Dated phylogeny and ancestral range estimation of sand scorpions (Buthidae: Buthacus) reveal Early Miocene divergence across land bridges connecting Africa and Asia

Shlomo Cain, Stephanie F. Loria, Rachel Ben-Shlomo, Lorenzo Prendini, Eran Gefen

Research output: Contribution to journalArticlepeer-review

Abstract

Sand scorpions of the genus Buthacus Birula, 1908 (Buthidae C.L. Koch, 1837) are widespread in the sandy deserts of the Palearctic region, occurring from the Atlantic coast of West Africa across the Sahara, and throughout the Middle East to Central Asia. The limits of Buthacus, its two species groups, and many of its species remain unclear, and in need of revision using modern systematic methods. The study presented here set out to investigate the phylogeny and biogeography of the Buthacus species occurring in the Levant, last studied in 1980. A phylogenetic analysis was performed on 104 terminals, including six species collected from more than thirty localities in Israel and other countries in the region. Three mitochondrial and two nuclear gene loci were sequenced for a total of 2218 aligned base-pairs. Morphological datasets comprising 22 qualitative and 48 quantitative morphological characters were compiled. Molecular and morphological datasets were analyzed separately and simultaneously with Bayesian Inference, Maximum Likelihood, and parsimony. Divergence time and ancestral range estimation analyses were performed, to understand dispersal and diversification. The results support a revised classification of Levantine Buthacus, and invalidate the traditional species groups of Buthacus, instead recovering two geographically-delimited clades, an African clade and an Asian clade, approximately separated by the Jordan Valley (the Jordan Rift Valley or Syro-African Depression), the northernmost part of the Great Rift Valley. The divergence between these clades occurred in the Early Miocene (ca. 19 Ma) in the Levant, coinciding temporally with the existence of two land bridges, which allowed faunal exchange between Africa and Asia.

Original languageEnglish
Article number107212
JournalMolecular Phylogenetics and Evolution
Volume164
DOIs
StatePublished - Nov 2021

Bibliographical note

Funding Information:
This research was funded by U.S.-Israel Binational Science Foundation award (BSF #2014046) to L.P. and E.G. 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 H?rweg (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, Fenik S. Hussen, Colleen Irby, Manel Khammasi, David Kotter, Reut Less, Snir Livne, Yael Lubin, Mor Lugassi, Asia Novikova, Itamar Ofer, Yael Olek, Ridha Ouina, Avner Rinot, Nitzan Segev, Boaz Shacham, Stav Talal, Itay Tesler, 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, 2017/41728 to E.G. and 2017/41779 to Efrat Gavish-Regev); 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; Sa?d 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, 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; Steve Thurston (AMNH) for assistance with photography; Pio Colmenares and Lou Sorkin (AMNH) for assistance with import/export logistics from the U.S.

Funding Information:
This research was funded by U.S.-Israel Binational Science Foundation award (BSF #2014046) to L.P. and E.G. The 26 th 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 Hӧrweg (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, Fenik S. Hussen, Colleen Irby, Manel Khammasi, David Kotter, Reut Less, Snir Livne, Yael Lubin, Mor Lugassi, Asia Novikova, Itamar Ofer, Yael Olek, Ridha Ouina, Avner Rinot, Nitzan Segev, Boaz Shacham, Stav Talal, Itay Tesler, 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, 2017/41728 to E.G., and 2017/41779 to Efrat Gavish-Regev); 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, 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; Steve Thurston (AMNH) for assistance with photography; Pio Colmenares and Lou Sorkin (AMNH) for assistance with import/export logistics from the U.S.

Publisher Copyright:
© 2021 Elsevier Inc.

Keywords

  • Biogeography
  • Buthidae
  • Chelicerata
  • Phylogeny
  • Scorpiones
  • Systematics

ASJC Scopus subject areas

  • Ecology, Evolution, Behavior and Systematics
  • Molecular Biology
  • Genetics

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