratypes: 1 adult M and 1 adult F (ARPC), same locality.

Distribution

known only from the type locality.

Buthus albengai Lourenço, 2003

Buthus albengai: Lourenço 2003: 902–904, fig. 70–74; Lourenço and Geniez 2005: 5; Aboumaâd et al. 2014: 6; Touloun et al. 2014: 76; Lourenço 2016b: fig. 3.

Type material

1 F holotype (MHNG), Ito Plateau (approx. 33.51°, -5.3°), Ifrane, Morocco. Paratypes: 3 F, same locality; 2 juv., Ifrane (Cedars woods); 1 M, 1 F juv., region north of Kenifra (all in MHNG).

Distribution

known from an area in Morocco that extends from ifrane to Kenifra.

Remarks

It is one of the largest known Buthus species. Records by Habel et al. (2012) south of the High-Atlas are most likely misidentifications.

Figure 12. 

Map of North Africa Maghrebian Buthus species’ type localities (numbers according to the Catalogue and Table 2) whenever known or the best possible approximation at present.

Buthus amri Lourenço, Yağmur & Duhem, 2010

Buthus amri: Lourenço, Yağmur and Duhem 2010: 96–99, fig. 1–5; Lourenço 2013: 65; Lourenço and Rossi 2013: 9; Amr 2015: 186.

Type material

1 M holotype (MNHN), Wadi Rum Desert (29.5363°, 35.4136°), Aqaba, Jordan. Paratypes: 1 adult F (MNHN), 2 adult F, 1 subadult F, 2 subadult M, 3 juv. (MTAS), all from the same locality.

Distribution

known only from the type locality.

Buthus atlantis Pocock, 1889

= Tityus tenuimanusBanks 1910: 189 (synonymized by Lourenço and Francke 1984: 428). 1 F holotype (MCZ), Buena Vista Lake, California, USA (incorrect locality).

Buthus atlantis: Pocock 1889b: 340–341, pl. XV, fig. 4; Birula 1896: 244; Kraepelin 1891: 197, 199; Birula 1903: 107–108; Werner 1932: 300–305; Vachon 1949a: 162–169,fig. 345, 347, 349, 351, 354, 355, 357–362; Vachon 1952a: 254–255, fig. 345, 347, 349, 351, 354, 355, 357–362; Malhomme 1954: 25; Bücherl 1964: 57; Pérez 1974: 22; Levy and Amitai 1980: 15; EIHennawy 1992: 98, 119; Kovařík 1998: 106; Fet and Lowe 2000: 91; Lourenço 2005a: 233–234; Lourenço and Geniez 2005: 5; Stockmann and Ythier 2010: 360–361; Stockmann 2015: fig. 5.

Buthus Atlantis (sic): Aboumaâd et al. 2014: 5.

Buthus atlantis atlantis: Vachon 1949a: 166–168; Vachon 1952a: 252, 254; Le Corroller 1967: 63; Kovařík 1995: 20; Fet and Lowe 2000: 91; Lourenço 2003: 883–885, fig. 18–22; Touloun et al. 2001: 2; Gantenbein and Largiadèr 2003: 120, 122; Touloun 2012: 43, fig. 9A.

Buthus occitanus atlantis: Kraepelin 1899: 26–27; Werner 1934b: 86–87, fig. 5; Schenkel 1949: 186–187.

Buthus (Buthus) atlantis: Birula 1910: 145; Birula 1917a: 213, 223.

Tityus tenuimanus: Cox 1921: 12; Ewing 1928: 22; Mello-Leitão 1931: 121, 140; MelloLeitão 1939: 60, 64, 71; Comstock 1940: 27; Mello-Leitão 1945: 308; Gertsch and Soleglad 1966: 2; Hjelle 1972: 28; Lourenço and Francke 1984: 427, fig. 10–12.

Type material

1 F holotype (NHMUK), Essaouira (formerly Mogador) (approx. 31.49°, -9.76°), Morocco.

Distribution

known to occur only in sandy dune habitats close to the Atlantic Ocean in Morocco, between Essaouira and Agadir.

Remarks

It is the largest known Buthus species.

Buthus aures Lourenço & Sadine, 2016

Buthus aures Lourenço and Sadine 2016: 14–17, fig. 4–13.

Type material

1 M holotype (MNHN), Batna region (35.5319°, 5.9194°), Aurès Mountains, Algeria. 1 M paratype (UGA), same locality.

Distribution

known only from the type locality.

Buthus awashensis Kovařík, 2011

Buthus awashensis: Kovařík 2011: 1–3, 5–8, fig. 5–16.

Buthus occitanus (MIS): Kovařík and Whitman 2005 (part): 106.

Type material

1 M holotype (FKCP), Metahara (approx. 8.900°, 39.900°), Oromia, Ethiopia. Paratypes: 34 M, 34 F, 36 juv. (FKCP), all from the same locality; 1 M (FKCP), Dire Dawa, Ethiopia.

Distribution

know only from two Ethiopian localities, more than 200 km apart.

Remarks

The pedipalp chela length-to-width ratio given by the author for the type material suggest that some animals exhibit sexual dimorphism while others do not. If this is true, the utility of this ratio as a diagnostic character in Buthus would be compromised. Alternatively, it may be due simply to the use of immature specimens.

Buthus barcaeus Birula, 1909

Buthus occitanus barcaeus: Birula 1909: 508–511. fig. A, C; Borelli 1914a: 155–156; Borelli 1924: 5–7; Borelli 1928: 351; Caporiacco 1932: 395; Borelli 1934: 169; Caporiacco 1937: 345; Pérez 1974: 23; Levy and Amitai 1980: 16; El-Hennawy 1992: 98, 120; Kovařík 1998: 106; Fet and Lowe 2000: 95; Kovařík 2002: 5.

Buthus (Buthus) occitanus barcaeus: Birula 1910: 156; Birula 1917a: 223.

Buthus barcaeus: Kovařík 2006: 3, fig. 6; Kaltsas et al. 2008: 215; Lourenço and Cloudsley-Thompson 2012: 15; Lourenço and Simon 2012: 11; Rossi, Tropea and Yağmur 2013: 3–5, 7.

Type material

4 M, 1 F juv., syntypes (ZIN), Barca (approx. 32.48°, 20.83°), 5 km E from Benghazi (Cyrenaica), Libya.

Distribution

know from several localities along the Mediterranean coast of Libya.

Remarks

The specimens present in the MNHN (F n° 4896), captured in Barca and identified by Vachon in 1974, have no intermediary keel on the fourth metasomal segment, which casts doubts about the use of the character in the diagnosis of B. barcaeus.

Buthus berberensis Pocock, 1900

= Buthus occitanus zeylensisPocock 1900b: 56–57 (synonymized by Levy and Amitai 1980: 16). 1 F holotype (NHMUK), Zeyla (northwestern Somaliland), Somalia.

Buthus occitanus berberensis: Birula 1903: 106–107; Birula 1909: 510; Birula 1910: 118; Kraepelin 1903: 558; Borelli 1904:·2–3; Giltay 1929: 196; Moriggi 1941: 84; Lamoral and Reynders 1975: 505; Levy and Amitai 1980: 16; EI-Hennawy: 1992: 98, 120; Kovařík 1998: 106; Fet and Lowe 2000: 95; Kovařík 2003 (part): 138.

Buthus occitanus zeylensis: Kraepelin 1903: 558–559; Borelli 1919: 363; Borelli 1931: 218; Caporiacco 1936: 137; Moriggi 1941: 84; Lamoral and Reynders 1975: 505–506; EI-Hennawy 1992: 98, 121–122; Kovařík 1998: 106.

Buthus berberensis: Lourenço 2008: 46; Kovařík 2011: 4–6.

Buthus (Buthus) occitanus berberensis: Birula 1917a: 123.

Type material

1 M holotype (NHMUK), Somaliland, Somalia.

Distribution

know from Djibouti, Eritrea, Ethiopia and Somalia, although Birula (1903) recorded toponyms that are old and difficult to map.

Remarks

Lourenço (2008) stated that B. o. zeylensis might be a distinct species from B. berberensis, but that further material was required to confirm this possibility. Kovařík (2011) considered B. o. zeylensis a colour morph of B. berberensis present in juveniles and some males.

Buthus bonito Lourenço & Geniez, 2005

https://science.mnhn.fr/institution/mnhn/collection/rs/item/rs8669 https://science.mnhn.fr/institution/mnhn/collection/rs/item/rs8670

= Buthus sabulicolaTouloun 2012: 46, 48–58, fig.10, 13, 14 (Syn. n.). 1 F holotype (MNHN), Khnifiss lagoon, Tan-Tan Province, Morocco.

Buthus bonito: Lourenço and Geniez 2005: 1–5, fig. 1–8, 10; Touloun et al. 2008: 3–4, fig.1; Stockmann and Ythier 2010: 362–363; Pedroso et al. 2013: 300; Aboumaâd et al. 2014: 6; Touloun et al. 2016: 880, fig. 2D.

Type material

1 M holotype (MNHN N° RS8669), Khnifiss lagoon (approx. 27.93°, -12.34°), Tarfaya, Morocco. Paratypes: 2 F (MNHN N° RS8670), from the same locality.

Distribution

known from the Atlantic coast of Morocco south of Tan-Tan extending almost to Dakhla in the Western Sahara (Touloun et al. 2016).

Remarks

Although the type material of B. sabulicola was collected in 2002 by Touloun, Stockmann and Slimani, the species was not formally described until the publication of the PhD thesis of Oulaid Touloun in 2012. The type specimens of B. bonito and B. sabulicola are from the exact same locality, the Khnifiss lagoon, and both descriptions are almost identical. Touloun et al. (2016), probably by mistake, indicated that the fifth metasomal segment and telson are darkened in B. bonito, but the trait does not appear in the accompanying figure.

Buthus boumalenii Touloun & Boumezzough, 2011

Buthus boumalenii: Touloun and Boumezzough 2011a: 183–186, fig. 2–7; Pedroso et al. 2013: 300; Aboumaâd et al. 2014: 6; El Hidan et al. 2016: 4.

Type material

1 F holotype (UCAM), Tineghir (approx. 31.366°, -5.905°), Boumalene, Morocco. Paratypes: 1 M (UCAM), 1 M, 1 F (MNHN, N° RS8891), all from the same locality.

Distribution

known only from the Boumalne region of Morocco (El Hidan et al. 2016).

Remarks

This species is the only known representative of a phylogenetic lineage present east of the High Atlas Mountains of Morocco. Because of its phylogenetic uniqueness, the conservation of this species should have top priority.

Buthus brignolii Lourenço, 2003

Buthus brignolii: Lourenço 2003: 905–907, fig. 75–79; Rossi and Tropea 2016a: 4.

Type material

1 F holotype (MHNG), Djebel Meidob (approx. 15.21°, 26.44°), Darfur, Sudan.

Distribution

known only from the type locality.

Remarks

it is one of the four known “inland island” species of Buthus that have been found in the Mountainous regions in the heart of the Sahara Desert.

Buthus centroafricanus Lourenço, 2016

Buthus centroafricanus: Lourenço 2016a: 73–77, fig. 1–11.

Type material

1 M holotype (MNHN), Between Bria and Yalinga (as Jalinga) (approx. 6.52°, 22.62°), Province Haute-Kotto, Central African Republic.

Distribution

known only from the type locality.

Buthus chambiensis Kovařík, 2006

Buthus chambiensis: Kovařík 2006: 1–3, fig. 2–5; Rossi, Tropea and Yağmur 2013: 3, 7.

Type material

1 M holotype (FKCP), Djebel Chambi Mountain (approx. 35.17°, 8.56°), Kasserine Province, Tunisia. Paratypes: 1 M juv., 2 F, 1 juv., all from the same locality.

Distribution

known only from the type locality.

Buthus confluens Lourenço, Touloun & Boumezzough, 2012

Buthus confluens: Lourenço, Touloun and Boumezzough 2012: 22–24, fig. 1–11; Touloun et al. 2014: 76–77.

Type material

1 M holotype (MNHN N° RS8919), Alhamra (35.39529°, -05.37181°), Tétouan, Morocco. Paratypes: 1 F (MNHN N° RS8920), 1 M (UCAM), all from the same locality.

Distribution

known from several localities in the Tingitana Pensinsula of Morocco, but also further to the south.

Remarks

Based on the colour pattern and pigmentation, the original authors suggested that B. confluens was the closest phylogenetic relative in Morocco to B. ibericus, from the Iberian Peninsula. However, because the presence of three dark bands on the metasoma is shared among several Buthus species, this claim should be further confirmed with additional data.

Buthus draa Lourenço & Slimani, 2004

Buthus draa: Lourenço and Slimani 2004: 166–169, fig. 1–7; Lourenço, Sun and Zhu 2009: p. 72, fig. 2–6; Stockmann and Ythier 2010: 362–363; Touloun and Boumezzough 2011b: 186; Habel et al. 2012: 2, 4; Sousa et al. 2012: 68–69; Pedroso et al. 2013: 300; Yang et al. 2013: 2; Aboumaâd et al. 2014: 6; El Hidan et al. 2016: 4.

Buthus occitanus tunetanus neeli (MIS): Touloun et al. 1999: 1–2.

Buthus tassili (MIS): Touloun 2012: 37, 40–41, fig.7.

Type material

1 M holotype (UCAM), Taznakht (30.51853°, -7.02595°), Ouarzazate, Morocco. Paratypes: 1 M, 2 F (UCAM), 1 M, 1 F (ZMH), 2 M, 1 F (MNHN), all from the same locality; 1 M (ZMH), Aït Bassou; 2 F juv. (ZMH), Aït Ounzar Oulad Aissa; 1 M (ZMH), near Agdez; 1 M (ZMH), Oulad Hlal. The ZMH accession number for all paratypes is A7/03.

Distribution

B. draa can be found in the upper part of the Draa River, probably at elevations below 1500 m a.s.l.

Remarks

B. draa shares with B. tassili and B. nigrovesiculosus the presence of a darkened fifth metasoma segment and telson.

Buthus dunlopi Kovařík, 2006

Buthus dunlopi: Kovařík 2006: 2–3, 6, fig. 7–8; Rossi, Tropea and Yağmur 2013: 5, 7.

Type material

1 M holotype (FKCP), Remada (approx. 32.31°, 10.39°), Tataouine, Tunisia. Paratypes: 1 M, 3 F (FKCP), same locality.

Distribution

known only from the type locality.

Buthus duprei Rossi & Tropea, 2016

Buthus duprei Rossi and Tropea 2016b: 25–28, fig. 1–12.

Type material

1 M holotype (MCVR), Port Sudan (approx. 19.59°, 37.19°), Sudan. Paratype: 1 M juv. (ARPC N° 0809), same locality.

Distribution

known only from the type locality.

Buthus egyptiensis Lourenço, 2012

Buthus egyptiensis: Lourenço and Cloudsley-Thompson 2012: 12–16, fig. 1–7; Lourenço and Simon 2012: 12; Rossi 2013: 191–192; Rossi, Tropea and Yağmur 2013: 4, 7.

Type material

1 F holotype (ZMH N° A20/12), Siwa (approx. 29.17°, 25.46°), Egypt.

Distribution

known only from the type locality.

Remarks

One of the four known “inland island” species of Buthus that have been found within the Sahara Desert, although in this case from an Oasis. It is also one of the largest known Buthus species.

Buthus elhennawyi Lourenço, 2005

Buthus elhennawyi: Lourenço 2005b: 246–249, fig. 1–7; Lourenço and Leguin 2012: 8.

Type material

1 M holotype (ZMH N° A42/05), Fété-Olé (as Félé-Olé) (16.233°, -15.099°), Ferlo, Senegal. Paratype: 1 M (MNHN N° RS8637), Rosi (as Rossi), Niger.

Distribution

this species is known from Niger and Senegal, from a single locality in each country, which are almost 2,000 Km apart.

Remarks

We used the location of Fété-Olé given in Vincke et al. (2010), a locality that has been part of long term ecological studies, to map this locality, instead of the original spelling “Félé-Olé”.

Buthus elizabethae Lourenço, 2005

Buthus elizabethae: Lourenço 2005a: 230–235, fig. 1–12, Lourenço 2005b: 249.

Type material

1 M holotype (ZMH N° A36/05), S.W. of Gaoual (approx. 11.71°, -13.22°), Boké, Guinea. Paratypes: 1 F (ZMH N° A37/05), same locality; 1 M, 1 F (MNHN), Niokolo-Koba National Park, Senegal.

Distribution

this species is known from Guinea and Senegal,

Remarks

Given the geographical proximity, it is possible that the Buthus material reported to have been found in Guinea-Bissau might very well correspond to this species. Unfortunately the Guinea-Bissau material was lost in a fire, and hence only newly collected material could confirm this possibility.

Buthus elmoutaouakili Lourenço & Qi, 2006

Buthus elmoutaouakili: Lourenço and Qi 2006: 288–291, fig. 1–11; Habel et al. 2012 (part): 2, 3; Husemann et al. 2012 (part): 2, 4–5; Touloun and Boumezzough 2011b: 11–12, fig. 2C; Aboumaâd et al. 2014: 6.

Buthus occitanus mardochei alluaudi: Vachon 1949c: 363–367, fig. 409–416; Vachon 1952a: 291–295, fig. 409–416; Le Corroller 1967: 63; Pérez 1974: 23; Touloun 2012: 39, 57.

Type material

1 M holotype (ZMH N° A18/06), Ait Baha (approx. 30.07°, -9.15°), Chtouka Aït Baha, Morocco.

Distribution

this species seems to be widely distributed across the western portion of the Anti-Atlas, although some misidentifications with Buthus parroti cannot be excluded.

Remarks

According to ICZN article 45.5, Vachon’s (1949) infrasubspecific name is unavailable. Although the name was published before 1961, it was only used as infrasubspecific by all subsequent authors.

Buthus elongatus Rossi, 2012

Buthus elongatus: Rossi 2012: 273–278, fig. 1–6; Teruel and Melic 2015: 5–9.

Type material

1 adult M holotype (MZUF N° 1432), Sierra Blanca (36.533°, 4.900°), Marbella, Malaga Province, Spain. Paratypes: 1 adult F (ARPC), same locality; 1 M, 1 F (MZUF N° 875), Playa del Alicate (36.499°, 4.818), Marbella, Malaga Province, Spain.

Distribution

this species is known from the southern Iberian Mediterranean coast, close to Marbella.

Remarks

The second locality given by Rossi as Alicante (sic), had a typographic error, as the coordinates given by the author, together with their map in Fig. 7, provide sufficient evidence for the correct mapping of this locality. Both localities are under severe anthropomorphic pressure.

Buthus hassanini Lourenço, Duhem & Cloudsley-Thompson, 2012

Buthus hassanini: Lourenço et al. 2012: 319–321, 323, fig. 35–42.

Type material

1 F (MNHN N° RS8927), Biti Tehëc (approx. 17.187°, 22.288°), Ennedi Plateau, Chad.

Distribution

known only from the type locality.

Remarks

Another of the four known “inland island” species of Buthus that have been found in a Mountainous region in the heart of the Sahara Desert. The type locality was pinpointed following the map provided by the authors (fig. 90).

Buthus ibericus Lourenço & Vachon, 2004, nomen protectum

= Buthus halius (C. L. Koch, 1839) (nomen oblitum) (comb. n., syn. n.). Holotype lost according to Fet and Lowe (2000), Portugal.

Buthus ibericus Lourenço & Vachon, 2004: 88–91, fig. 31–42, Fernández 2004: 222; Teruel and Pérez-Bote 2005: 273–276, fig. 1; Armas and González-Moliné 2009: 553–554; Fet 2010: 4; Sousa et al. 2010: 207; Rossi 2012: 274–275, 277–278; Pedroso et al. 2013: 300; Teruel and Melic 2015: 6–9.

Androctonus halius C. L. Koch 1839a: 69–70, pl.CLXIII, fig. 383; Gervais 1844a: 43; C. L. Koch 1850: 90; Simon 1879: 96.

Buthus occitanus (MIS): Berejano and Pérez-Bote 2002: 59.

Type material

1 M holotype (MNHN N° RS8605), San José del Valle (36.6247°, -5.6646°), Cádiz, Spain. Paratypes: 2 F (MNHN, N° RS8654), 1 M, 2 F (CRBA, N° CRBA-21826), and 2 F (MNCN N° 20.02/14857), all from the same locality.

Distribution

this species seems to have a wide distribution range in the western part of the Iberian Peninsula, although the limits of its distribution remain poorly defined.

Remarks

B. ibericus was first described from Spain and subsequently reported for Portugal (e.g. Sousa et al. 2010, Rossi 2012). Simon (1879) synonymized Androctonus halius with B. occitanus based on its type locality, which was wrongly stated to be Spain (page 98), because Simon considered B. occitanus (as B. europaeus) to be the only good species in Spain. Koch’s original description is not by itself enough to synonymize both species. However, Koch’s fig. 383 illustration of Androctonus halius includes a basal lobe in the movable finger (Fig. 13a and B), which is the key diagnostic character for B. ibericus (Lourenço and Vachon 2004, Rossi 2012), together with its type locality (Portugal), supports this synonymy. Nonetheless, according to the I.C.Z.N. article 23.9, the junior synonym can remain valid to maintain taxonomic stability. To our knowledge the name B. halius has not been used since 1879 (article 23.9.1.1), and more than 25 works have been published in the past 12 years by more than 10 authors using the name B. ibericus (article 23.9.1.2). Not all works are cited here because they are neither taxonomic nor faunistic. As such we propose to maintain as valid the junior synonym B. ibericus (nomen protectum) according to prevailing usage (article 23.9.1), and to consider the senior synonym B. halius a nomen oblitum. Rossi, 2012 (page 278), for B. ibericus, states erroneously “Sexual dimorphism is not noticeable in the chela manus”, which is in contrast to the original descriptions of both Koch (1839) and Lourenço and Vachon (2004) and to the complementary description in Teruel and Pérez-Bote (2005). In both works it can be observed that the male manus is more bulbous than the female’s, which results in the males having a smaller length to width ratio. The type locality of B. ibericus was precisely located following the indications of Iñigo Sánchez, the original collector.

Figure 13. 

Reproduction of C. L. Koch’s 1839 B. halius figure 382 (plate CLXIII). Right pedipalp che-lae detail from three Buthus species published in the same work: A B. paris (as Androctonus clytoneus), fig. 384 (same plate) B B. halius with an arrow pinpointing the basal lobe C B. paris, fig. 352 (pl. CLI). All images were taken as provided by the pdf copy available in the BHL, which was made available by the Ernst Mayr Library of the Museum of Comparative Zoology, Harvard University.

Buthus intermedius (Ehrenberg in Hemprich and Ehrenberg 1829)

Androctonus (Leirus) tunetanus intermedius: Ehrenberg in Hemprich and Ehrenberg 1829: 354; Braunwalder and Fet 1998: 33–34.

Androctonus (Leiurus) tunetanus intumescens (MIS): Kovařík 2006: 10.

Androctonus (Liurus) tunetanus intermedius: Ehrenberg in Hemprich and Ehrenberg 1831: (pages not numbered).

Androctonus occitanus intermedius: Gervais 1844a: 42.

Buthus (Buthus) occitanus intermedius: Birula 1917a: 228.

Buthus occitanus intermedius: Peréz 1974: 23.

Buthus intermedius (Ehrenberg): Lourenço 2008: 46–47.

Type material

1 F (in bad conditions) (ZMB N° 146), Al Luhayyah (as Lohaie), Yemen.

Distribution

Known only from the type locality.

Remarks

Fet and Lowe (2000) considered the locality as probably wrong since at that time no other Buthus had been collected again in Yemen. However, Lourenço's (2008) Buthus yemenensis revalidated Ehrenberg species’ by providing concrete proof for the existence of Buthus species in this country. Doubst about B. intermedius type locality were the only evidence given by Kovařík (2006) for its synonimization with Buthus intumescens (Ehrenberg in Hemprich and Ehrenberg 1829).

Buthus intumescens (Ehrenberg in Hemprich and Ehrenberg 1829)

Androctonus (Leiurus) tunetanus intumescens: Ehrenberg in Hemprich and Ehrenberg 1829: 354; Braunwalder and Fet 1998: 33.

Androctonus (Liurus) tunetanus intumescens: Ehrenberg in Hemprich and Ehrenberg 1831 (pages not numbered); Moritz and Fischer 1980: 316.

Androctonus occitanus intumescens: Gervais 1844a: 42.

Buthus intumescens: Kovařík 2006 (part): 10–11, 15, fig. 20; Kaltsas et al. 2008 (part): 215; Rossi 2013: 191–192; Rossi, Tropea and Yağmur 2013: 3, 6–8.

Type material

1 (sex unknown) (in bad conditions) (ZMB N° 145), Egypt.

Remarks

known only from a single specimen.

Buthus israelis Shulov & Amitai, 1959

Buthus occitanus mardochei israelis: Shulov and Amitai 1959: 219–225, fig. 1–3.

Buthus occitanus israelis: Pérez 1974: 23; Vachon and Kinzelbach 1987: 101; Fet and Lowe 2000: 95; Skutelsky 1995: 46; Skutelsky 1996: 50

Buthus occitanus israelis (Shulov & Amitai, 1959): Levy and Amitai 1980: 16–21, fig. 25–29; EI-Hennawy 1992: 101, 120; Kovařík 2006: 10; Lourenço, Yağmur and Duhem 2010: 96.

Buthus intumescens (MIS): Kovařík 2006 (part): 10–11.

Buthus israelis: Lourenço, Yağmur and Duhem 2010: 96–97; Yağmur, Koç and Lourenço 2011: 29.

Buthus israelis (Shulov & Amitai, 1959): Rossi 2013: 191–192; Rossi, Tropea and Yağmur 2013: 2–3, 6–7.

Buthus occitanus mardochei israelis: Vachon 1966: 211; Kovařík 2006: 10; Lourenço, Yağmur and Duhem 2010: 95.

Buthus occitanus typicus: Bodenheimer 1937: 235.

Type material

holotype lost (sex unknown), Mash’abbe Sade (as Mashavei Sadé) (approx. 31°, 34.78°), Negev desert, Israel.

Distribution

This species has been recorded in Egypt (the Sinai Peninsula) and Israel. Notwithstanding, Kovařík (2006) considered this species to be a junior synonym of B. intumescens.

Remarks

Levy and Amitai (1980) did not designate any neotype when they re-described the species. In contrast to the several infrasubspecific taxa described by Vachon that are unavailable according to the ICZN, B. o. israelis is an available name according to the ICZN article 45.6.4.1 (“a name that is infrasubspecific under Article 45.6.4 is nevertheless deemed to be subspecific from its original publication if, before 1985, it was either adopted as the valid name of a species or subspecies or was treated as a senior homonym”), which is the present case as Levy and Amitai redescribed this taxon before 1985, and thus articles 45.5 and 45.5.1 do not apply. Several authors wrongly report the original authors of the species in parenthesis. The use of parenthesis is only to be made when a species is changed from one genus to another (ICZN article 51.3), which is clearly not the case with B. israelis.

Buthus jianxinae Lourenço, 2005

Buthus jianxinae: Lourenço 2005c: 22–23, fig. 1–12.

Type material

1 M holotype (MNHN N° RS8175), Loka (approx. 4.21°, 30.91°), Equatoria, South Sudan.

Distribution

known only from the type locality.

Buthus karoraensis Rossi & Tropea, 2016

Buthus karoraensis: Rossi and Tropea 2016a: 4–7, fig. 1–13; Rossi and Tropea 2016b: 25.

Buthus occitanus berberensis (MIS): Kovařík 2003 (part): 138.

Buthus occitanus (MIS): Kovařík and Whitman 2005 (part): 106.

Type material

1 M holotype (MZUF N° 610), Karora (17.703°, 38.365°) (small enclave in the Eritrean-Sudan border), Eritrea. Paratypes: 1 M, 4 F (MZUF N° 610); 1 M (MCSNB: N° 12749), 1 F (MCSNB: N° 12748), all from the same locality.

Distribution

known only from the type locality.

Buthus kunti Yağmur, Koç & Lourenço, 2011

Buthus kunti: Yağmur, Koç and Lourenço 2011: 29–33, fig. 1–12.

Buthus europaeus (MIS): Simon 1879: 97.

Buthus occitanus (MIS): Kraepelin 1891 (part): 199.

Buthus sp.: Levy and Amitai 1980: 21.

Type material

1 F holotype (MTAS), Rizokarpaso (Dipkarpaz) (35.58472°, 34.42306°), Karpaz Region, Cyprus. Paratypes: 1 M juv. (MTAS), Zafer. 1 M juv. (MNHN N° RS8892), Güzelyurt.

Distribution

the species is only known from the northern portion of Cyprus.

Remarks

according to Yağmur, Koç and Lourenço (2011), this species is rare in the island.

Buthus labuschagnei Lourenço, 2015

Buthus labuschagnei Lourenço 2015: 22–24, fig. 13–22.

Type material

1 F holotype (MNHN N° RS8992), Zakouma (Zakouma National Park) (approx. 10.89°, 19.82°), Salamat Region, Chad.

Distribution

known only from the type locality.

Buthus lienhardi Lourenço, 2003

Buthus lienhardi: Lourenço 2003: 899–902, fig. 62–69; Stockmann and Ythier 2010: 362–363; Touloun and Boumezzough 2011a: 186; Touloun 2012: 37, fig.5C; Aboumaâd et al. 2014: 6.

Buthus occitanus tunetatus Lepineyi: Vachon 1949: 353–359, fig. 393–400; Vachon 1952a: 281–286, fig. 393–400;

Buthus occitanus tunetatus lepineyi: Malhomme 1954: 29–30; Le Corroller 1967: 63; Peréz 1974: 22; Touloun et al. 1999: 1; Touloun et al. 2001: 2; Touloun 2012: 37.

Buthus occitanus tunetatus (MIS): Touloun 2012: 104, 108.

Type material

1 M holotype (MHNG), Oukaimeden (approx. 31.201°, -7.861°), Marrakech, Morocco. Paratypes: 1 F, 2 juv. (MHNG), same locality.

Distribution

the species is known from a wide range across the High-Atlas Mountains.

Remarks

Vachon (1949) infrasubspecific name is not available as explained previously.

Buthus lourencoi Rossi, Tropea & Yağmur, 2013

Buthus lourencoi: Rossi et al. 2013: 2–3, fig. 3–10.

Buthus occitanus (MIS): Kovařík and Whitman 2005 (part): 106.

Type material

1 adult F holotype (MZUF N° 783), Mellaha (approx. 32.896°, 13.285°), Tripoli, Libya

Distribution

known only from the type locality.

Remarks

The type locality is now part of the large city of Tripoli. It is fairly unlikely that the species still occur within the boundaries of the city given the level of urban development. However, Mellaha, which was originally a military airport, is now the Mitiga International Airport, where large patches of unconstructed ground that may be suitable fot the species still exist.

Buthus malhommei Vachon, 1949

Buthus occitanus malhommei Vachon 1949: 376; Vachon 1952a: 304–308, fig. 433–444; Fet and Lowe 2000: 95; Touloun et al. 2001: 2; Touloun 2012: 35, 104, 108, fig. 5A.

Buthus malhommei: Lourenço 2003: 887–889, fig. 33–38; Stockmann and Ythier 2010: 364–365 (MIS); Sousa et al. 2012: 68–69; Aboumaâd et al. 2014: 5.

Type material

3 M, 3 F, 7 juv., syntypes (MNHN), Mechra ben Abbou (approx. 32.646°, -7.800°), Settat, Morocco.

Distribution

Toulon (2012) greatly expanded the known distribution of this species along the basin of the Oum er Rbia River.

Buthus mardochei Simon, 1878

Buthus mardoche (IOS): Simon 1878: 159–160; Simon 1879: 100; Kraepelin 1891: 199; Birula, 1896: 244

Buthus (Buthus) mardoche (IOS): Birula 1910: 145–146; Birula 1917a: 223 (“dubious species”); Werner 1932: 300–305.

Buthus occitanus mardochei: Vachon 1949c (part): 358–363, fig. 400–408; Vachon 1952a (part): 286–295, fig. 401–408; Malhomme 1954: 28–29; Pérez 1974: 22; Levy and Amitai 1980: 16; El-Hennawy 1992: 98, 120; Kovařík 1995: 20; Gantenbein et al. 1998a: 51; Gantenbein et al. 1998b: 33–39; Kovařík 1998: 106; Fet and Lowe 2000: 96; Touloun et al. 2001: 2; Gantenbein and Largiadèr 2003 (part): 120, 122.

Buthus occitanus mardochei mardochei : Le Corroller, 1967. 63; Touloun 2012: 39, 104, 108, fig.5D.

Buthus mardochei: Lourenço 2003: 889, fig. 39; Stockmann and Ythier 2010: 364–365; Aboumaâd et al. 2014: 5.

Type material

1 F (MNHN N° RS1771, damaged), southern Morocco.

Distribution

this species appears to have a distribution parallel to that of B. atlantis, between Essaouira and Agadir, but is found further inland and away from the Atlantic coast.

Remarks

Vachon (1949d: 358) corrected what he considered Simon's incorrect original spelling of “mardoche” to “mardochei”, since the form was named as a patronym after its collector, Rabbi Mardoché. Nevertheless, it is our understanding that this was an unjustified emendation, because the ICZN article 31.1 admit the use of a noun in apposition as was the case with “mardoche”, however the ICZN article 33.2.3.1 admits the prevalence of this emendation as it continues to be attributed to “the original author and date” and is “in prevailing usage” and as such we refrain from any change to the name. Vachon (1949, 1952) also established that the species occurs roughly between Essaouira and Agadir, but not near the coast where it is replaced by B. atlantis.

Buthus mariefranceae Lourenço, 2003

Buthus mariefranceae: Lourenço 2003: 889–893, fig. 40–46; Lourenço and Qi 2006: 291; Stockmann and Ythier 2010: 364–365; Sousa et al. 2012: 68–69; Touloun 2012: 40; Pedroso et al. 2013: 300; Aboumaâd et al. 2014: 6.

Buthus occitanus mardochei mimeuri: Vachon 1949c: 367–373, fig. 417–425; Vachon 1952a: 295–301, fig. 417–425; Le Corroller 1967: 63; Pérez 1974: 23; Touloun 2012: 40, 57.

Type material

1 F holotype (MHNG), Tan-Tan (approx. 28.43°, -11.1°), Guelmim Region, Morocco. Paratypes: 5 M, 4 F, 2 F juv. (MHNG), Goulimine.

Distribution

this species has a large distribution in Morocco, east and south of the Anti-Atlas Mountain.

Remarks

Vachon (1949) infrasubspecific name is not available as explained above.

Buthus maroccanus Birula, 1903

= Prionurus tingitanus: Pallary 1928a: 350–351, fig. 4 (synonymized by Vachon 1949b: 281). Syntype, sex unknown (MNHN), Rabat, Morocco (Vachon, 1949, 1952).

Buthus occitanus maroccanus: Birula 1903: 106.

Buthus europaeus (MIS): Hirst 1925 (part): 416.

Buthus (Buthus) occitanus maroccanus: Birula 1910: 145; Birula 1917a: 223.

Buthus occitanus maroccanus: Giltay 1929: 196; Werner 1929: 31–32.

Buthus maroccanus: Werner 1932: 299; Werner 1934b: 84; Vachon 1949b: 281–287, fig. 364–371; Vachon 1952a: 255–261, fig. 364–371; Foley 1951: 33; Bücherl 1964: 57; Pérez 1974: 22: l.evy and Amitai 1980: 15; El-Hennawy 1992: 98, 119; Kovařík 1995: 20; Kovařík 1998: 106; Fet and Lowe 2000: 92; Sousa et al. 2012: 68–69; Stockmann and Ythier 2010: 366–367; Aboumaâd et al. 2014: 6..

Buthus marocanus (ISS): Le Corroller 1967: 63.

Type material

3 M, F, syntypes (ZIN), Morocco; 1 specimen syntype (ZIN), locality unknown (Fet and Lowe 2000).

Distribution

all known specimens have been captured in the Rabat Region (approx. 28.43°, -11.10°).

Remarks

it remains the only known Buthus species with a uniformly darkened body.

Buthus montanus Lourenço & Vachon, 2004

Buthus montanus Lourenço and Vachon 2004:84, 86–87, 91, fig. 16–30; Fernández 2004: 222; Fet 2010: 4; Rossi 2012: 274, 277–278; Teruel and Melic 2015: 5–9.

Type material

1 M holotype (MNHN N° RS8604), Sierra Nevada (between Puerto de la Ragua and Cerro Pelado) (approx. 37.11°, -3.14°), Granada Region, Spain. Paratypes: 1 M, 3 F (MNHN N° RS8653), same locality.

Distribution

known only from the type locality.

Buthus nigrovesiculosus Hirst, 1925, stat. n.

Buthus europaeus nigrovesiculosus: Hirst 1925: 416.

Buthus occitanus nigrovesiculosus: Pérez 1974: 22; Fet and Lowe 2000: 96.

Type material

1 M (adult?), 1 juv., syntypes (NHMUK), Boste (approx. 23.79°, -15.68°), Rio de Oro (Western Sahara), now Morocco.

Distribution

known only from the type locality.

Remarks

Hirst identified this North African species as a subspecies of B. occitanus, but as currently circumscribed, B. occitanus does not occur in North Africa (Gantenbein and Largiadèr 2003, Sousa et al. 2012). The original description of B. nigrovesiculosus suggests morphological similarities to B. draa and B. tassili Lourenço, 2002. These three species have a dark, almost black, fifth segment of the metasoma (Fig. 14 and 15A, less clear in the male type, obvious in the juvenile, syntypes in the NHMUK). Males of these three species also show slender pedipalp chelae. The type series of B. nigrovesiculosus includes only two animals, and more material is necessary to correctly evaluate the relationship between these three species. Nevertheless the males of B. nigrovesiculosus can be distinguished from males of the other two species by a higher pectinal tooth count (Fig. 10 and 15B, 36 versus <32 in the other two species), and from B. tassili by having a squared first metasomal segment.

Figure 14. 

Photo of the syntypes of B. nigrovesiculosus (NHMUK). Photo by Sérgio Henriques.

Figure 15. 

Detailed morphology of the larger syntype of B. nigrovesiculosus (NHMUK) A Lateral view of the telson and two terminal segments of the metasoma B Ventral view of the mesosoma, with pectines clearly visible C External lateral view of the terminal half of the right pedipalp che-la. All photos by Sérgio Henriques.

Buthus occidentalis Lourenço, Sun & Zhu, 2009

Buthus occidentalis: Lourenço et al. 2009: 72–74, fig. 7–19; Yang et al. 2013: 2.

Type material

1 F holotype (MNHN N° RS8844), Dakhlet Nouadhibou Region, in the coastal area (approx.20.28°, -16.24°), Mauritania. Paratypes: 1 M, 1 F juv. (MNHN N° 8845); 1 F, 1 M juv. (MHBU), all from the same locality.

Distribution

known only from the type locality.

Buthus occitanus (Amoreux, 1789), restricted distribution

= Scorpio rufus: Amoreux 1789a: 42–43 (synonymized by Amoreux 1789b, as the first revisor, ICZN article 24.2.2).

= Androctonus ajax: C. L. Koch 1839b: 53, pl. CXCIII, fig. 467 (synonymized by Simon 1879: 96). Type lost: Spain.

= Androctonus eurialus: C. L. Koch 1839b: 25–27, pl. CLXXXVII (not pl. CXXXVIl, as listed in the text), fig. 448 (synonymized by Simon 1879: 96). Type lost, France.

= Androctonus eurilochus: C. L. Koch 1839b: 27–28, pl. CLXXXVII, fig. 449 (synonymized by Kraepelin 1891: 196). Type lost, locality unknown.

= Buthus europaeus: Thorell 1876b: 7. Type is the lost Linnaeus (1748, 1754), specimen, purportedly from Italy (see the above “type species of Buthus“ section and Fet et al. 2002).

tridentatus = Buthus europaeus tridentatus: Franganillo 1918: 122–123 (syn. n.). Type unknown, Janvier, Navarre, Spain.

Scorpio occitanus: Amoreux 1789a: 42–43, pl. I, fig. 2; Amoreux 1789b: 10–16, pl. I. fig. 3; Herbst 1800: 73–82: Latreille 1804: 122; Latreille 1806: 132; Maccary 1810: 5–48; Audouin 1826: 172–173, pl. VIII, fig. 1; Audouin 1827: 410–411, pl. VIII, fig. 1; Dufour 1856 (part): 570.

Androctonus ajax: C. L. Koch 1850: 90.

Androctonus euryalus (ISS): C. L. Koch 1850: 90.

Androctonus eurylochus (ISS): C. L. Koch 1850: 90; Kraepelin 1891: 196.

Androctonus (Prionurus) occitanus: Lankester 1885: 380.

Buthus europaeus: Karsch 1879a: 18; Simon 1879: 96–98; Pavesi 1880: 312–313; Simon 1880c: 29; Pavesi 1884: 450; Pavesi 1885: 197, 199; Simon 1885: 51; Pocock 1889a: 116; Thorell 1893: 358–359; Birula 1896: 241–243; Birula 1900c: 9; Hirst 1925 (part): 415–416; Gadeau de Kerville 1926: 71; Bacelar 1928: 191; Hugues 1933: 487–488.

Buthus occitanicus (ISS): Dalla Torre 1905: 3; Táborský 1934: 40.

Buthus occitanus: Leach 1815: 391; Risso 1826: 156–157; Peters 1861b: 513; Karsch 1881a: 89; Kraepelin 1891 (part): 196–199 (part), pl. I, fig. 5, plI., fig. 18; Kraepelin 1895: 80; Kraepelin 1899: 26; Kraepelin 1901a (part): 266; Werner 1902 (part): 598; Birula 1910: 118–120; Masi 1912: 101; Borelli 1914b: 460; Lampe 1917: 191; Pavlovsky 1924: 77; Pavlovsky 1925: 140; Werner 1925: 209; Werner 1936: 173; Schenkel 1938: 4; Feytaud 1940: 38–39; Vachon 1940: 242–247, 254–258, fig. 1–9, 29, 33, 61–64; Käsmer 1941: 231; Denis 1948: 155–156; Vachon 1948c: 61, fig. 5; Vachon 1949a: 156–160, fig. 331–344, 348, 372–380; Vachon 1950b: fig. 591; Vachon 1951a: fig. 641, 657, 663, 679, 687, 696; Vachon 1951b: 621–623; Vachon 1952a: 264; Vachon 1952b: 274–279; Vachon 1961: 31–32; Bücherl 1964: 57; Pérez 1974: 22; Vachon 1974: 873; Goulliart 1979: 2; Levy and Amitai 1980 (part): 15–16; Kinzelbach 1982: 53; Prost 1982: 5; Mari et al. 1987; Sissom 1990: 92, fig. 3.17C, L; Kovařík 1992a (part): 183; Reichholf and Steinbach 1992: 33, fig. 4–5; Crucitti 1993: 51; Crucitti et al. 1994: 57–66; Vincent 1994: 6; Crucitti and Chinè 1995: 15–26; Braunwalder 1997b: 3; Crucitti and Chinè 1997: 195–200; Cloudsley-Thompson and Lourenço 1998: 1–2; Kovařík 1998: 106; Kovařík 1999 (part): 39, 42, fig. 3; Fet and Lowe 2000 (part): 92–94; Lourenço 2003 (part): 884, 886–887, fig. 27–32; Lourenço and Vachon 2004: 83–85, fig. 1–15; Kovařík and Whitman 2005 (part): 106; Teruel and Pérez-Bote 2005: 276; Castilla and Pons 2007: 258; Dupré et al. 2008: (pages unnumbered); Sousa et al. 2010: 207; Colombo 2011: 1; Rossi 2012: 274–278; Pedroso et al. 2013: 300; Rossi, Tropea and Yağmur 2013: 3; Martin-Eauclaire et al. 2014: 56; Teruel and Melic 2015: 6–9; Lourenço 2016b: fig. 2.

Buthus occitanus occitanus: Birula 1910: 118; Hadži 1929: 31; Vachon 1949a: 156–160, fig. 331–344; Vachon 1949c: 336; Vachon 1952a (part): 264; Le Corroller 1967: 63; Fet and Lowe 2000 (part): 94–95; Gantenbein and Largiadèr 2003 (part): 120, 122.

Buthus occitanus tridentatus: Fet and Lowe 2000: 97; Kovařík 2001: 79;

Buthus (Buthus) occitanus: Birula 1909b: 507; Birula 1910: 143; Birula 1914b: 644–664; Birula 1917a: 22, 38–39, 199, 213.

Buthus (Buthus) occitanus occitanus: Roewer 1943: 206.

Buthus cf. occitanus: Piñero et al. 2013: 88.

Scorpio australis (MIS): Asso 1784: 146, Tab. I, fig. 2.

Scorpio (Androctonus) occitanus: Gervais 1844a: 42–44; pl. XXIII, fig. 4.

Scorpio occitanicus (ISS): Serres 1822: 65.

Scorpion Occitanus (ISS): Latreille 1817: 105–106.

Type material

type unknown, Souvignargues, Occitanie Region, France.

Distribution

Traditionally, the distribution of B. occitanus was considered to span from the Moroccan Atlantic shores in North Africa to the Middle East in Asia and to Southern-Western Europe. However, following the description of new species in the genus, the present distribution of B. occitanus has been restricted to NE Spain and SW France. Several molecular phylogenetic studies have demonstrated that the species range does not extend beyond Western Europe (Gantenbein and Largiadèr 2003, Sousa et al. 2010, 2012, Pedroso et al. 2013). Lourenço and Vachon (2004) and Rossi (2012) include redescriptions of B. occitanus that include only European animals, but Vachon (1952a) also included material from Morocco, which was most likely not conspecific.

Remarks

All material collected outside of the range here proposed should be considered as Buthus sp. Only the re-examination of those specimens could reveal their appropriate identity. Vachon (1952a) included animals from the west (Atlantic) coast of Morocco, from Kenitra to El Jadida, within his definition of B. o. occitanus. As explained above this material is now considered not to be part of B. occitanus and thus remains unnamed. We opted to leave Androctonus eurilochus in synonymy with B. occitanus, despite the uncertain about the species provenance, to help to stabilise the genus’ taxonomy. To further bring stability to Buthus taxonomy we propose Franganillo’s subspecies, B. o. tridentatus, as a junior synonym of B. occitanus. Although the type specimen is not known (if it ever existed as such), its type locality is clearly stated as Javier, in Navarre, Spain. Extensive sampling on the left bank of the Ebro River (Sousa 2017) indicates that only B. occitanus occurs in this part of Spain and hence we here propose this new synonymy.

Buthus orientalis Lourenço & Simon, 2012

Buthus orientalis: Lourenço and Simon 2012: 10–14, fig. 1–12; Rossi 2013: 191–192; Rossi, Tropea and Yağmur 2013: 5, 7.

Type material

1 F holotype (MNHN N° RS8910), Alexandria (approx. 31.17°, 29.91°), Egypt. Paratypes: 7 M, 13 F (MNHN, N° RS6623), same locality.

Distribution

known only from the type locality.

Buthus paris (C. L. Koch, 1839)

= Androctonus clytoneus: C. L. Koch 1839a: 70–72, pl. CLXIII, fig. 384 (synonymized by Vachon 1949c: 380–381). Types lost; Africa.

Androctonus paris: C. L. Koch 1839a: 25–28, pl. CLI, fig. 352; C. L. Koch 1850: 90.

Androctonus clytonicus (ISS): Gervais 1844a: 43.

Androctonus clytoneus: C. L. Koch 1850: 90.

Buthus occitanus paris: Birula 1903: 107; Birula 1910: 118, 155; Giltay 1929: 196; Werner 1932: 300–305; Vachon 1949c: 380–388, fig. 356, 400, 445–455; Vachon 1951b: 621; Vachon 1952a: 308–316, fig. 356, 400 445–455; Malhomme 1954: 29; Arroyo 1961: 186–189; Le Corroller 1967: 63; Peréz 1974: 23; Levy and Amitai 1980: 16; El-Hennawy 1992: 98, 121; Kovařík 1995: 20; Gantenbein et al. 1998a: 51; Kovařík 1998: 106; Fet and Lowe 2000: 96; Touloun et al. 2001: 2; Gantenbein and Largiadèr 2003: 120, 122; Touloun 2012: 35, 104, 108, fig.5B.

Buthus (Buthus) occitanus paris: Birula 1910. 145, 155; Birula 1917a: 223.

Buthus paris: Lourenço 2003: 896–897, fig. 52–56; Kovařík 2006: 2, 6, 8, 15, fig. 10–11; Lourenço 2013: 65–66; Rossi, Tropea and Yağmur 2013: 3, 5, 7; Aboumaâd et al. 2014: 6; Touloun et al. 2014: 77–78; Lourenço and Sadine 2016: 14–15.

Type material

Holotype lost according to Fet and Lowe (2000), Algeria. Vachon (1949c, 1952a) wrote that the types came from Alger without further explanation.

Distribution

the species is currently distributed across Algeria, Morocco and Tunisia.

Remarks

Because of the description of new species from Algeria, and the lack of both type specimen and locality (beyond the country), a neotype for B. paris is necessary to stabilize the taxonomy of Algerian Buthus, which may challenge the status of some newly described species. This is further complicated by recent diagnoses of B. paris (at least in part: Lourenço 2003, Kovařík 2006, Lourenço and Sadine 2016) that differ from those offered by Vachon (1952a). The differences between the different diagnoses include the number of rows in the movable finger, the aspect ratio of the first metasomal segment, the body chaetotaxie, the aculeus to vesicle length and the type of sexual dimorphism of the pedipalp chelae. Vachon (1952a) studied a large number of specimens from the entire Maghreb region, unrivalled by any subsequent study, which leads us to consider Vachon’s description as the “gold standard”. Vachon himself stated that most of the specimens used in his redescriptions were stored at the MNHN. If this material is ever located, it should have priority in the designation of a neotype. Although Vachon (1952a) did not formally describe any varieties within B. paris, he split the specimens that compose the species into three regions: 1) The typical region (from Algiers to northern Tunisia); 2) Specimens from the Oujda region (Morocco); 3) Those from northern Morocco and the Middle Atlas flanks. Interestingly the split of B. paris into these three regions corresponds well with the distribution of the genetic variability in the cox1 gene (Gantenbein and Largiadèr 2003, Sousa et al. 2012, Pedroso et al. 2013), placing B. paris in two different groups: in occitanus which include all B. paris specimens from regions two and three, and tunetanus, which include B. paris specimens from the typical region one, along the split of the two cox1 groups in the middle of Algeria (Fig. 7). If confirmed, this will mean that B. paris does not occur in Morocco. It is unclear if the variety from the third region above might correspond to B. confluens Lourenço, Touloun & Boumezzough, 2012, although these authors (page 22) refrained from suggesting this possibility because they could not find any of the material used by Vachon, purportedly to be in the MNHN, to describe this variety.

Buthus parroti Vachon, 1949, stat. n.

Buthus atlantis parroti: Vachon 1949a: 168–169, fig. 346, 350, 352–354, 356, 363; Vachon 1952a: 254–255, fig. 346, 350, 352, 353, 354, 356, 363; Le Corroller 1967: 63; Pérez, 1974 1974: 22; El-Hennawy 1992: 98, 119; Kovařík 1995: 20; Kovařík 1998: 106; Lourenço 2003: 883–885, fig. 23–26; Fet and Lowe 2000: 91–92; Touloun 2012: 43, fig. 9B; Pedroso et al. 2013: 300.

Type material

1 F (MNHN N° RS1870), 1F, 12 juv., syntypes (MNHN), Forest house (approx. 30.31°, -9.33°), Ademine Forest, 40 Km S.W. of Agadir, Morocco; 2 M, 1 F juv., syntypes (MNHN), Taroudant (approx. 30.46°, -8.87°), Morocco.

Distribution

Known only from the Sous River Valley. Type localities in Fet and Lowe (2000) were mixed up.

Remarks

B. parroti was first described as a subspecies of B. atlantis. The two species occupy different habitats in Western Morocco; B. parroti is a forest species and B. atlantis is a sand dune dweller (Vachon 1952a). Furthermore, B. atlantis is clearly larger than B. parroti. Additionaly, the first metasomal segment of B. parroti is wider than long, the aculeus is shorter than the vesicle and the anal arch has only two lobes (Vachon 1952a), while in B. atlantis this segment is longer than wide, the aculeus is as long as or longer than the vesicle and the anal arch has three lobes (Vachon 1952a). Three additional Buthus species occur in the same area of Morocco as B. parroti, namely B. elmoutaouakili Lourenço & Qi, 2006, B. mardochei Simon, 1878 and B. mariefranceae. B. parroti can be distinguished from all three species by the presence of macrosetae in the terguites (Vachon 1952a). Moreover, it can be distinguished from B. mariefranceae by its larger size and absence of a dark fifth metasomal segment. Mesosoma colour pattern is not clear for B. parroti; the examined specimen at the MNHN (RS1870) (Fig. 16) appears to have two very faint darker stripes, while B. mariefranceae has two very well marked mesosomal dark stripes. B. parroti males show slender pedipalp chelae than females, while there is little if any sexual dimorphism in B. mardochei.

Buthus prudenti Lourenço & Leguin, 2012

Buthus prudenti: Lourenço and Leguin 2012: 2–6, 8, fig. 1–14; Lourenço 2016a: 76.

Type material

1 M holotype (MNHN N° RS8913), Ouro Labaré (9.38715°, 13.83447°), Bénoué, Cameroon. Paratypes: 7 M, 8 F, same locality; 11 paratypes (MNHN N° RS8914, RS8915), 4 paratypes (CBGP).

Distribution

known only from the type locality.

Remarks

type locality toponym derived from the coordinates given in the original description, as the type locality given was only Region of Sanguéré-Djoi, Cameroon.

Figure 16. 

Photo of a B. parroti female syntype (MNHN Nº RS1870), from the Ademine Forest, 04-1939, S.W. of Agadir, Morocco (Vachon 1952b).

Buthus pusillus Lourenço, 2013

Buthus pusillus: Lourenço 2013: 64–67, fig. 1–9; Lourenço and Sadine 2016: 14; Rossi, Tropea and Yağmur 2013: 5, 7.

Type material

1 M holotype (ZMH N° A11/13), Tizi Oumalou (36.5102°, 4.3390°), Tizi Ouzou Province, Djurdjura Mountains, Algeria. Paratype: 1 M juv. (ZMH N° A12/13), same location.

Distribution

known only from the type locality.

Remarks

The locality we present here corresponds to the coordinates given in the paper, although these do not agree with the elevation also reported in the paper: 2150 m a.s.l. This is very close to the maximum altitude of the highest peaks of the Djurdjura Mountains, and much higher than the 935 m a.s.l. of Tizi Oumalou.

Buthus rochati Lourenço, 2003

Buthus rochati: Lourenço 2003: 893–896, fig. 47–51;Aboumaâd et al. 2014: 6.

Buthus occitanus mardochei panousei: Vachon 1949c: 373–376, fig. 426–432; Vachon 1952a: 301–304, fig. 426–432; Le Corroller 1967: 63; Pérez 1974: 23; Touloun 2012: 57.

Type material

1 M holotype (MHNG), Tafnidilt Region (as Tafnidit) (approx. 28.56°, -11.03°), Guelmim Region, Morocco. Paratypes: 1 M, 1 F (MHNG); same locality; 6 F (MHNG), west of Tafnidilt Region, Draa River valley; 3 M, 1 F (MHNG), unknown locality.

Distribution

known distribution confined to the Tafnidilt region of Morocco.

Remarks

Lourenço (2003) does not mention the most remarkable diagnostic character given by Vachon (1952a), the interrupted dorso-median keel of the pedipalp patella. However Lourenço's Figure 49 illustrates this character, as it is a copy of Vachon's original drawings, and thus we consider it as part of the species diagnose and the most reliable diagnostic character for B. rochati. Vachon (1949) infrasubspecific name is not available as is explained above.

Buthus saharicus Saddine, Bissati & Lourenço, 2015

Buthus saharicus: Saddine et al. 2015: 47–49, fig. 6–8; Lourenço 2016b: fig. 4.

Type material

1 F holotype (MNHN), Ghardaïa Region (approx. 32.300°, 3.833°), in Wadi bed, Algeria. Paratypes: 1 M (UGA), 1 F juv. (MNHN), same locality.

Distribution

known only from the type locality.

Remarks

Saddine et al. (2015) claimed that B. saharicus was the “first true deserticolous species found in Algeria”, a bold claim given that B. tunetanus (sensu Vachon 1949, part) had already been recorded for Beni Abbés by Vachon (1949, 1952), also a desert location in central west Algeria, and albeit neither localities are Erg Desert areas, Beni Abbés is in the border of the Grand Erg Occidental.

Buthus tassili Lourenço, 2002

Buthus tassili: Lourenço 2002: 113–115, fig. 10, 12, 14; Lourenço 2003: 906–909, fig. 80–86; Rossi, Tropea and Yağmur 2013: 3–5, 7.

Buthus occitanus (MIS): Pallary 1929: 134, 140; Pallary 1934: 98–99.

Buthus occitanus tunetanus neeli: Gysin 1969: 65–71, fig. 1–5; Peréz 1974: 22; Touloun 2012: 37, 40.

Buthus occitanus tunetatus , “Spécimens des régions montagneuses centrales du Sahara”: Vachon 1952a: 279.

Type material

1 M holotype (MNHN N° RS8501), Tin Tazarif (approx. 24.466°, 10.466°), Illizi, Algeria. Paratype: 1 F (MNHN N° RS8622), same locality.

Distribution

This species is known from a wide area around the Hoggar and Tassili N’Ajjer Mountains, including at least one locality in Libya.

Remarks

The Tin Tazarif coordinates given here, standing at 880 m a.s.l., do not match the altitude given for the point by Lourenço (2002), of 1.800 m a.s.l., but correspond well to the map location given by the author in Figure 6 of the same article. Nevertheless there are several locations in the Tassili N’Ajjer Mountains at or above 1.800 m a.s.l., located closer to Jebel Azao, its highest peak. The species ranges from the Hoggar Mountains to the Tassili N’Ajjer Mountains, including Ghat in Libya. Gysin’s name, B. o. t. neeli, is not available under the ICZN article 10.2 and 45.5, as already stated by Fet and Lowe (2000). Therefore, it cannot enter formal synonymy. Nevertheless, because Gysin’s description (1969) brings relevant taxonomic information (figures and new localities) for B. tassili, we have decided to clearly state this new informal synonymy. The specimens studied of both species come from the Algerian Hoggar Mountains and share a typical darkened fifth segment of the metasoma (see B. nigrovesiculosus above).

Buthus trinacrius Lourenço & Rossi, 2013

Buthus trinacrius: Lourenço and Rossi 2013: 10–12, fig. 1–9.

Buthus europaeus (MIS): Simon 1879: 97; Simon 1910 (part): 69.

Buthus occitanus (MIS): Kraepelin (1901): 266.

Type material

1 M holotype, Palermo Province? (approx. 38.05°, 13.32°), Sicily. Paratypes: 1 M, 1 F. All type material in bad conditions (MNHN N° RS3247).

Distribution

Recorded from Sicily.

Remarks

Although Lourenço and Rossi (2013) report that the collector is not mentioned in Simon's notes, Kraepelin (1901), in his list of all the scorpion material present in the MNHN Paris, writes that Letourneur collected the Buthus material from Sicily and Corfu (Greece), which causes doubts regarding the correct collection locality of the specimens used to describe this species. Simon (1879) had doubts about the actual existence of Buthus in Sicily (when examining the specimens that would eventually be designated as type material for B. trinacrius): “il habite probablement aussi le midi de l’Italie et la Sicile”, and Buthus has never been found in mainland Italy. Furthermore, the authors also claimed that Simon (1910) “referred to the almost impossibility to distinguish Buthus populations from North of Africa with those from Spain and Sicily”, which is only partially correct. Simon (1910) solely refered to the distinction of Buthus populations of Algeria and Spain from those of Egypt, although in a subsequent paper, Simon does state that Buthus exist in Sicily without any further comments.

Buthus tunetatus (Herbst, 1800)

Scorpio tunetanus Herbst 1800: 68–69, pl. III, fig. 3 (not pl. II, fig. 2, as listed in the text); Latreille 1804: 122–124.

Androctonus (Leiurus) tunetanus: Ehrenberg in Hemprich and Ehrenberg 1829: 354.

Androctonus (Leiurus) tunetanus genuinus: Ehrenberg in Hemprich and Ehrenberg 1829: 354.

Androctonus (Liurus) tunetanus: Ehrenberg in Hemprich and Ehrenberg 1831 (pages unnumbered)

Androctonus (Liurus) tunetanus genuinus: Ehrenberg in Hemprich and Ehrenberg 1831 (pages unnumbered).

Androctonus tunetanus: C. L. Koch 1845: 15–19, pl. CCCCI (sic), fig. 968; C. L. Koch 1850: 90.

Buthus occitanus (MIS): Kovařík and Whitman 2005 (part): 106.

Buthus occitanus tunetanus: Birula 1903: 107; Birula 1910: 118; Borelli 1914a: 154–155; Borelli 1914b: 461; Borelli 1924: 4–5; Borelli 1928: 351; Giltay 1929: 196–197; Werner 1929: 30–31; Caporiacco 1932: 395–396; Schenkel 1932: 379–380; Werner 1932: 300–305; Pallary 1934: 99; Borelli 1934: 169; Werner 1934b: 84–85, fig. 4; Werner 1936: 173; Caporiacco 1937: 345; Schenkel 1949: 186; Vachon 1949c: 344–353, fig. 381–393; Vachon 1951a: fig. 670; Vachon 1952a: 272–281, fig. 381–393, 670; Vachon 1966: 211; Peréz 1974: 22; Levy and Amitai 1980: 16; El-Hennawy 1992: 98, 121; Kovařík 1995: 20; Kovařík 1997: 179; Gantenbein et al. 1998a: 51; Gantenbein et al. 1998b: 33–39; Kovařík 1998: 106; Fet and Lowe 2000: 97; Lourenço 2002, p. 113, 115, fig. 8–9, 11, 13; Kovařík 2002: 6; Gantenbein and Largiadèr 2003: 120, 122; Ben Othmen et al. 2004: 257; Touloun 2012: 37, 41.

Buthus (Buthus) occitanus tunetanus: Birula 1908: 123–124; Birula 1909: 507–508, fig. B; Birula 1910: 156–157; Birula 1917a: 223; Roewer 1943: 206.

Buthus tunetanus: Simon 1872: 251–252; Lourenço 2003: 897–899, fig. 57–61; Kovařík 2006: 2, 8, 10, 15, fig. 16–19.; Sadine et al. 2011: 6; Lourenço and Cloudsley-Thompson 2012: 13–16, fig. 8; Lourenço and Simon 2012: 12; Lourenço 2013: 65–66; Rossi, Tropea and Yağmur 2013: 4–5, 7

Scorpion Tunetanus (ISS): Latreille 1817: 106.

Type material

Types lost according to Fet and Lowe (2000), Tunisia.

Distribution

The species is currently distributed across Algeria, Libya, Morocco and Tunisia, and doubtfully in the island of Malta.

Remarks

Because of the description of new species from Tunisia, and the lack of both type specimen and locality (beyond the country), a neotype for B. tunetanus is necessary to stabilize the taxonomy of Tunisian Buthus. As explained for B. paris, this is further complicated by recent diagnoses of B. tunetanus that differ from those offered by Vachon (1952a), and as such if Vachon’s B. tuntetanus material is found in the MNHN it should be given priority in the future designation of a neotype. Vachon (1952a) did not formally described any variety of B. tunetanus, but he again split the specimens that compose the species into four regions: 1) the typical region, corresponding to north and central Tunisia; 2) the southern montane region of Algeria, specimens from which have subsequently been described as B. tassili; 3) the Algerian Saharan Atlas and the southern region of the High Plateau; and 4) the disjunct desert regions of southern Tunisia, western central Algeria and eastern central Morocco. It is unclear whether region 3 or 4 might either correspond to B. dunlopi or B. saharicus. As explained in Fet and Lowe (2000), the name A. (Leiurus) t. genuinus refers to the nominotypical form of the species and as such the adjective “genuinus” is not an available subspecific name.

Buthus yemenensis Lourenço, 2008

Buthus yemenensis: Lourenço 2008: 47–50, fig. 1–7.

Type material

1 F holotype (ZMH N° A33/08), Ma'bar (approx. 14.8°, 44.3°), Dhamar, Yemen.

Distribution

Known only from the type locality.

Buthus sp.

Buthus albengai (MIS): Habel et al. 2012: 3–4.

Buthus europaeus (MIS): Simon 1899: 85; Simon 1910 (part): 68–70, fig. 5, 8.

Buthus malhommei (MIS): Habel et al. 2012: 3–4.

Buthus occitanus (MIS): Karsch 1881b: 8 (Libya); Pocock 1899: 834 (Africa); Kraepelin 1901a (part): 266; Werner 1902 (part): 598; Chaignon 1904: 83–84 (Tunisia); Tullgren 1909: 2–3 (Egypt); Borelli 1924 (Libya): 4; King 1925: 81 (Sudan); Gough and Hirs 1927: 5, fig. 9 (Egypt); Pallary 1934: 98–99; Werner 1934a: 269, fig. 330 (Morocco); Pallary 1938: 281–282; Sergent 1938: 519–520, pl. 49; Monard 1939: 82–83 (Guinea-Bissau); Moriggi 1941: 84; Sergent 1941a: 355, fig. 1E, 2.7, pl. 35, fig. 7; Sergent 1941b: 447, plate 37; Vachon 1941: 52; Frade 1947: 268 (Guinea-Bissau); Vachon 1952a (part): 262–271, fig. 331–344, 348, 372–380, 591, 641, 657, 663, 679, 687, 696; Vachon 1953: 1021–1024, fig. 12 (Mauritania); Malhomme 1954: 28 (Morocco); Belfield 1956: 44; Kinzelbach 1975: 14, fig. 1; Lamoral and Reynders 1975: 505 (Africa); Levy and Amitai 1980 (part): 15–16; Kinzelbach 1984: 100 (Asia); Kinzelbach 1985: map II (Asia); El-Hennawy 1987: 17 (Egypt); Amr et al. 1988: 374 (Jordan); Michalis and Dolkeras 1989: 265–266 (Greece); El-Hennawy 1992: 98, 101, 119–120 (Arabia); Kovařík 1992a (part): 183; Kovařík 1992b: 90 (Iraq); Amr and EI-Oran 1994: 181 (Jordan); Kovařík 1997: 179 (Maghreb); Fet and Lowe 2000 (part): 92–94; Kovařík 2002: 5; Lourenço 2003: 884 (Morocco) Soleglad and Fet 2003: 7 (Morocco); Kaltsas et al. 2008: 215–216 (Libya); Sadine et al. 2012: 33; El-Hennawy 2013: 260; Aboumaâd et al. 2014: 5.

Buthus occitanus occitanus (MIS): Pocock 1895: 299 (Egypt); Vachon 1952a (part): 262–271, fig. 331–344, 372–379, 400, 554; Levy and Amitai 1980: 16 (Africa); Kovařík 1995: 20 (Morocco); Kovařík 1997: 179 (Ghana); Gantenbein et al. 1998a: 51, Fet and Lowe 2000 (part): 94–95; Gantenbein and Largiadèr 2003 (part): 120, 122.

Buthus occitanus occitanus, “Afrique occidentale française”: Vachon 1952a: 270.

Buthus occitanus occitanus, “Cote Occidentale du Maroc”: Vachon 1952a: 268–269, fig. 373–378.

Scorpio occitanus: Dufour 1856 (part): 570.

not Buthus: Vachon 1955: 372 (El Fâcher, Djebel Meidob, Darfur, Sudan) (Vachon said that this material was close to but not part of the genus Buthus).

Note

We futher propose to transfer a species from the genus Buthus to the genus Androctonus.

Androctonus barbouri (Werner, 1932), comb. n.

Buthus barbouri: Werner 1932: 300, fig. 141; Vachon 1949b: 287–288, fig. 371; Vachon 1952a: 261–262; Perez 1974: 22; El-Hennawy 1992: 98, 119; Kovařík 1995: 20; Kovařík 1998: 106; Fet and Lowe 2000: 92.

Type material

1 M (type probably lost), Agadir (approx. 30.43°, -9.60°), Morocco.

Distribution

known only from the type locality, Agadir, just north of the Sous River mouth, in southern Morocco.

Remarks

The species B. barbouri was described by Werner from Agadir, Morocco. However, this species cannot be linked to any of the Buthus species known from the reported type locality. This problem was already recognized by Vachon (1952a) that considered this species to have been “imperfectly described”, but then failed to make any taxonomic or nomenclatural act to fix it. Buthus barbouri and B. marrocanus are the only fully dark Buthus species known from Morocco. B. marrocanus was described from the Rabat region, 500 km distant from Agadir, and is distinguished from the former species by the shape of the inferior lateral keels of the fifth segment of the metasoma and by the number of pectinal teeth (Vachon, 1952a). Because of the chaetotaxy of the pedipalps Vachon (1952a) considered B. barbouri to be closer to Androctonus mauritanicus (Pocock, 1902). In agreement with these diagnostic characters, and Vachon’s (1952) opinion, we transfer B. barbouri to the genus Androctonus, with the new combination Androctonus barbouri (Werner, 1932). The relationship between A. barbouri and A. mauritanicus bourdoni Vachon, 1948, a subspecies of A. mauritanicus (Pocock, 1902), another black Androctonus species that also occurs in the Sous River valley, should be investigated.

Acknowledgments

Pedro Sousa is grateful to Arie van der Meijden for the opportunity he was given to sample for Buthus and other scorpions in Morocco, Tunisia, and Egypt, and also for the financial support to visit the MNHN, as part of an unrelated research project. Pedro Sousa would also like to thank Mark Judson, Elise-Anne Leguin and Wilson Lourenço who welcomed him at the MNHN. José Templado Gonzalez, Javier Sanchez Almazan and Carolina Noreña Janssen also welcomed Pedro Sousa during is stay at the MNCN.

We would like to thank our friend Sérgio Henriques who provided us with photographs of B. nigrovesiculosus with the kind permission of Janet Beccaloni from the NHMUK; and to extend our thank to Anja Friederichs and Jason Dunlop from the ZMB who provided us with photographs of B. intermedius and B. intumescens. We would also express our thanks to Oulaid Toulon who provided clear information about the location of the type material of B. draa and to Iñigo Sánchez who gave us the coodinates of the type locality of B. ibericus.

For their help in providing us with otherwise difficult-to-obtain bibliography, we thank Wilson Lourenço and Andrea Rossi, as well as Alison Corley, Elisa Mora, Gerard Dupré, Petr Dolejš and Petr Kment (Czech National Museum of Natural History, Prague). Isabel Sá from the Library of the Faculty of Sciences of Porto´s University helped us to obtain publications located in the Library of Coimbra University. Porto’s Public Library helped us to obtain some difficult-to-locate Portuguese articles. Several users of the webpage etimologias.dechile.net/ provided useful comments regarding Buthus etymology. We also express our gratitude to Miguel Angel Zarazaga for his expert opinion in the interpretation of several articles of the ICZN regarding nomenclature problems in Buthus, although all conclusions about such matters in the present manuscript are our responsibility alone. We extend our thanks to Jan Ove Rein for keeping an updated and freely available list of valid scorpion species names (www.ntnu.no/ub/scorpion-files/).

Wilson Lourenço as Editor, Lionel Monod, and an additional anonymous reviewer provided helpful comments on an earlier draft of the manuscript.

This work was partially supported by FEDER through the COMPETE program, Portuguese national funds through the Portuguese Fundação para a Ciência e Tecnologia (FCT), financed by the Programa Operacional Potencial Humano (POPH) – Quadro de Referência Estratégico Nacional (QREN) from the European Social Fund and Portuguese Ministério da Educação e Ciência: PS, PhD grant SFRH/BD/74934/2010; DJH, IF-contract IF/01627/2014.

References

  • Aboumaâd B, Iba N, Dersi N (2014) L'envenimation scorpionique au Maroc: scorpions du genre Androctonus, Buthus et Hottentota. Bulletin de la Société de pathologie exotique 107: 1–9. https://doi.org/10.1007/s13149-014-0332-7
  • Amoreux P-J (1789a) Description méthodique d'une espèce de scorpion roux commune à Souvignargues en Languedoc et détails historiques à ce sujet. Observations sur la Physique, sur l'Histoire naturelle et sur les Arts. 35: 9–16. http://biodiversitylibrary.org/page/6071194
  • Amoreux P-J (1789b) Notice des insectes de la France, réputés venimeux. Tirée des ecrits des Naturalistes, des Médecins, and de l'Observation. Rue et Hôtel Serpente, Paris, 302 pp. http://biodiversitylibrary.org/page/34452155
  • Amr ZS (2015) Scorpionism and Dangerous Species of Jordan. In: Gopalakrishnakone P, Schwartz EF, Vega RCR de la, Possani LD (Eds) Scorpion Venoms. Springer, Dordrecht, 181–200. https://doi.org/10.1007/978-94-007-6404-0
  • Armas LF de, González-Moliné AL (2009) Primer registro de Buthus ibericus Lourenço and Vachon, 2004 (Scorpiones: Buthidae) para la provincia de Huelva, España. Boletín de la Sociedad Entomológica Aragonesa 45: 553−555.
  • Arroyo H (1961) Etude sur la variété oranaise du scorpion Buthus occitanus (Amoreux, 1789). Archives de l’Institut Pasteur d’Algérie 39: 186–189.
  • Audouin V (1826) Explication sommaire des planches d'Arachnides de l'Égypte et de la Syrie, publiées par Jules-César Savigny, Membre de l'Institut; offrant un exposé des caractères natureles des genres, avec la distinction des espèces. In: Description de l'Egypte, ou recueil des observations et des recherches qui ont été faites en Égypte pendant l'expédition de l'armée française, publié par les ordes de Sa Majesté l'empereur Napoléon le Grand. Histoire Naturelle. L'Imprimerie Impériale, Paris, 99–186. http://digi.ub.uni-heidelberg.de/diglit/jomard1809bd5_1_1?sid=b2853cf0c40b3c782fdc22a191d9050b
  • Audouin V (1827) Explication sommaire des planches d'Arachnides de l'Égypte et de la Syrie, publiées par Jules-César Savigny, Membre de l'Institut; offrant un exposé des caractères naturels des genres avec la distinction des espèces. In: Description de l'Égypte ou recueil des observations et des recherches qui ont été faites en Égypte pendant l'expedition de l'armée française. Seconde Édition, Dédiée au Roi, Histoire naturelle, Zoologie, Animaux invertébrés (suite). pp 291–430. Imprimerie de C. L. F. Panckoucke, Paris, 291–430. http://gallica.bnf.fr/ark:/12148/bpt6k280218/f1.image.r=description de l’egypte.
  • Bacelar A (1928) Aracnídios portuguêses III Catálogo Sistemático dos Aracnídios de Portugal citados por diversos autores (1831–1926). Bulletin de la Société Portugaise des Sciences Naturelles 10: 169–203.
  • Banks N (1910) The Scorpions of California. Pomona College Journal of Entomology 2: 185–190.
  • Belfield W (1956) A preliminary check list of the West African scorpions and key for their identification. Journal of the West African Science Association 2: 41–47.
  • Ben Othmen A, Chatti N, Ben Ali-Haouas Z, Ouldbrahim I, Said K (2004) Allozymic differentiation of Tunisian populations of Androctonus species and Buthus occitanus (Scorpiones: Buthidae). Biological Journal of the Linnean Society 81: 255–265. https://doi.org/10.1111/j.1095-8312.2003.00285.x
  • Berejano I, Pérez-Bote JL (2002) Determinación de una presa atípica en la dieta de Buthus occitanius (Amoreaux, 1789) (Scorpiones: Buthidae). Revista Ibérica de Aracnología 5: 59–60.
  • Birula AA (1896) Miscelanea scorpiologica. I. Zur synonymie der russischen Skorpione. Annuaire du Musée Zoologique de l'Académie Impériale des Sciences de Saint-Pétersbourg 1: 229–245.
  • Birula AA (1900) Scorpiones mediterranei Musei Zoologici Mosquensis. Izvestiya Imperatorskogo Obshchestva Lyubitelei Prirody, Istorii, Antropologii i Etnografii 98: 8–20.
  • Birula AA (1903) Bemerkungen über einige neue oder wenig bekaunte Skorpionnenformen Nord-Afrikas. Bulletin de l'Académie Impériale des Sciences de St-Pétersbourg [Izviestiia Imperatorskoi akademii nauk] 19: 105–113.
  • Birula AA (1908) Ergebnisse der mit Subvention aus der Erbschaft Treitl unternommenen zoologischen Forschungsreise Dr. F. Werner's nach dem Ägyptischen Sudan und Nord-Uganda. XIV. Scorpiones und Solifugae. Sitzungsberichte der Kaiserlichen Akademie der Wissenschaften, Mathematisch-Naturwissenschaftliche Klasse, Wien 117: 121–152. http://www.zobodat.at/publikation_articles.php?id=148339
  • Birula AA (1909) Scorpione und Solifugen von Tripolis und Barka. Nach der Sammlung von Dr. Bruno Klaptocz im jahre 19-06. Zoologische Jahrbücher. Abteilung für Systematik, Geographie und Biologie der Tiere 28: 505–522.
  • Birula AA (1914) Ergebnisse einer von Prof. Franz Werner im Sommer 1910 mit Unterstützung aus dem Legate Wedl ausgeführten zoologischen Forschungsreise nach Algerien. VI. Skorpione und Solifugen. Sitzungsberichte der Kaiserlichen Akademie der Wissenschaften, Mathematisch-Naturwissenschaftliche Klasse, Wien 123: 633–668. http://www.zobodat.at/publikation_articles.php?id=148652.
  • Birula AA (1917a) [authored as Byalynitskii-Birulya, A. A.] Arachnoidea Arthrogastra Caucasica. Pars I. Scorpiones. Zapiski Kavkazskogo Muzeya, Seriia A [Mémoires du Musée du Caucase, Série A] 5: 1–253. [English translation: Byalynitskii-Birulya AA, 1964. Arthrogastric Arachnids of Caucasia. Part I. Scorpions. Israel Program for Scientific Translations, Jerusalem. 170 pp.]
  • Birula AA (1917b) [authored as Byalynitskii-Birulya AA] Faune de la Russie et des pays limitrophes fondee principalement sur les collections du Musée Zoologique de l'Académie des Sciences de Russie. Arachnides (Arachnoidea), Petrograd, 227 pp. [English translation: Byalynitskii-Birulya AA, 1965. Fauna of Russia and Adjacent Countries. Arachnoidea. Vol. I. Scorpions. Israel Program for Scientific Translations, Jerusalem, 154 pp.]
  • Bodenheimer FS (1937) Prodromus Faunae Palestinae. Essai sur les éléments zoogéographiques et historiques du sud-ouest du sous-règne paléarctique. Mémoires p. Imprimerie de l'Institut Français d'Archéologie Orientale, Le Caire, 286 pp.
  • Borelli A (1904) Di alcuni scorpioni della Colonia Eritrea. Bollettino dei Musei di Zoologia ed Anatomia Comparata della R. Università di Torino 19: 1–5.
  • Borelli A (1914a) Contributo allo studio della fauna Libica. Materiali raccolti nelle zone di Misurata e Homs (1912-13). Scorpioni. Annali del Museo Civico di Storia Naturale di Genova 6: 148–159.
  • Borelli A (1914b) Gli scorpioni del Museo Civico di Storia Naturale di Milano. Atti della Società Italiana di Scienze Naturali e del Museo Civico di Storia Naturale in Milano 53: 456–464.
  • Borelli A (1924) Missione zoologica del Dr. E. Festa in Cirenaica. XVI. Scorpioni e Solifughi. Bollettino dei Musei di Zoologia ed Anatomia Comparata della R. Università di Torino 39: 1–16.
  • Borelli A (1928) Risultati zoologici della Missione inviata dalla R. Societa Geografica Italiana per l’explorazione dell’oasi di Giarabud (1926-1927). Scorpioni e Solifughi. Annali del Museo civico di Storia Naturale Giacomo Doria 52: 346–355.
  • Borelli A (1934) Scorpiones. In: Zavattari E (Ed.) Prodromo della fauna della Libia. Tipografia già Cooperativa, Pavia, 169–173.
  • Braunwalder ME (1997) A note on Pierre-Joseph Amoreux (1741-1824) and his publications on Buthus occitanus. Newsletter of the British Arachnological Society 78: 3–4.
  • Braunwalder ME, Fet V (1998) On publications about scorpions (Arachnida, Scorpiones) by Hemprich and Ehrenberg (1828–1831). Bulletin of the British Arachnological Society 11: 29–35.
  • Bücherl W (1964) Distribuição geográfica dos Aracnóides peçonhentos temíveis. Memórias do Instituto Butantan 31: 55–66.
  • Caporiacco L di (1932) Spedizione scientifica all’Oasi di Cufra (Marzo-Luglio 1931). Scorpioni e Solifugi. Annali del Museo civico di Storia Naturale Giacomo Doria 55: 395–408.
  • Caporiacco L di (1937) Risultati scientifici della Missione del Prof. G. Scortecci nel Fezzan e sui Tassili (1936). Scorpioni e Solifughi. Atti della Società Italiana di Scienze Naturali e del Museo Civico di Storia Naturale in Milano 76: 340–354.
  • Chaignon H de (1904) Contributions à l'Histoire Naturelle de la Tunisie. Bulletin de la Société d'Histoire Naturelle d'Autun 17: 1–166.
  • Cloudsley-Thompson JL (1956) Studies in diurnal rhythms VI. Bioclimatic observations in Tunisia and their significance in relation to the physiology of the fauna, especially woodlice, centipedes, scorpions, and beetles. The Annals and Magazine of Natural History 12: 305–329. https://doi.org/10.1080/00222935608655817
  • Cloudsley-Thompson JL, Lourenço WR (1998) On the meaning of the trivial name of the Scorpion Buthus occitanus (Amoreux, 1789) with notes on its distribution and variability. Newsletter of the British Arachnological Society 81: 1–2.
  • Comstock JH (1940) Order Scorpionida. In: Comstock JH (Ed.) The spider book, revised and edited by W. J. Gertsch. Cornell Univ. Press, Ithaca, 21–32.
  • Cox FA (1921) A list of California Arachnida. III. The Scorpionida. Journal of Entomology and Zoology, Pomona College 14: 12–13.
  • Crucitti P (1993) Problemi di conservazione degli scorpioni europei. Ambiente Risorte Salute 22: 51–55.
  • Crucitti P, Chine A (1995) Variability and anomalies in the scorpion Buthus occitanus occitanus (Scorpiones, Buthidae). Bollettino dell Associazione Romana di Entomologia 49: 15–26.
  • Crucitti P, Chine A (1997) Rapporto sessi in Buthus occitanus occitanus (Scorpiones, Buthidae). Bollettino della Società Entomologica Italiana 129: 129.
  • Dalla Torre KW von (1905) Die skorpione Tirols. Entomologisches Jahrbuch 1905: 1–4.
  • De Geer C (1778) Cinquieme Mémoire. Des Scorpions and des Faux-Scorpions. In: Mémoires pour servir à l'histoire des Insectes. Tome 7e. Imprimerie Pierre Hesselberg, Stockholm, 325–367. https://doi.org/10.5962/bhl.title.14802
  • Denis J (1948) Le scorpion languedocien dans les Pyrénées-orientales. Bulletin de la Société Linnéenne de Lyon 17: 155–156.
  • Dufour ML (1856) Histoire anatomique et physiologique des Scorpions. Mémoires présentés par divers savants à l'Académie des sciences de l'Institut de France 14: 561–656. doi: ark:/12148/bpt6k3319c
  • Dupré G (2016) Dictionary of scientific scorpion names. Arachnides 78: 1–68.
  • Dupré G, Lambert N, L'Association “Les Ecologistes D'Euzière” (2008) Le Scorpion Languedocien Buthus occitanus (Scorpiones, Buthidae). Sa répartition en France. Arachnides Supplément [not numbered].
  • El Hidan, Abdelmonaim M, Touloun O, Boumezzough A (2016) New data on the diversity of scorpion fauna in the oases of south eastern Morocco. Serket 15: 1–7.
  • El-Hennawy HK (1987) A simplified key to Egyptian scorpion species (Arachnida: Scorpionida). Serket 1: 15–17.
  • El-Hennawy HK (1992) A catalogue of the scorpions described from the Arab countries (1758-1990) (Arachnida: Scorpionida). Serket 2: 95–153.
  • El-Hennawy HK (2013) Preliminary list of Lebanese spiders and other arachnids (except ticks and mites). Serket 13: 228–275.
  • Ewing HE (1928) The scorpions of the western part of the United States, with notes on those occuring in Northern Mexico. Proceedings of the U.S. Natural Museum 73: 1–24.
  • Fabricius JC (1781) Species Insectorum, exhibentes eorum differentias specificas, synonyma auctorum, loca natalia, metamorphosin adjectis observationibus, descriptionibus. Tom. 1. Impensis Carol. Ernest. Bohnii, Hamburgi and Kilonii, 552 pp.
  • Fernández J (2004) Noticia de nuevos táxones para la ciencia en el ámbito Íbero-Balear y Macaronésico. Nuevos táxones animales descritos en la península Ibérica y Macaronesia desde 1994 (VIII). Graellsia 60: 217–243.
  • Fet V (2010) Scorpions of Europe. Acta Zoologica Bulgarica 62: 3–12.
  • Fet V, Braunwalder ME, Cameron HD (2002) Scorpions (Arachnida, Scorpiones) described by Linnaeus. Bulletin of the British Arachnological Society 12: 176–182.
  • Fet V, Lowe G (2000) Genus Buthus Leach, 1815. In: Fet V, Sissom WD, Lowe G, Braunwalder ME (Eds) Catalog of the Scorpions of the World (1758-1998). The New York Entomological Society, New York, 91–97.
  • Fet V, Sissom WD (2000) Family Euscorpiidae Laurie, 1893. In: Fet V, Sissom WD, Lowe G, Braunwalder ME (Eds) Catalog of the Scorpions of the World (1758–1998). The New York Entomological Society, New York, 355–380.
  • Feytaud J (1940) Les scorpions de France. Revue de Zoologie Agricole et Appliquée 39: 33–43.
  • Foley H (1951) Les scorpions noirs de l'Afrique du Nord. Bulletin de la Société d'Histoire Naturelle de l'Afrique du Nord 42: 33.
  • Frade F (1948) Trabalhos da Missão Zoológica da Guiné. X. Escorpiões, Solífugos e Pedipalpos da Guiné Portuguesa. Anais da Junta das Missões Geográficas e de Investigações Coloniais, Lisboa 3: 5–18.
  • Francke OF (1985) Conspectus genericus scorpionorum 1758–1982 (Arachnida: Scorpiones). Occasional Papers of the Museum, Texas Tech University 98: 1–32.
  • Franganillo P (1918) Arácnidos nuevos o hallados por primera vez en España. Boletín de la Sociedad Entomológica de España 1: 120–123.
  • Gadeau de Kerville H (1926) Scorpions. In: Voyage zoologique d’Henri Gadeau de Kerville en Syrie (avril-juin 1908): tome premier, récit du voyage et liste méthodique des animaux récoltés en Syrie, par Henri Gadeau de Kerville; mémoires sur l’Éponge, les Copépodes, les Ostracodes, les Isopodes, les Opiliones, les Myriopodes, les Thysanoures, les Odonates, les Orthoptères, les Hyménoptères mellifères, les Diptères et les Oligochètes récoltés pendant ce voyage, par Émile Topsent, C. Dwight Marsh, Eugène Daday de Deés, Mme Harriet Richardson Searle, William Sôrensen, le Comte Carl Attems, le Docteur Filippo Silvestri, R. Martin, Ignacio Bolivar, J. Pérez, le Docteur Joseph Villeneuve, Jacques Surcouf et le Docteur Wilhelm Michaelsen. J.-B. Baillière et fills, Paris, 71–72.
  • Gantenbein B, Braunwalder ME, Scholl A (1998) Allozyme studies on scorpions from the Aegean Region and from Morocco. Abstracts XIV International Congress Arachnology \ 22 Annual Meeting of the American Arachnological Society, Chicago, 51.
  • Gantenbein B, Büchi L, Braunwalder ME, Scholl A (1998) The genetic population structure of Euscorpius germanus (C.L. Koch) (Scorpiones: Chactidae) in Switzerland. In: Selden PA (Ed.) Proceedings of the 17th European Colloquium of Arachnology, Edinburg, 1997. British Arachnological Society, Burnham Beeches, Bucks, 33–40.
  • Gantenbein B, Largiadèr CR (2003) The phylogeographic importance of the Strait of Gibraltar as a gene flow barrier in terrestrial arthropods: a case study with the scorpion Buthus occitanus as model organism. Molecular Phylogenetics and Evolution 28: 119–130. https://doi.org/10.1016/S1055-7903(03)00031-9
  • Gertsch WJ, Soleglad ME (1966) The scorpions of the Vejovis boreus group (subgenus Paruroctonus) in North America (Scorpionida, Vaejovidae). American Museum Novitates 2278: 1–54.
  • Gervais P (1844a) Ordre III. Scorpionides. II. Scorpions. In: Walckenaer CA, Gervais P (Eds) Histoire naturelle des Insectes. Aptères. Librairie Encyclopédique de Roret, Paris, 14–74.
  • Gervais P (1844b) Remarques sur la famille des scorpions et description de plusieurs espèces nouvelles de la collection du Muséum. Archives du Muséum d'Histoire Naturelle, Paris 4: 201–240.
  • Giltay L (1929) Mission saharienne Augieras-Draper, 1927–1928. Scorpions. Bulletin du Muséum National d'Histoire Naturelle, Paris, 2eme série 1: 193–197.
  • Gough HL, Hirst S (1927) Key to the identification of Egyptian scorpions. A method of identifying Egyptian scorpions from the fifth caudal segment and stings. Bulletin of the Ministry of Agriculture of Egypt, Technical Scientific Service 76: 1–7.
  • Goulliart M, Frechin E (1979) Les scorpions de France. Bulletin de la Société Entomologique du Nord de la France 213: 1–3.
  • Gysin J (1969) Une nouvelle variété de scorpion au Hoggar Buthus occitanus (Am.) ssp tunetanus (Herbst) var neeli var. nov. Archives de l'Institut Pasteur d'Algérie 47: 65–72.
  • Habel JC, Husemann M, Schmitt T, Zachos FE, Honnen A-C, Petersen B, Parmakelis A, Stathi I (2012) Microallopatry caused strong diversification in Buthus scorpions (Scorpiones: Buthidae) in the Atlas Mountains (NW Africa). PloS One 7: e29403. https://doi.org/10.1371/journal.pone.0029403
  • Hadži J (1929) Skorpije Schmidtove zbirke (Euscorpius italicus polytrichus n. ssp. i ostale nove rase). (in Slovenian). Glasnik Muzejskega Drustva za Slovenijo 10: 30–41.
  • Hemprich FW, Ehrenberg CG (1828) Arachnoidea. Plates I + II. In: Symbolae Physicae seu Icones et Descriptiones Animalium Evertebratorum sepositis Insectis quae ex ilinere per Africam borealem et Asiam occidentalem. Friderici Guilelmi Hemprich et Chrisliuni Godofredi Ehrenberg, studio novae aut illustratae redierunt. Pe. Berolini ex officina Academica, Venditur a Mittlero.). https://doi.org/10.5962/bhl.title.107403
  • Hemprich FW, Ehrenberg CG (1829) Vorläufige Uebersicht der in Nord-Afrika und West-Asien einheimischen Scorpione und deren geographischen Verbreitung, nach den eigenen Beobachtungen von Dr. Hemprich und Dr. Ehrenberg. Verhandungen der Gessellschaft naturforschende Freunde in Berlin 1: 348–362. https://books.google.pt/books?id=8OoTAAAAQAAJ
  • Hemprich FW, Ehrenberg CG (1831) Animalia articulata. Arachnoidea. Scorpiones africani et asiatici. In: Hemprich et Ehrenberg Symbolae Physicae. Animalia evertebrata exclusis insectis. Percensuit Dr. C. G. Ehrenberg. Series prima cum tabularum decade prima. Continent Animalia Africana et Asiatica 162. Berolini ex officina Academica, 12 (unnumbered). https://doi.org/10.5962/bhl.title.107403
  • Hendrixson BE (2006) Buthid scorpions of Saudi Arabia, with notes on other families (Scorpiones: Buthidae, Liochelidae, Scorpionidae). Fauna of Arabia 21: 33–120.
  • Herbst JFW (1800) Natursystem der ungeflügelten Insekten. Naturgeschichte der Skorpionen. Gottlieb August Lange, Berlin, 86 pp.
  • Hjelle JT (1972) Scorpions of the Northern California Coast Ranges (Arachnida: Scorpionida). Occasional Papers of the California Academy of Sciences 92: 1–59.
  • Hofmann JJ, Hackius J, Boutesteyn C, Vander AP, Luchtmans J (1698) Lexicon Universale, Historiam Sacram Et Profanam Omnis aevi, omniumque Gentium; Chronologiam Ad Haec Usque Tempora; Geographiam Et Veteris Et Novi Orbis; Principum Per Omnes Terras Familiarum [...] Genealogiam; Tum Mythologiam, Ritus, Caerimonias, Omnemqu. Editio abs. Hofmann JJ (Ed. ) Johannes du Vivié, Leyden, 1072 pp. http://www.uni-mannheim.de/mateo/camenaref/hofmann.html.
  • Hull-Williams V (1987) Some notes on the scorpion Buthus occitanus collected in Morocco, April 1987, by V. Hull-Williams and A. Smith. Journal of the British Tarantula Society 2: 12–15.
  • Husemann M, Schmitt T, Stathi I, Habel JC (2012) Evolution and radiation in the scorpion Buthus elmoutaouakili Lourenco and Qi 2006 (Scorpiones: Buthidae) at the foothills of the Atlas Mountains (North Africa). Journal of Heredity 103: 1–9. https://doi.org/10.1093/jhered/esr130
  • International Commission on Zoological Nomenclature (ICZN) (1957) Direction 60. Designation under the Plenary Powers of a type species for the genus Scorpio Linnaeus, 1758 (Class Arachnida) in harmony with accustomed usage and suppression under the same Powers of the specific name europaeus Linnaeus, 1758, as published in the combination Scorpio europaeus (amendment of a Ruling given in Opinion 104). Opinions and declarations rendered by the International Commission on Zoological Nomenclature 1E: 1–12.
  • Karsch F (1881a) Uebersicht der Europäienschen Skorpione. Berliner Entomologische Zeitschrift 25: 89–91.
  • Karsch F (1881b) Verzeichniss der während der Rohfs'schen africanischen Expedition erbeuteten Myriopoden und Arachniden. Archive für Naturgeschichte 47: 1–14.
  • Karsch F (1886) Skorpionologische Beiträge. I. Ueber einen sizilianischen Skorpion. II. Uebersicht der Gruppe Buthinae (Androctonina). III. Ueber einen neuen Opisthacanthus (Peters). Berliner entomologische Zeitschrift 30: 75–79.
  • Karsch F (1891) Arachniden von Ceylon und von Minikoy gesammelt von den Herren Doctoren P. und F. Sarasin. III. Ordo Scorpiones. Berliner entomologische Zeitschrift 36: 305–307.
  • Kästner A (1941) Ordung der Arachnida: Scorpiones. In: Krumbach T (Ed.) Handbuch der Zoologie. Walter de Gruyter Verlag 3 (1), 117–240.
  • Kinzelbach R (1975) Die Skorpione der Ägäis. Beiträge zur systematik, phylogenie und biogeographie. Zoologische Jahrbücher, Abteilung für Systematik 3 102: 12–50.
  • Kinzelbach R (1982) Die Skorpionssammlung des Naturhistorischen Museums der Stadt Mainz. Teil I: Europa Und Anatolien. Mainzer Naturwisserschaftlichen Archiv 20: 49–66.
  • Kinzelbach R (1984) Die Skorpionssammlung des Naturhistorischen Museums der Stadt Mainz. Teil II: Vorderasien. Mainzer Naturwisserschaftlichen Archiv 22: 97–106.
  • Kinzelbach R (1985) Vorderer Orient. Skorpione (Arachnida: Scorpiones). Tübingen Atlas der Vorderen Orients (TAVO). Karte A VI 14.2. Ludwig Reichert Verlag, Wiesbaden.
  • Kovařík F (1992a) A check list of scorpions (Arachnida: Scorpiones) in the collections of the zoological department, National Museum in Prague. Acta Societas Zoologicae Bohemicae 56: 181–186.
  • Kovařík F (1992b) Buthus occitanus (Amoreuxi, 1789) and Orthochirus innesi Simon, 1910 (Scorpionidea, Buthidae) from Iraq. Casopis Národního Muzea, Rada Prirodovedná 159: 90.
  • Kovařík F (1995) First report of Orthochirus innesi (Scorpionida: Buthidae) from Morocco. Klapalekiana 31: 19–21.
  • Kovařík F (1997) A check-list of scorpions (Arachnida) in the collection of the Hungarian Natural History Museum, Budapest. Annales Historico-Naturales Musei Nationalis Hungarici 89: 177–185.
  • Kovařík F (1998) Štiří. Madagaskar, Jihlava, 176 pp.
  • Kovařík F (1999) Review of European scorpions, with a key to species. Serket 6: 38–44.
  • Kovařík F (2001) Catalog of the Scorpions of the World (1758-1998) by V. Fet, W. D. Sissom, G. Lowe, and M. Braunwalder (New York Entomological Society, 2000: 690 pp.). Discussion and supplement for 1999 and part of 2000. Serket 7: 78–93.
  • Kovařík F (2002) A checklist of scorpions (Arachnida) in the collection of the Forschungsinstitut und Naturmuseum Senckenberg, Frankfurt am Main, Germany. Serket 8: 1–23.
  • Kovařík F (2003) Scorpions of Djibouti, Eritrea, Ethiopia, and Somalia (Arachnida: Scorpiones), with a key and descriptions of three new species. Acta Societatis Zoologicae Bohemicae 67: 133–159. http://kovarex.com/scorpio/pdf/Kovarik2003c.pdf.
  • Kovařík F, Whitman S (2004) Cataloghi del Museo di Storia Naturale dell'Università di Firenze - sezione di zoologia «La Specola» XXII. Arachnida Scorpiones. Tipi. Addenda (1998-2004) e checklist della collezione (Euscorpiinae esclusi). Atti della Società Toscana di Scienze Naturali, Memorie, Serie B 111: 103–119.
  • Kraepelin K (1891) Revision der Skorpione. I. Die familie der Androctonidae. Jahrbuch der Hamburgischen wissenschaftlichen Anstalten 8: 143–286.
  • Kraepelin K (1895) Nachtrag zu Theil I der Revision der Scorpione. Jahrbuch der Hamburgischen wissenschaftlichen Anstalten 12: 73–96.
  • Kraepelin K (1901) Catalogue des Scorpions des collections du Muséum d'histoire naturelle de Paris. Bulletin du Muséum d'Histoire Naturelle, Paris 7: 265–274.
  • Kraepelin K (1903) Scorpione und Solifugen Nordost-Afrikas, gesammelt 1900 und 1901 von Carlo Freiherrn von Erlanger und Oscar Neumann. Zoologische Jahrbücher. Abteilung für Systematik, Geographie und Biologie der Tiere 18: 557–578.
  • Lamoral BH, Reynders SC (1975) A catalogue of the scorpions described from the Ethiopian faunal region up to December 1973. Annals of the Natal Museum 22: 489–576.
  • Lampe E (1917) Katalog der Skorpione, Pedipalpen und Solifugen des Naturhistorischen Museums der Residenzstadt Wiesbaden. Jahrbücher des Nassauischen Verein für Naturkunde, Wiesbaden 70: 185–208.
  • Lankester ER (1885) Notes on certain points in the anatomy and generic characters of scorpions. Transactions of the Zoological Society of London 11: 372–384.
  • Latreille PA (1804) Histoire des Scorpions. In: Histoire naturelle, générale et particulière des Crustacés et des Insectes. Ouvrage faisant suite aux oeuvres de Leclerc de Buffon, et partie de cours complet d'histoire naturelle rédigée par C. S. Sonnin, member de plusieurs Sociétés savantes. Imprimerie de F. Dufart, Paris, 110–129.
  • Latreille PA (1806) Familia Tertia. Scorpionides. In: Genera Crustaceorum et Insectorum secundum ordinem naturalem in familias disposita, iconibus exemplisque plurimis explicata. Amand Koenig, Parisiis and Argentorati, 130–132.
  • Latreille PA (1817) Tome III. Contenant les Crustacés, les Arachnides et les Insectes. In: Cuvier G (Ed.) Le règne animal distribué d'après son organisation, pour servir de base a l'histoire naturelle des animaux et d'introduction a l'anatomie comparée. Imprimerie de A. Belin, Paris, 653.
  • Laurie M (1896) Further notes on the anatomy and development of scorpions, and their bearing on the classification of the Order. Annals and Magazine of Natural History 18: 121–133.
  • Le Coroller Y (1967) Les scorpions du Maroc. Archives de l'Institut Pasteur d'Algérie 45: 62–71.
  • Leach WE (1815) A tabular View of the external Characters of Four Classes of Animals, which Linné arranged under INSECTA; with the Distribution of the Genera composing Three of these Classes into Orders, andc. and Descriptions of several New Genera and Species. Transactions of the Linnean Society of London 11: 306–400. https://doi.org/10.1111/j.1096-3642.1813.tb00065.x
  • Levy G, Amitai P (1980) Fauna Palaestina. Arachnida I: Scorpiones. The Israel Academy of Sciences an d Humanit es, Jerusalem, 172 pp.
  • Linnaeus C (1748) Systema naturae, sistens regna tria naturae, in classes et ordines, genera et species. Editio Sexta, Emendata et Aucta. Impensis Godofr. Kiesewetteri, Stockholmiae, 224 pp. https://doi.org/10.5962/bhl.title.36934
  • Linnaeus C (1754) Museum S:ae R:ae M:tis Adolphi Friderici Regis [etc] in quo Animalia rariora imprimis, et exotica quadrupedia, aves, amphibia, pisces, insecta, vermes describuntur et determinantur, Latine et Svetice cum iconibus. E Typographia Regia, Holmiae [Stockholm], 96 pp. doi: ark:/12148/bpt6k972707
  • Linnaeus C (1758) Scorpio. In: Systema naturæ per regna tria naturæ, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Tomus I. Editio decima, reformata. Laurentii Salvii, Holmiae [Stockholm], 624–625. https://doi.org/10.5962/bhl.title.542
  • Lönnberg E (1897) Om Skorpionernas och Pedipalpernas geografiska utbredning. Entomologisk Tidskrift 18: 193–211.
  • Lönnberg E (1898) A revision of the Linnaean type specimens of scorpions and pedipalps in the Zoological Museum of the Royal University of Upsala. The Annals and Magazine of Natural History (5th series) (7) 1: 82–89.
  • Lourenço WR (2002) Considérations sur les modèles de distribution et différentiation du genre Buthus Leach, 1815, avec la description d'une nouvelle espèce des montagnes du Tassili des Ajjer, Algérie (Scorpiones, Buthidae). Biogeographica 78: 109–127.
  • Lourenço WR (2003) Compléments à la faune de scorpions (Arachnida) de l'Afrique du Nord, avec des considérations sur le genre Buthus Leach, 1815. Revue Suisse de Zoologie 110: 875–912.
  • Lourenço WR (2005a) Description of a new scorpion species of the genus Buthus Leach, 1815 (Scorpiones, Buthidae) from Guinea and Senegal in Western Africa. Entomologische Mitteilungen aus dem Zoologischen Museum Hamburg 14: 229–236.
  • Lourenço WR (2005b) A new species of the genus Buthus Leach, 1815 (Scorpiones, Buthidae) from Senegal and Niger in Western Africa. Entomologische Mitteilungen aus dem Zoologischen Museum Hamburg 14: 245–251.
  • Lourenço WR (2005c) Description Of Three New Species Of Scorpion From Sudan (Scorpiones, Buthidae). Boletín de la Sociedad Entomológica Aragonesa 36: 21–28.
  • Lourenço WR (2005d) Nouvelles considérations taxonomiques sur les espèces du genre Androctonus Ehrenberg, 1828 et description de deux nouvelles espèces (Scorpiones, Buthidae). Revue suisse de Zoologie 12: 145–171. https://doi.org/10.5962/bhl.part.80291
  • Lourenço WR (2008) About the presence of the genus Buthus Leach, 1815 in the Arabian Peninsula and description of a new species (Scorpiones, Buthidae). Entomologische Mitteilungen aus dem Zoologischen Museum Hamburg 15: 45–52.
  • Lourenço WR (2013) A new species of Buthus Leach, 1815 from Algeria (Scorpiones, Buthidae). Entomologische Mitteilungen aus dem Zoologischen Museum Hamburg 16: 63–68.
  • Lourenço WR (2015) Deux nouvelles espèces de scorpions de la famille des Buthidae C. L. Koch, 1837 collectées dans le Parc National de Zakouma au Tchad. Revista Ibérica de Aracnología 26: 19–24.
  • Lourenço WR (2016a) A new species of the genus Buthus Leach, 1815 (Scorpiones: Buthidae) from dry forest formations in Central. Serket 15: 71–79.
  • Lourenço WR (2016b) Scorpion incidents, misidentification cases and possible implications for the final interpretation of results. Journal of Venomous Animals and Toxins including Tropical Diseases 22: 1–25. https://doi.org/10.1186/s40409-016-0075-6
  • Lourenço WR, Cloudsley-Thompson JL (2012) A new species of Buthus Leach, 1815 from Egypt (Scorpiones, Buthidae). Entomologische Mitteilungen aus dem Zoologischen Museum Hamburg 16: 11–18.
  • Lourenço WR, Francke OF (1984) The identities of Tityus floridanus and Tityus tenuimanus (Scorpiones, Buthidae). The Florida Entomologist 67: 424–429.
  • Lourenço WR, Qi J-X (2006) A new species of Buthus Leach, 1815 from Morocco (Scorpiones, Buthidae). Entomologische Mitteilungen aus dem Zoologischen Museum Hamburg 14: 287–292.
  • Lourenço WR, Rossi A (2013) Confirmation of a new species of Buthus Leach, 1815 from Sicily (Scorpiones, Buthidae). Biogeographical implications. Revista Ibérica de Aracnología 22: 9–14.
  • Lourenço WR, Sadine SE (2016) One more new species of Buthus Leach; 1815 from Algeria (Scorpiones: Buthidae). Revista Ibérica de Aracnología 28: 13–17.
  • Lourenço WR, Simon E (2012) Confirmation of a new species of Buthus Leach, 1815 from Alexandria, Egypt (Scorpiones, Buthidae). Serket 13: 8–15.
  • Lourenço WR, Slimani T (2004) Description of a new scorpion species of the genus Buthus Leach, 1815 (Scorpiones, Buthidae) from Morocco. Entomologische Mitteilungen aus dem Zoologischen Museum Hamburg 14: 165–170.
  • Lourenço WR, Touloun O, Boumezzough A (2012) Un nouveau Buthus Leach, 1815 (Scorpiones, Buthidae) du Nord du Maroc; possible lien entre les populations Marocaines et Européennes. Revista Ibérica de Aracnología 21: 21–25.
  • Lourenço WR, Vachon M (2004) Considérations sur le genre Buthus Leach, 1815 en Espagne, et description de deux nouvelles espèces (Scorpiones, Buthidae). Revista Ibérica de Aracnología 9: 81–94.
  • Maccary A (1810) Mémoire sur le scorpion qui se trouve sur la montagne de Cette, dé partement de l'Hérault, son venin, et l'usage qu'on pourrait en faire en Médecine. Imprimerie de Froullé, Paris, 48 pp.
  • Malhomme J (1954) Aperçu sur la distribution des scorpions de la région de Marrakech. Bulletin de la Société des Sciences Naturelles et Physiques du Maroc 34: 25–30.
  • Mari EG, Ferragut E, Navarro V, Laborda R, Comelles JC (1987) Nueva aportación al concimiento de los artrópodos de las Islas Columbretes. In: Alonso Matilla LA, Carretero JL, Garcia-Carrascosa A. (Eds) Islas Columbretes. Contribución al estudio de su medio natural. Generalitat Valenciana, Valencia, 155–179. http://www.llig.gva.es/es/1361-9788478903511.html.
  • Martin-Eauclaire M-F, Bosmans F, Céard B, Diochot S, Bougis PE (2014) A first exploration of the venom of the Buthus occitanus scorpion found in southern France. Toxicon 79: 55–63. https://doi.org/10.1016/j.toxicon.2014.01.002
  • Masi L (1912) Note sugli scorpioni appartenenti al R. Museo zoologico di Roma. Memorie della Società Entomologica Italiana 3: 88–108, 120–144.
  • Maupertuis PL (1731) Expériences sur les Scorpions. Histoire de l'Académie royale des sciences, avec les mémoires de mathématique et de physique 1731: 223–229. http://biodiversitylibrary.org/item/86706.
  • Mello-Leitão C de (1931) Divisão e distribuição do gênero Tityus Koch. Annaes da Academia Brasileira de Ciências 3: 119–150.
  • Mello-Leitão C de (1939) Revisão do gênero Tityus. Physis 17: 57–76.
  • Mello-Leitão C de (1945) Escorpiões Sul-Americanos. Arquivos do Museu Nacional, Rio de Janeiro 40: 1–468.
  • Michalis K, Dolkeras P (1989) Beitrag zur Kenntnis der Skorpione Thessaliens und Epirus (Nordgriechenland). Entomologische Mitteilungen aus dem Zoologischen Museum Hamburg 9: 259–270.
  • Monard A (1939) Résultats de la mission scientifique du Dr Monard en Guinée Portugaise. 1937-1938. IV. Scorpions. Arquivos do Museu Bocage 10: 81–85.
  • Moriggi M (1941) Gli scorpioni dell’ Africa Orientale Italiana. Rivista di Biologia Coloniale, Roma 4: 77–103.
  • Moritz M, Fischer SC (1980) Die Typen der Arachniden-Sammlung des Zoologischen Museums Berlin. III. Scorpiones. Mitteilungen aus dem Zoologischen Museum in Berlin 2: 309–326.
  • Müller K (1848) Fragmenta Historicum Graecorum. Ambrosio Firmin Didot, Paris, 630 pp.
  • Nenilin AB, Fet V (1992) Zoogeographical analysis of the world scorpion fauna (Arachnida, Scorpiones). Arthropoda Selecta 1: 3–31.
  • Noël F (1824) Dictionnaire historique des personnages célèbres de l'antiquité, princes, généraux, philosophes, poëtes, artistes, artistes etc.; des dieux, héros de la fable; des villes, fleuves, etc. Seconde Éd. Le Normant Père, Paris, 581 pp.
  • Olson DM, Dinerstein E, Wikramanayake ED, Burgess ND, Powell GVN, Underwood EC, D’amico J a, Itoua I, Strand HE, Morrison JC, Loucks CJ, Allnutt TF, Ricketts TH, Kura Y, Lamoreux JF, Wettengel WW, Hedao P, Kassem KR (2001) Terrestrial Ecoregions of the World: A New Map of Life on Earth. BioScience 51: 933–938. https://doi.org/10.1641/0006-3568(2001)051[0933:TEOTWA]2.0.CO;2
  • Pallary P (1928) Description de quatre scorpions nouveaux de la Berbérie. Bulletin du Muséum National d'Histoire Naturelle, Paris, 1ere série 5: 346–351.
  • Pallary P (1929) Les scorpions du Sahara central. Bulletin de la Société d'Histoire Naturelle de l'Afrique du Nord 20: 133–141.
  • Pallary P (1934) Scorpions du Sahara central. Mission du Hoggar III (Février à Mai 1928). Mémoires de la Société d'Histoire Naturelle d'Afrique du Nord 4: 90–100.
  • Pallary P (1938) Sur des scorpions de la Berbérie, de la Syrie et du Congo. Archives de l'Institut Pasteur d'Algérie 16: 279–282.
  • Pavesi P (1880) Studi sugli Aracnidi Africani. I. Aracnidi di Tunisia. Annali del Museo Civico di Storia Naturale di Genova 15: 283–388.
  • Pavesi P (1884) Materiali per lo studio della fauna Tunisia raccolta da G. e L. Doria: Aracnidi. Annali del Museo Civico di Storia Naturale di Genova 20: 446–486.
  • Pavesi P (1885) Aracnidi raccolti dal conte Bouturlin ad Assab e Massaua. Bollettino della Società Entomologica Italiana 17: 197–200.
  • Pavlovsky EN (1924) On the morphology of the male genital apparatus in scorpions. Trudy Leningradskogo Obshchestva Yestestvoispytatelei 53: 17–86.
  • Pavlovsky EN (1925) Zur morphologie des weiblichen Genitalapparates und zur Embryologie der Skorpione. Annuaire du Musée Zoologique de l'Académie des Sciences d'URSS 26: 137–205.
  • Pedroso D, Sousa P, Harris DJ, van der Meijden A (2013) Phylogeography of Buthus Leach, 1815 (Scorpiones: Buthidae): a multigene molecular approach reveals a further complex evolutionary history in the Maghreb. African Invertebrates 48: 298–308. https://doi.org/10.3377/004.048.0216
  • Pérez S (1974) Un inventario preliminar de los escorpiones de la region Paleartica y claves para la identification de los generos de la region Paleartica Occidental. Cátedra de Artrópodos 7: 1–45.
  • Peters W (1861) Ueber eine neue Eintheilung der Skorpione und ueber die von ihm in Mossambique gesammelten Arten von Skorpionen, aus welchem hier ein Auszug mitgetheilt wird. Monatsberichte der Königlich preussischen Akademie der Wissenschaften zu Berlin: 507–516.
  • Piñero FS, Tenorio FU, García FJM (2013) Foraging of Buthus occitanus (Scorpiones: Buthidae) on shrub branches in an arid area of southeastern Spain. Journal of Arachnology 41: 88–90. https://doi.org/10.1636/Hi11-90.1
  • Pocock RI (1889a) Arachnida, Chilopoda and Crustacea. In “III. On the Zoology of the Afghan Commission Delimitation. by J. E. T. Aitchison [Ed.].” (p. 53-142). Transactions of the Linnaean Society of London, 2nd Ser–ies, Zoology 5: 110–121.
  • Pocock RI (1889b) Notes on some Buthidae, new and old. Annals and Magazine of Natural History 3: 334–351.
  • Pocock RI (1890) A revision of the genera of Scorpions of the family Buthidae with descriptions of some South-African species. Proceedings of the Zoological Society of London 10: 114–141.
  • Pocock RI (1893) Notes on the classification of Scorpions, followed by some observations upon synonymy, with descriptions of new genera and species. Annals and Magazine of Natural History 6: 303–330.
  • Pocock RI (1895) On the Arachnida and Myriapoda obtained by Dr. Anderson’s collector during Mr. T. Bent’s expedition to the Hadramaut, South Arabia, with a Supplement upon the Scorpions obtained by Dr. Anderson in Egypt and the Eastern Soudan. Addendum: List of the Scorpi. Zoological Journal of the Linnaean Society, London 25: 292–316.
  • Pocock RI (1899) On the Scorpions, Pedipalps and Spiders from tropical West-Africa represented in the collection of the British Museum. Proceedings of the Zoological Society of London 1899: 833–885. http://biodiversitylibrary.org/page/31572831.
  • Pocock RI (1900a) Arachnida. Fauna of British India including Ceylon and Burma. Taylor and Francis, London, 279 pp.
  • Pocock RI (1900b) On a Collection of Insects and Arachnids made in 1895 and 1897, by Mr. C. V. A. Peel, F.Z.S., in Somaliland, with Descriptions of new Species. 10. General List of the Scorpions of Somaliland and the Boran Country. Philosophical Magazine series 3 32: 54–63. https://doi.org/10.1111/j.1096-3642.1890.tb01694.x
  • Polis GA (1990) The Biology of Scorpions. Polis GA (Ed). Stanford University Press, Standford, 587 pp.
  • Prost A (1982) L’identification des scorpions de Haute-Volta. Notes et Documents Voltaiques 13: 4–10.
  • Reichholf JH, Steinbach G eds. (1992) Skorpione. In: Die Grosse Bertelsmann Lexikothek. Naturenzyklopädie Europas. Tome 4. Bertelsmann Verlag, 32–35.
  • Risso A (1826) Les Scorpionides – Histoire Naturelle des principales productions de l'Europe méridionale et particulièrement de celles des environs de Nice et des Alpes Maritimes. Tome cinquième. F.-G. Levrault, Paris, 155–157.
  • Roewer CF (1943) Über eine neuerworbene Sammlung von Skorpionen des Natur-Museums Senckenberg. Senckenbergiana 26: 205–244.
  • Rossi A (2012) Notes on the distribution of the species of the genus Buthus (Leach, 1815) (Scorpiones, Buthidae) in Europe, with a description of a new species from Spain. Bulletin of the British Arachnological Society 15: 273–279.
  • Rossi A (2017) The genus Buthus Leach , 1815 in the basin countries of the Gulf of Guinea with the description of a new species from Ghana (Scorpiones: Buthidae). Onychium 13: 9–15. https://doi.org/10.5281/zenodo.546330
  • Rossi A, Tropea G (2016a) 1 On the presence of the genus Buthus Leach, 1815 in Sudan with the description of a new species from the enclave of Karora (Scorpiones: Buthidae). Onychium 12: 3–10. https://doi.org/10.13156/arac.2012.15.1.273
  • Rossi A, Tropea G (2016b) A complementary study on the genus Buthus Leach, 1815 in Sudan with the description of a new species (Scorpiones: Buthidae). Arachnida - Rivista Aracnologica Italiana 8: 24–31.
  • Sabaj MH (2016) Standard symbolic codes for institutional resource collections in herpetology and ichthyology: an Online Reference. Version 6.5 (16 August 2016). American Society of Ichthyologists and Herpetologists, Washington, DC. http://www.asih.org/.
  • Sadílek D, Nguyen P, Koç H, Kovařík F, Yağmur EA, St’áhlavský F (2015) Molecular cytogenetics of Androctonus scorpions: an oasis of calm in the turbulent karyotype evolution of the diverse family Buthidae. Biological Journal of the Linnean Society 115: 69–76.
  • Sadine SE, Alioua Y, Chenchouni H (2012) First data on scorpion diversity and ecological distribution in the National Park of Belezma, Northeast Algeria. Serket 13: 27–37.
  • Sadine SE, Bissati S, Lourenço WR (2015) The first true deserticolous species of Buthus Leach, 1815 from Algeria (Scorpiones: Buthidae); Ecological and biogeographic considerations. Comptes Rendus Biologies 339: 44–49. https://doi.org/10.1016/j.crvi.2015.10.002
  • Sadine SE, Samia B, Ould Elhadj MD (2011) Premieres donnees sur la diversite scorpionique dans la region du Souf (Algerie). Arachnides 61: 1–10.
  • Schenkel E (1932) Notizen üeber einige Scorpione und Solifugen. Revue Suisse de Zoologie 39: 375–396.
  • Schenkel E (1938) Spinnentiere von der Iberischen Halbinsel, gesammelt von Prof. Dr O. Lundblad, 1935. Arkiv för Zoologi 30A: 1–29.
  • Schenkel E (1949) Mitteilungen über Spinnentiere. Skorpione aus Marokko. Verhandlungen der Naturforschenden Gesellschaft in Basel 60: 186–201.
  • Sergent E (1938) Iconographie des Scorpions de l'Afrique du Nord. Archives de l'Institut Pasteur d'Algérie 16: 513–522.
  • Sergent E (1941a) Sur le postabdomen (queue) de quelques scorpions de l'Afrique du Nord. Archives de l'Institut Pasteur d'Algérie 19: 353–357.
  • Sergent E (1941b) Sur le scorpion commun de l'Atlas blidéen, Buthus occitanus Amx. Archives de l'Institut Pasteur d'Algérie 19: 447–448.
  • Serre M de (1822) Essai pour server a l’histoire des animaux du midi de la France. Gabon, Paris and Montpellier, 95 pp.
  • Sharma PP, Fernández R, Esposito LA, González-Santillán E, Monod L (2015) Phylogenomic resolution of scorpions reveals multilevel discordance with morphological phylogenetic signal. Proceedings of the Royal Society B: Biological Sciences 282: 2014295. https://doi.org/10.1098/rspb.2014.2953
  • Shulov A, Amitai P (1959) Observations sur les Scorpions. Buthus occitanus ssp. mardochei var. israelis var. nov. Archives de l'Institut Pasteur d'Algérie 37: 218–225.
  • Simon E (1872) Arachnides de Syrie rapportés par M. Charles Piochard de la Brûlerie (Scorpions et Galeodes). Annales de la Société Entomologiqe de France, 5eme série 2: 245–266.
  • Simon E (1878) Études arachnologiques. 7e Mémoire. XIII. Descriptions de deux espèces de Scorpions. Annales de la Société Entomologiqe de France, 5eme série 8: 158–160.
  • Simon E (1879) 3e Ordre. Scorpiones Thorell. In: Les Arachnides de France. VII. Les ordes des Chernetes, Scorpiones et Opiliones. Librairie Encyclopédique de Roret, Paris, 79–115.
  • Simon E (1880) M. E. Simon montre quelques Scorpions qui lui ont été donnés par notre confrère M. Reiche, de la part de M. F. de Saulcy, qui les a reçus de Mossoul (ancienne Ninive), sur le Tigre, en Mésopotamie. Annales de la Société Entomologiqe de France, 5eme série 10: 29.
  • Simon E (1885) Ètude sur les Arachnides recueillis en Tunisie en 1883 et 1884 par MM. A. Letourneux, M. Sédillot et Valery Mayet, membres de la Mission de l'Exploration scientifique de la Tunisie. In: Exploration scientifique de la Tunisie. Zoologie – Arachnides. Imprimerie Nationale, Paris, 55 pp.
  • Simon E (1899) Liste des arachnides recueillis en Algérie par M. P. Lesne et descrition d'une espèce nouvelle. Bulletin du Muséum d'Histoire Naturelle, Paris 5: 82–87.
  • Simon E (1910) Révision des Scorpions d'Egypte. Bulletin de la Société Entomologique de d'Egypte 2: 57–87.
  • Sissom WD (1990) Systematics, biogeography and paleontology. In: Polis GA (Ed.) Biology of scorpions. Stanford University Press, Stanford, 64–160.
  • Skutelsky O (1995) Flexibility in foraging tactics of Buthus occitanus scorpions as a response to above-ground activity of termites. Journal of Arachnology 23: 46–47.
  • Skutelsky O (1996) Predation risk and state-dependent foraging in scorpions: effects of moonlight on foraging in the scorpion Buthus occitanus. Animal Behaviour 52: 49–57. https://doi.org/10.1006/anbe.1996.0151
  • Sousa P, Froufe E, Alves PC, Harris DJ (2010) Genetic diversity within scorpions of the genus Buthus from the Iberian Peninsula: mitochondrial DNA sequence data indicate additional distinct cryptic lineages. Journal of Arachnology 38: 206–211. https://doi.org/10.1636/H08-98.1
  • Sousa P, Harris DJ, Froufe E, van der Meijden A (2012) Phylogeographic patterns of Buthus scorpions (Scorpiones: Buthidae) in the Maghreb and South-Western Europe based on CO1 mtDNA sequences. Journal of Zoology 288: 66–75. https://doi.org/10.1111/j.1469-7998.2012.00925.x
  • Sousa P (2017) Resilient arthropods: Buthus scorpions as a model to understand the role of past and future climatic changes on Iberian Biodiversity. PhD Thesis, Porto, Universidade do Porto, Portugal.
  • Stahnke HL (1972) A key to the genera of Buthidae (Scorpionida). Entomological News 83: 121–133.
  • Táborský K (1934) Sur les espèces du genre Buthus Leach du Muséum National de Prague (in Czech). Sborník Zoologického Oddĕlení Národního Musea v Praze [Acta Zoologica Musaei Nationalis Pragae] 1: 39–40.
  • Teruel R, Melic A (2015) Orden Scorpiones. Revista IDE@ - SEA 18: 1–17.
  • Teruel R, Pérez-Bote JL (2005) Complementos a la descripción de Buthus ibericus Lourenço and Vachon 2004 (Scorpiones: Buthidae). Boletín de la Sociedad Entomológica Aragonesa 37: 273–275.
  • Thorell T (1876a) Études scorpiologiques. Atti della Società Italiana di Scienze Naturali 19: 75–272.
  • Thorell T (1893) Scorpiones exotici R. Musei Historiea Naturalis Florentini. Bolletino della Società Entomologica Italiana 25: 356–387.
  • Touloun O (2012) Chapitre III. Ecologie et repartition de la scorpiofaune du Sud Ouest Marocain. In: Les peuplements de scorpions du sud ouest marocain: Ecologie, Biogéographie et Epidémiologie des envenimations. Éditions universitaires européennes, Saarbrücken, 30–82.
  • Touloun O, Boumezzough A (2011b) Contribution à l'inventaire et à la répartition des scorpions de la province de Sidi Ifni (Maroc). Poiretia 3: 8–15.
  • Touloun O, El Hidan MA, Boumezzough A (2016) Species composition and geographical distribution of Saharan scorpion fauna, Morocco. Asian Pacific Journal of Tropical Disease 6: 878–881. https://doi.org/10.1016/S2222-1808(16)61150-7
  • Touloun O, Elhidan MA, Boumezzough A (2014) Inventory and distribution of scorpion fauna in eastern Morocco (Arachnida, Scorpiones). Serket 14: 72–79.
  • Touloun O, Slimani T, Boumezzough A (1999) Decouverte au Maroc de Buthus occitanus tunetanus var. neeli Gysin, 1969 (Scorpiones, Buthidae). Arachnides 41: 28–30.
  • Tullgren A (1909) Solifugae, Scorpiones und Chelonethi aus Ägypten und dem Sudan. In: Results of the Swedish Zoological Expedition to Egypt and the White Nile 1901, under the direction of L.A. Jägerskiöld. Part III. The Library of the Royal University of Uppsala, Uppsala, 1–12.
  • Vachon M (1940) Sur la systématique des scorpions. Mémoires du Muséum national d'Histoire naturelle, Nouvelle Série, Paris 13: 241–260.
  • Vachon M (1941) Remarques biogéographiques sur quelques Scorpions et Pseudoscorpions prédésertiques. Compte-rendu sommaire des Séances de la Société de Biogéographie 18: 50–53.
  • Vachon M (1948a) Études sur les Scorpions. III. Déscription des Scorpions du Nord de l'Afrique. Archives de l'Institut Pasteur d'Algérie 26: 162–208.
  • Vachon M (1948b) 3Études sur les Scorpions. III (suite). Déscription des Scorpions du Nord de l'Afrique. Archives de l'Institut Pasteur d'Algérie 26: 288–316.
  • Vachon M (1948c) Scorpions récoltés dans l'ile de Crète par Mr le docteur Otto Von Wettstein. Annalen des Naturhistorischen Museums in Wien 56: 61–69. http://www.jstor.org/stable/41768480.
  • Vachon M (1949a) Études sur les Scorpions. III (suite). Déscription des Scorpions du Nord de l'Afrique. Archives de l'Institut Pasteur d'Algérie 27: 134–169.
  • Vachon M (1949b) Études sur les Scorpions. III (suite). Déscription des Scorpions du Nord de l'Afrique. Archives de l'Institut Pasteur d'Algérie 27: 281–288.
  • Vachon M (1949c) Études sur les Scorpions. III (suite). Déscription des Scorpions du Nord de l'Afrique. Archives de l'Institut Pasteur d'Algérie 27: 334–396.
  • Vachon M (1950a) Études sur les Scorpions. III (suite). Détermination des scorpions du Nord-Ouest de l'Afrique. Archives de l'Institut Pasteur d'Algérie 28: 152–216.
  • Vachon M (1950b) Études sur les Scorpions. (suite). Déscription des Scorpions du Nord-ouest de l'Afrique. Archives de l'Institut Pasteur d'Algérie 28: 382–413.
  • Vachon M (1951a) Études sur les Scorpions. (suite et fin). Détermination des Scorpions du Nord-Ouest de l'Afrique. Archives de l'Institut Pasteur d'Algérie 29: 46–104.
  • Vachon M (1951b) Scorpions collectés au Maroc par MM. P. Strinati et V. Aellen (Mission scientifique Suisse au Maroc, août-septembre 1950). Bulletin du Muséum National d'Histoire Naturelle, Paris, 2eme série 23: 621–623.
  • Vachon M (1952a) Études sur les scorpions. (published 1948–1951 in Archives de l'Institut Pasteur d'Algérie, 1948, 26: 25–90, 162–208, 288–316, 441–481; 1949, 27: 66–100, 134–169, 281–288, 334–396; 1950, 28: 152–216, 382–413; 1951, 29: 46–104). Institut Pasteur d'Algérie, Alger, 482 pp. https://www.ntnu.no/ub/scorpion-files/vachon.php.
  • Vachon M (1952b) Le Scorpion Languedocien et sa répartition en France à propos de sa présence dans le Département de l'Ardèche. Bulletin du Muséum National d'Histoire Naturelle, Paris, 2eme série 24: 274–279.
  • Vachon M (1953) Contribution à l'étude du peuplement de la Mauritanie. Scorpions. Bulletin de l'Institut français d'Afrique Noire 15: 1012–1028.
  • Vachon M (1955) Remarques préliminaires sur la faune des scorpions du Soudan Oriental. Bulletin du Muséum National d'Histoire Naturelle, Paris, 2e Série 27: 371–373.
  • Vachon M (1958) Scorpionidea (Chelicerata) de I'Afghanistan. The 3rd Danish Expedition to Central Asia. (Zoological Results 23). Videnskabelige Meddelelser fra Dansk Naturhistorisk Forening i Købehavn 120: 121–187.
  • Vachon M (1961) Le Parc national de Niokolo-Koba. III. Scorpions. Mémoires de l'Institut Français d'Afrique Noire 62: 31–32.
  • Vachon M (1963) De l'utilité, en systématique, d'une nomenclature des dents de chelicères chez les Scorpions. Bulletin du Muséum National d'Histoire Naturelle, Paris, 2e Série 35: 161–166.
  • Vachon M (1966) Liste des scorpions connus en Egypte, Arabie, Israeli, Liban, Syrie, Jordanie, Turquie, Irak, Iran. Toxicon 4: 209–218.
  • Vachon M (1975) Sur l'utilisation de la trichobothriotaxie du bras des pédipalpes des scorpions (Arachnides) dans le classement des genres de la famille des Buthidae Simon. Comptes rendus hebdomadaires des séances de l'Académie des sciences, Série D, Sciences naturelles 281: 1597–1599.
  • Vachon M, Kinzelbach R (1987) On the taxonomy and distribution of the scorpions of the Middle East. In: Krupp F, Schneider W, Kinzelbach R (Eds) Proceeding of the Symposium on the Fauna and Zoogeography of the Middle East, Mainz, 1985. (Beihefte zum Tübingen Atls des Vorderen Orients). Reichert Verlag, Wiesbaden, 91–103.
  • Vigna Taglianti A, Audisio PA, Biondi M, Bologna MA, Carpaneto GM, De Biase A, Fattorini S, Piattella E, Sindaco R, Venchi A, Zapparoli M (1999) A proposal for a chorotype classification of the Near East fauna, in the framework of the Western Palearctic region. Biogeographia 20: 31–59.
  • Vincent M (1994) Observation de Buthus occitanus (Amoreux, 1789) (Scorpionida, Buthidae) dans la région de Manosque (Alpes de Haute-Provence). Arachnides 22: 6–7.
  • Volschenk ES, Mattoni CI, Prendini L (2008) Comparative anatomy of the mesosomal organs of scorpions (Chelicerata, Scorpiones), with implications for the phylogeny of the order. Zoological Journal of the Linnean Society 154: 651–675. https://doi.org/10.1111/j.1096-3642.2008.00426.x
  • Werner F (1902) Die Scorpione, Pedipalpen und Solifugen in der zoologisch-vergleichend-anatomischen Sammlung der Universität Wien. Verhandlungen der Kaiserlich-Königlichen Zoologish-Botanischen Gesellschaft in Wien 52: 595–608.
  • Werner F (1925) Skorpione und Solifugen aus dem östlichen und nördlichen Spanien, gesammelt von Dr. F. Haas in den Jahren 1914–1918. Senckenbergiana 7: 209–210.
  • Werner F (1929) Wissenschaftliche Ergebnisse einer zoologischen Forschungsreise nach Westalgerien und Marokko. III. Scorpiones. Sitzungsberichte der Kaiserlichen Akademie der Wissenschaften, Mathematisch-Naturwissenschaftliche Klasse, Wien 138: 1–34. http://www.zobodat.at/publikation_articles.php?id=169321.
  • Werner F (1934a) Scorpiones, Pedipalpi. In: Bronn HG (Ed.) Klassen und Ordnungen des Tierreichs. Band 5: Arthropoda, Abteilung 4: Arachnoidea, Buch 8. Akademische Verlagsgesellschaft, Leipzig, 490 pp.
  • Werner F (1934b) Über skorpione aus Marokko. Verhandlungen der Naturforschenden Gesellschaft in Basel 45: 82–88.
  • Werner F (1936) Neu-Eingänge von Skorpionen im Zoologischen Museum in Hamburg. Festschrift zum 60. Geburtstage von Professor Dr. Embrik. Strand, Riga 2: 171–193.