Systematic Revision of the Sand Scorpions, Genus Buthacus Birula, 1908 (Buthidae C.L. Koch, 1837) of the Levant, with Redescription of Buthacus arenicola (Simon, 1885) from Algeria and Tunisia

Shlomo Cain, Eran Gefen, Lorenzo Prendini

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Scorpions of the genus ButhacusBirula, 1908 (Buthidae C.L. Koch, 1837), commonly known as "sand scorpions,"are widespread in the sandy deserts of the Palearctic, from West Africa to India. Although many new species of Buthacus were described in recent years, no modern revision exists for the genus and the limits of many infrageneric taxa remain unclear. The present contribution addresses the species of Buthacus recorded from the Levant, defined here as the region of the Middle East including Syria, Lebanon, Jordan, Israel, the Palestinian territories, and the Sinai Peninsula (Egypt). Prior to this study, five species and subspecies, including several synonyms, were recognized from the region. Based on extensive new collections, a reassessment of the morphology (including multivariate statistical analysis), and a phylogenetic analysis of morphological and DNA sequence data, published elsewhere, seven species of Buthacus are now recognized from the Levant, raising the number of species in the genus to 30. Three new species are described: Buthacus amitaii, sp. nov., endemic to Israel; Buthacus arava, sp. nov., endemic to Israel and Jordan; and Buthacus levyi, sp. nov., endemic to Egypt, Israel, and perhaps Libya. Buthacus arenicola (Simon, 1885) is redescribed and restricted to northeastern Algeria and central Tunisia, and Buthacus leptochelys (Ehrenberg, 1829) redescribed and restricted to Egypt, Sudan, and perhaps Libya. Buthacus armasiLourenço, 2013, stat. rev., from southern Algeria, and Buthacus spatzi (Birula, 1911), stat. rev., from southern Tunisia and western Libya, are revalidated, and Buthacus fuscataPallary, 1929, stat. nov. et stat. rev., from southern Algeria, revalidated and elevated to the rank of species. Buthacus nitzaniLevy et al., 1973, stat. nov., currently restricted to Israel but probably present in the Sinai Peninsula (Egypt), is elevated to the rank of species. Buthacus tadmorensis (Simon, 1892), stat. rev., recorded from Iran, Iraq, Jordan, Syria, and Turkey, and Buthacus yotvatensisLevy et al., 1973, stat. rev., endemic to Israel and Jordan, are redescribed and revalidated. Three new synonyms are presented: Androctonus (Leiurus) macrocentrus Ehrenberg, 1828 = Buthacus leptochelys (Ehrenberg, 1829), syn. nov.; Buthus pietschmanniPenther, 1912 = Buthacus tadmorensis (Simon, 1892), syn. nov.; Buthacus granosusBorelli, 1929 = Buthacus leptochelys (Ehrenberg, 1829), syn. nov. Buthacus arenicola and the seven species of Buthacus occurring in the Levant are diagnosed and illustrated to modern standards, with updated distribution maps. A list of the currently recognized species of Buthacus, and a key to identification of the species occurring in the Levant are also presented.

Original languageEnglish
Pages (from-to)1-136
Number of pages136
JournalBulletin of the American Museum of Natural History
Issue number1
StatePublished - 1 Apr 2021

Bibliographical note

Funding Information:
This research was funded by U.S.-Israel Binational Science Foundation award 47441 to E.G. and L.P. The 26th European Congress of Arachnology and Ben-Gurion University of the Negev (BGU), Israel, partially supported L.P.’s visit to Israel in 2011, and the Israel Taxonomy Initiative and BGU (Jacob Blaustein Center for Scientific Cooperation) supported his visit in 2013. We thank Efrat Gavish-Regev (HUJ), Sergei Zonstein and Ariel-Leib-Leonid Friedman (SMNH), and Christoph Hörweg (NHMW) for lending type and nontype material from collections in their care; Jason Dunlop (ZMB) and Elise-Anne Leguin (MNHN) for images of type material; Rotem Agmon, Ayelet Allon, Nadav Amir, Zuhair Amr, Essam Attia, Jamel Babay, Tharina Bird, Noach Braun, Efrat Gavish-Regev, Gal Geisler, Alexander V. Gromov, Fenik S. Hussen, Colleen Irby, Manel Khammasi, Halil Koç, David Kotter, Reut Less, Snir Livne, Yael Lubin, Mor Lugassi, Shahrokh Navidpour, Asia Novikova, Itamar Ofer, Yael Olek, Ridha Ouina, Avner Rinot, Nitzan Segev, Boaz Shacham, Stav Talal, Itay Tesler, Ersen A. Yağmur, and Yoram Zvik for donating material and/or assisting the authors with fieldwork; the Israel Nature and National Parks Protection Authority for permission to collect and export scorpions from Israel (permits 2011/38157 and 2013/39974 to L.P. 2015/41081 and 2017/41728 to E.G.); Essam Attia and Hisham K. El-Hennawy for hospitality, logistics and permissions in Egypt; Zuhair Amr and the Royal Jordanian Society for the Conservation of Nature for hospitality, logistics, and permissions in Jordan; Saïd Nouira, Jamel Babay, and Ridha Ouina for hospitality and logistics in Tunisia; Diogo Casellato, Deborah Chin, Ofelia Delgado-Hernandez and Tarang Sharma for generating DNA sequence data at the AMNH; Andrew Fatiukha, Rachel Ben-Shlomo, Stav Talal, and Tamar Krugman for assistance with generating sequence data at the Institute of Evolution, University of Haifa; Emile Cain and Myriam Freund for assisting with French text translations; Stephanie Loria (AMNH) for assistance with phylogenetic analysis; Steve Thurston (AMNH) for assistance with photography and preparing the plates; Pio Colmenares and Lou Sorkin (AMNH) for assistance with import/export logistics from the U.S.; and Oscar F. Francke and an anonymous reviewer for constructive comments on a previous draft of the manuscript.

Publisher Copyright:
Copyright © American Museum of Natural History 2021.

ASJC Scopus subject areas

  • Ecology
  • Agricultural and Biological Sciences (miscellaneous)


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