Catalogue |
Corresponding author: Pedro Sousa ( prsousa@gmail.com ) Academic editor: Wilson Lourenço
© 2017 Pedro Sousa, Miquel Arnedo, D. James Harris.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Sousa P, Arnedo MA, Harris JD (2017) Updated catalogue and taxonomic notes on the Old-World scorpion genus Buthus Leach, 1815 (Scorpiones, Buthidae). ZooKeys 686: 15-84. https://doi.org/10.3897/zookeys.686.12206
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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.
Taxonomy, new synonymy, new combination, new status, Geographic distribution, Africa, Asia, Europe, diagnostic characters
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.
Buthus exhibits a wide distribution range, spanning over two biogeographic realms, the Palearctic (Western) and the Afrotropical (
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.
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
Buthus is the type genus of the Buthidae C. L. Koch, 1837 (
Buthus was first proposed by
The poor description of
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 (
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 (
During the last 15 years, the rate of description of new Buthus species has increased exponentially (Fig.
Nomenclature and measurements follow Stahnke (1970), except for trichobothriotaxy (
Most references prior to 1998 cited by
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
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
MCVR Museo Civico di Storia Naturale di Verona, Verona, Italy
MTAS * Museum of the Turkish Arachnology Society, Ankara, Turkey
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
ZINZoological Institute, Russian Academy of Sciences, St. Petersburg, Russia (formerly ZISP)
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
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
Original drawings of the habits of Scorpio europaeus Linnaeus, 1758 (A) and S. occitanus (B), according to
Finally, although Buthus is considered the nominal genus of Buthidae,
The revised classification of
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
There are no taxonomically distinct groups within the genus Buthus, although two “species complexes” are generally recognised.
Recently, based on the information provided by a cox1 mtDNA tree,
Current composition of the groups proposed by
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.
Map representing the five phylogenetic Buthus cox1 groups in the Maghreb. Groups as defined by
Several morphological traits have been used by recent authors as diagnostic characters (in the sense of
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.
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.
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.
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
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
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.
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
Confirming the fast pace of new Buthus species descriptions, a new species, Buthus danyii Rossi, 2017 was published from Ghana (
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.
Buthus
:
(by original designation). Scorpio occitanus Amoreux, 1789 [=Buthus occitanus (Amoreux, 1789)].
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 (
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
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
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 |
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 (
Buthus
adrianae
:
1 adult M holotype (
known only from the type locality.
Buthus
albengai
:
1 F holotype (
known from an area in Morocco that extends from ifrane to Kenifra.
It is one of the largest known Buthus species. Records by
Buthus
amri
: Lourenço, Yağmur and Duhem 2010: 96–99, fig. 1–5;
1 M holotype (
known only from the type locality.
= Tityus tenuimanus
Buthus
atlantis
:
Buthus
Atlantis
(sic):
Buthus
atlantis
atlantis
:
Buthus
occitanus
atlantis
:
Buthus (Buthus) atlantis
:
Tityus
tenuimanus
:
1 F holotype (
known to occur only in sandy dune habitats close to the Atlantic Ocean in Morocco, between Essaouira and Agadir.
It is the largest known Buthus species.
Buthus
aures
1 M holotype (
known only from the type locality.
Buthus
awashensis
:
Buthus occitanus (MIS): Kovařík and Whitman 2005 (part): 106.
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.
know only from two Ethiopian localities, more than 200 km apart.
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
occitanus
barcaeus
:
Buthus (Buthus) occitanus barcaeus
:
Buthus
barcaeus
:
4 M, 1 F juv., syntypes (
know from several localities along the Mediterranean coast of Libya.
The specimens present in the
= Buthus occitanus zeylensis
Buthus
occitanus
berberensis
:
Buthus
occitanus
zeylensis
:
Buthus
berberensis
:
Buthus (Buthus) occitanus berberensis
:
1 M holotype (
know from Djibouti, Eritrea, Ethiopia and Somalia, although
= Buthus sabulicola
Buthus
bonito
:
1 M holotype (
known from the Atlantic coast of Morocco south of Tan-Tan extending almost to Dakhla in the Western Sahara (
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.
Buthus
boumalenii
:
1 F holotype (UCAM), Tineghir (approx. 31.366°, -5.905°), Boumalene, Morocco. Paratypes: 1 M (UCAM), 1 M, 1 F (
known only from the Boumalne region of Morocco (El Hidan et al. 2016).
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
:
1 F holotype (
known only from the type locality.
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
:
1 M holotype (
known only from the type locality.
Buthus
chambiensis
:
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.
known only from the type locality.
Buthus
confluens
: Lourenço, Touloun and Boumezzough 2012: 22–24, fig. 1–11;
1 M holotype (
known from several localities in the Tingitana Pensinsula of Morocco, but also further to the south.
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
:
Buthus
occitanus
tunetanus
neeli
(MIS):
Buthus
tassili
(MIS):
1 M holotype (UCAM), Taznakht (30.51853°, -7.02595°), Ouarzazate, Morocco. Paratypes: 1 M, 2 F (UCAM), 1 M, 1 F (
B. draa can be found in the upper part of the Draa River, probably at elevations below 1500 m a.s.l.
B. draa shares with B. tassili and B. nigrovesiculosus the presence of a darkened fifth metasoma segment and telson.
Buthus
dunlopi
:
1 M holotype (FKCP), Remada (approx. 32.31°, 10.39°), Tataouine, Tunisia. Paratypes: 1 M, 3 F (FKCP), same locality.
known only from the type locality.
Buthus
duprei
1 M holotype (
known only from the type locality.
Buthus
egyptiensis
:
1 F holotype (
known only from the type locality.
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
:
1 M holotype (
this species is known from Niger and Senegal, from a single locality in each country, which are almost 2,000 Km apart.
We used the location of Fété-Olé given in
Buthus
elizabethae
:
1 M holotype (
this species is known from Guinea and Senegal,
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
:
Buthus
occitanus
mardochei
alluaudi
:
1 M holotype (
this species seems to be widely distributed across the western portion of the Anti-Atlas, although some misidentifications with Buthus parroti cannot be excluded.
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
:
1 adult M holotype (
this species is known from the southern Iberian Mediterranean coast, close to Marbella.
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.
Buthus
hassanini
:
1 F (
known only from the type locality.
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 halius (C. L. Koch, 1839) (nomen oblitum) (comb. n., syn. n.). Holotype lost according to
Buthus
ibericus
Lourenço & Vachon, 2004: 88–91, fig. 31–42,
Androctonus
halius
C. L.
Buthus
occitanus
(MIS):
1 M holotype (
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.
B. ibericus was first described from Spain and subsequently reported for Portugal (e.g.
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.
Androctonus (Leirus) tunetanus intermedius
: Ehrenberg in
Androctonus (Leiurus) tunetanus intumescens
(MIS):
Androctonus (Liurus) tunetanus intermedius
: Ehrenberg in
Androctonus
occitanus
intermedius
:
Buthus (Buthus) occitanus intermedius
:
Buthus occitanus intermedius : Peréz 1974: 23.
Buthus
intermedius
(Ehrenberg):
1 F (in bad conditions) (
Known only from the type locality.
Androctonus (Leiurus) tunetanus intumescens
: Ehrenberg in
Androctonus (Liurus) tunetanus intumescens
: Ehrenberg in
Androctonus
occitanus
intumescens
:
Buthus
intumescens
:
1 (sex unknown) (in bad conditions) (
known only from a single specimen.
Buthus occitanus mardochei israelis
:
Buthus
occitanus
israelis
:
Buthus
occitanus
israelis
(Shulov & Amitai, 1959):
Buthus
intumescens
(MIS):
Buthus israelis : Lourenço, Yağmur and Duhem 2010: 96–97; Yağmur, Koç and Lourenço 2011: 29.
Buthus
israelis
(Shulov & Amitai, 1959):
Buthus occitanus mardochei israelis
:
Buthus
occitanus
typicus
:
holotype lost (sex unknown), Mash’abbe Sade (as Mashavei Sadé) (approx. 31°, 34.78°), Negev desert, Israel.
This species has been recorded in Egypt (the Sinai Peninsula) and Israel. Notwithstanding,
Buthus
jianxinae
:
1 M holotype (
known only from the type locality.
Buthus
karoraensis
:
Buthus
occitanus
berberensis
(MIS):
Buthus occitanus (MIS): Kovařík and Whitman 2005 (part): 106.
1 M holotype (
known only from the type locality.
Buthus kunti : Yağmur, Koç and Lourenço 2011: 29–33, fig. 1–12.
Buthus
europaeus
(MIS):
Buthus
occitanus
(MIS):
Buthus
sp
.:
1 F holotype (MTAS), Rizokarpaso (Dipkarpaz) (35.58472°, 34.42306°), Karpaz Region, Cyprus. Paratypes: 1 M juv. (MTAS), Zafer. 1 M juv. (
the species is only known from the northern portion of Cyprus.
according to Yağmur, Koç and Lourenço (2011), this species is rare in the island.
Buthus
labuschagnei
1 F holotype (
known only from the type locality.
Buthus
lienhardi
:
Buthus
occitanus
tunetatus
Lepineyi
: Vachon 1949: 353–359, fig. 393–400;
Buthus
occitanus
tunetatus
lepineyi
:
Buthus
occitanus
tunetatus
(MIS):
1 M holotype (
the species is known from a wide range across the High-Atlas Mountains.
Vachon (1949) infrasubspecific name is not available as explained previously.
Buthus
lourencoi
:
Buthus occitanus (MIS): Kovařík and Whitman 2005 (part): 106.
1 adult F holotype (
known only from the type locality.
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
occitanus
malhommei
Vachon 1949: 376;
Buthus
malhommei
:
3 M, 3 F, 7 juv., syntypes (
Toulon (2012) greatly expanded the known distribution of this species along the basin of the Oum er Rbia River.
Buthus
mardoche
(IOS):
Buthus (Buthus) mardoche
(IOS):
Buthus
occitanus
mardochei
:
Buthus
occitanus
mardochei
mardochei
: Le Corroller, 1967. 63;
Buthus
mardochei
:
1 F (
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.
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
:
Buthus
occitanus
mardochei
mimeuri
:
1 F holotype (
this species has a large distribution in Morocco, east and south of the Anti-Atlas Mountain.
Vachon (1949) infrasubspecific name is not available as explained above.
= Prionurus tingitanus:
Buthus
occitanus
maroccanus
:
Buthus
europaeus
(MIS):
Buthus (Buthus) occitanus maroccanus
:
Buthus
occitanus
maroccanus
:
Buthus
maroccanus
:
Buthus marocanus (ISS): Le Corroller 1967: 63.
3 M, F, syntypes (
all known specimens have been captured in the Rabat Region (approx. 28.43°, -11.10°).
it remains the only known Buthus species with a uniformly darkened body.
Buthus
montanus
1 M holotype (
known only from the type locality.
Buthus
europaeus
nigrovesiculosus
:
Buthus
occitanus
nigrovesiculosus
:
1 M (adult?), 1 juv., syntypes (
known only from the type locality.
Hirst identified this North African species as a subspecies of B. occitanus, but as currently circumscribed, B. occitanus does not occur in North Africa (
Detailed morphology of the larger syntype of B. nigrovesiculosus (
Buthus
occidentalis
:
1 F holotype (
known only from the type locality.
= Scorpio rufus:
= Androctonus ajax: C. L.
= Androctonus eurialus: C. L.
= Androctonus eurilochus: C. L.
= Buthus europaeus:
tridentatus
= Buthus europaeus tridentatus:
Scorpio
occitanus
:
Androctonus
ajax
: C. L.
Androctonus
euryalus
(ISS): C. L.
Androctonus
eurylochus
(ISS): C. L.
Androctonus (Prionurus) occitanus
:
Buthus
europaeus
:
Buthus
occitanicus
(ISS):
Buthus
occitanus
:
Buthus
occitanus
occitanus
:
Buthus
occitanus
tridentatus
:
Buthus (Buthus) occitanus
:
Buthus (Buthus) occitanus occitanus
:
Buthus
cf.
occitanus
:
Scorpio
australis
(MIS):
Scorpio (Androctonus) occitanus
:
Scorpio
occitanicus
(ISS):
Scorpion Occitanus (ISS):
type unknown, Souvignargues, Occitanie Region, France.
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 (
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.
Buthus
orientalis
:
1 F holotype (
known only from the type locality.
= Androctonus clytoneus: C. L.
Androctonus
paris
: C. L.
Androctonus
clytonicus
(ISS):
Androctonus
clytoneus
: C. L.
Buthus
occitanus
paris
:
Buthus (Buthus) occitanus paris
:
Buthus
paris
:
Holotype lost according to
the species is currently distributed across Algeria, Morocco and Tunisia.
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:
Buthus
atlantis
parroti
:
1 F (
Known only from the Sous River Valley. Type localities in
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 (
Buthus
prudenti
:
1 M holotype (
known only from the type locality.
Buthus
pusillus
:
1 M holotype (
known only from the type locality.
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
:
Buthus
occitanus
mardochei
panousei
:
1 M holotype (
known distribution confined to the Tafnidilt region of Morocco.
Buthus
saharicus
: Saddine et al. 2015: 47–49, fig. 6–8;
1 F holotype (
known only from the type locality.
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
:
Buthus
occitanus
(MIS):
Buthus
occitanus
tunetanus
neeli
:
Buthus
occitanus
tunetatus
, “Spécimens des régions montagneuses centrales du Sahara”:
1 M holotype (
This species is known from a wide area around the Hoggar and Tassili N’Ajjer Mountains, including at least one locality in Libya.
The Tin Tazarif coordinates given here, standing at 880 m a.s.l., do not match the altitude given for the point by
Buthus
trinacrius
:
Buthus
europaeus
(MIS):
Buthus
occitanus
(MIS):
1 M holotype, Palermo Province? (approx. 38.05°, 13.32°), Sicily. Paratypes: 1 M, 1 F. All type material in bad conditions (
Recorded from Sicily.
Although
Scorpio
tunetanus
Androctonus (Leiurus) tunetanus
: Ehrenberg in
Androctonus (Leiurus) tunetanus genuinus
: Ehrenberg in
Androctonus (Liurus) tunetanus
: Ehrenberg in
Androctonus (Liurus) tunetanus genuinus
: Ehrenberg in
Androctonus
tunetanus
: C. L.
Buthus occitanus (MIS): Kovařík and Whitman 2005 (part): 106.
Buthus
occitanus
tunetanus
:
Buthus (Buthus) occitanus tunetanus
:
Buthus
tunetanus
:
Scorpion Tunetanus (ISS):
Types lost according to
The species is currently distributed across Algeria, Libya, Morocco and Tunisia, and doubtfully in the island of Malta.
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
Buthus
yemenensis
:
1 F holotype (
Known only from the type locality.
Buthus
albengai
(MIS):
Buthus
europaeus
(MIS):
Buthus
malhommei
(MIS):
Buthus
occitanus
(MIS):
Buthus
occitanus
occitanus
(MIS):
Buthus
occitanus
occitanus
, “Afrique occidentale française”:
Buthus
occitanus
occitanus
, “Cote Occidentale du Maroc”:
Scorpio
occitanus
:
not Buthus:
We futher propose to transfer a species from the genus Buthus to the genus Androctonus.
Buthus
barbouri
:
1 M (type probably lost), Agadir (approx. 30.43°, -9.60°), Morocco.
known only from the type locality, Agadir, just north of the Sous River mouth, in southern Morocco.
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
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
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
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.