Research Article |
Corresponding author: Ahmed Badry ( ahmedbadry@azhar.edu.eg ) Academic editor: Wilson Lourenço
© 2023 Abdulmani H. Al-Qahtni, Abdullah M. Al-Salem, Fahad Mesfer, Manal S. Al Balawi, Wasayf S. Allahyani, Abdulaziz R. Alqahtani, Ahmed Badry.
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:
Al-Qahtni AH, Al-Salem AM, Mesfer F, Al Balawi MS, Allahyani WS, Alqahtani AR, Badry A (2023) A new species and a key to the genus Leiurus Ehrenberg, 1828 (Scorpiones, Buthidae) from Saudi Arabia. ZooKeys 1178: 293-312. https://doi.org/10.3897/zookeys.1178.109083
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A new species, Leiurus hadb Al-Qahtni, Al-Salem, Alqahtani & Badry, sp. nov., is described and illustrated from the Majami al-Hadb Protected Area in the Riyadh Province of Saudi Arabia. The new species is compared with species of Leiurus distributed in Saudi Arabia, especially L. arabicus Lowe, Yağmur & Kovařík, 2014. The integrated results indicate that the population found in Majami al-Hadb represents a distinct species, which is described herein. Moreover, the molecular analysis is conducted on the mitochondrial gene 16S rRNA to compare L. hadb sp. nov. with samples of L. arabicus and L. haenggii from Saudi Arabia. The analysis revealed a genetic divergence ranging from 6.0 to 12%. The combination of molecular evidence and morphological characteristics provides adequate support for recognizing the Majami al-Hadb population as a distinct species. Additionally, an identification key for the genus Leiurus found in Saudi Arabia is also provided.
Description, identification key, Majami al-Hadb Protected Area, molecular phylogeny scorpion, taxonomy
The genus Leiurus was first introduced by Ehrenberg in
In this study, we describe a new species of the genus Leiurus based on several specimens of the Majami al-Hadb Protected Area under an integrative taxonomic perspective, using morphological and molecular evidence.
A total of 11 specimens of Leiurus hadb sp. nov. was collected from Majami al-Hadb Protected Area in the Riyadh Province between 23 and 25 May 2023, using ultraviolet light at night. The specimens were preserved in 96% alcohol and photographed using a Canon EOS 6D Mark II. The photographs were edited using Adobe Photoshop software. The measurements were taken in mm on preserved specimens, according to
The genomic DNA was isolated from five scorpion specimens of L. hadb sp. nov., using Qiagen DNA extraction kits following the manufacturer’s instructions. The amplified 16S rRNA gene products were purified and sequenced using invertebrate universal primers, as determined, and sequenced on an ABI 3500 automated sequencer (Applied Biosystems Inc., USA) and following
Family Buthidae C.L. Koch, 1837
Genus Leiurus Ehrenberg, 1828
Holotype : ♂ Saudi Arabia, Riyadh: Majami al-Hadb Protected Area, Wadi Rawdat al-hadb (NCWM/Sco-2023:1020), 23.V.2023, 21.605867°N, 43.766051°E, 1045 m a.s.l., Badry A. leg. (NCWM/Sco-2023: 1020). Paratypes: 1♀, Wadi Rawdat al-hadb (NCWM/Sco-2023: 1021), 22.V.2023, 21.650736°N, 43.704530°E, 985 m. a.s.l., Badry A. leg.; 6♀ & 3 juv., Wadi Rawdat al-hadb, 23.V.2023, 21.605867°N, 43.766051°E, 1045 m a.s.l., Badry A. leg. (NCWM/Sco-2023: 1022-29).
Leiurus arabicus: Saudi Arabia, Riyadh, 24.253561°N, 46.890831°E, 1♀, Alqahtani AR & Badry A leg. (NCWM/Sco-2023: 1111).
The specific name is placed in apposition to the generic name and refers to Majami al-Hadb protected Area, National Center for Wildlife, where the new species was found.
Medium to large Leiurus, 66.5–113.00 mm in length, carapace L 6.7–10.6 mm; base color is yellow or yellow-orange, carapace and tergites with extensive dark pigmentation; ventromedian carinae of metasomas II and III with some vestigial blackish spots over ventral carinae; metasoma IV fuscous except anteriorly; metasoma V heavily blackish; carapace with area between anterior median carinae bearing scattered fine to medium granules, area between posterior median carinae with deep median furrow of carapace moderately flanked by lateral granules by arcs; medial intercarinal surfaces of tergites II and III smooth or lightly shagreened; posterior margin of coxa III smooth or with sparse fine granules; metasoma moderately slender, metasoma II L/W 1.66–1.86, metasoma III L/W 1.81–2.07, metasoma IV L/W 2.02–2.36; ventromedian carinae of metasoma II and III with 23–33 denticles (16/16 carinae); metasoma V with enlarged subtriangular or lobate denticles on ventrolateral carinae; pedipalps slender, patella L/W ♂ 3.62, ♀ 3.08–3.74; leg III patella L/D ♂ 4.10, ♀ 4.16–5.55; pectin teeth ♂ 36–37, ♀ 29–32; pectines long, narrow, pectine L/carapace L ♂ 1.40, ♀ 1.03–1.33, mid-pectine sensillar margin L/metasoma I W ♂ 0.22, ♀ 0.107–0.163; pectin basal piece smooth in females, smooth or slightly shagreened in males; leg III basitarsus with 8–10 retrosuperior setae; pedipalp chela fixed finger with trichobothrium db distal to est; sternite VII with area between median carinae smooth or with sparse fine granulation anteriorly, more heavily in males; sternite carination: males, sternite III with median carinae weak to obsolete, sternites IV and V with weak, finely granulated lateral carinae, obsolete median carinae; females, sternite III with median carinae weak or obsolete, sternites IV and V with lateral carinae moderate, median carinae weak or obsolete.
(based on holotype and paratypes). Morphometric values presented in Tables
Morphometric values (in mm) of the female holotype and one male paratype of Leiurus hadb sp. nov. from Majami al-Hadb Protected Area.
Leiurus hadb sp. nov. | Leiurus hadb sp. nov. | |
---|---|---|
♂ holotype | ♀ paratype | |
Total length (including telson) | 85.5 | 92.00 |
Carapace: | ||
- length | 8.5 | 9.4 |
- anterior width | 6.1 | 6.4 |
- posterior width | 10.6 | 11.2 |
Mesosoma length | 24.2 | 22.4 |
Metasomal segment I: | ||
- length | 7.2 | 8.1 |
- width | 5.3 | 6.4 |
Metasomal segment II: | ||
- length | 7.9 | 9.0 |
- width | 4.6 | 4.3 |
Metasomal segment III: | ||
- length | 8.1 | 9.4 |
- width | 4.2 | 5.2 |
Metasomal segment IV: | ||
- length | 7.9 | 9.7 |
- width | 3.8 | 4.8 |
Metasomal segment V: | ||
- length | 10.0 | 11.7 |
- width | 3.6 | 4.3 |
- depth | 3.5 | 4.0 |
Telson length | 9.4 | 9.1 |
Vesicle: | ||
- length | 5.3 | 6.1 |
- width | 3.5 | 4.1 |
Pedipalp: | ||
- Femur length | 9.7 | 10.1 |
- Femur width | 2.4 | 2.9 |
- Patella length | 10.5 | 11.3 |
- Patella width | 2.9 | 3.4 |
- Chela length | 18.6 | 19.9 |
- Chela width | 2.7 | 3.4 |
- chela manus ventral | ||
- length | 5.7 | 6.3 |
Movable finger: | ||
- length | 13.2 | 14.0 |
Morphometric ratios of the ♂ holotype and ♀ paratype of Leiurus hadb sp. nov. from Saudi Arabia, including the ranges, mean ± SD, and sample sizes (in parentheses) for length (L) and width (W).
Ratio | Leiurus hadb sp. nov. | Leiurus hadb sp. nov. |
---|---|---|
♂ holotype (N = 1) | ♀ paratype (N = 7) | |
Carapace W/ L | 1.24 | 1.14–1.24 1.19 ± 0.038 |
Pedipalp femur L/W | 4.04 | 3.33–4.16 3.79 ± 0.30 |
Pedipalp patella L/W = b | 3.62 | 3.08–3.743 3.35 ± 0.23 |
Pedipalp chela L/manus W | 6.88 | 3.18–6.66 5.74 ± 1.16 |
Pedipalp movable finger L/ manus ventral L | 2.31 | 2.08–2.87 2.28 ± 0.26 |
Pedipalp movable finger L/ carapace L | 2.31 | 2.08–2.87 2.28 ± 0.26 |
Pedipalp chela manus W/ carapace L | 0.31 | 0.30–0.66 0.37 ± 0.13 |
Leg III patella L/D = c | 4.10 | 4.15–5.54 4.83 ± 0.57 |
Pectine L/ carapace L | 1.43 | 1.03–1.32 1.14 ± 0.10 |
Metasoma I L/W | 1.35 | 1.26–1.39 1.31 ± 0.05 |
Metasoma II L/W | 1.71 | 1.65 ± 1.85 1.74 ± 0.07 |
Metasoma III L/W = a | 1.92 | 1.80–2.06 1.89 ± 0.10 |
Metasoma IV L/W | 2.07 | 2.02–2.36 2.20 ± 0.10 |
Metasoma V L/W | 2.77 | 2.57–2.72 2.24 ± 0.05 |
Mid-pectine sensillar margin L/ metasoma I W | 0.22 | 0.10–0.16 0.13 ± 0.01 |
Fs = a.b.c | 28.66 | 23.57–37.61 25.73 ± 6.36 |
Coloration. Base color yellow or yellow-orange, carapace and tergites with extensive dark pigmentation; carapace dark on anterior interocular area, carinae, and posterior margin, light around posterior median furrow, lateral flanks outside lateral carinae pale; pretergites I–VI dark posteriorly, maculate anteriorly, III–VI with pair of pale median spots; tergites I–VI fuscous on medial and mediolateral intercarinal surfaces, with pair of pale anterior median spots, lateral flanks pale; tergite VII with slight fuscosity in anterior median area; ventromedian carinae of metasomas II and III with some vestigial blackish spots over ventral carinae; metasoma IV fuscous except anteriorly; metasoma V heavily blackish.
Prosoma
(Figs
Chelicera. Dorsal surface of manus smooth, with six short, pale microsetae, four near apical margin, two subapical, each surrounded by granules; dorso-internal carina at base of fixed finger very strong, well granulated, terminating anteriorly with prominent granules projecting over front of manus; single macroseta in middle of dorso-internal carina; dorsal surface of movable finger smooth, with four pale microsetae; fingers with characteristic buthid dentition (Vachon, 1963); movable finger dorsal margin with five teeth: dorsal distal tine, subdistal, median and two basal teeth fused in bicuspid; ventral margin with three teeth: ventral distal tine, median and basal teeth; fixed finger margin with four teeth: distal tine, subdistal, median and basal teeth; ventral aspect of fixed finger with two teeth.
Pectines
(Figs
Mesosoma
(Figs
Metasoma
(Figs
Pedipalps
(Fig.
Trichobothriotaxy
(Fig.
Legs
(Fig.
Most collections came from vegetated wadis in Majami al-Hadb Protected Area arid deserts. The species is probably lapidicolous, living under rocks in wadis of both sedimentary and igneous hills and mountains (Fig.
The Mujam’a Al-Hadab protected area is situated in the southwest of the Al-Hamra Mountains, approximately 80 km from the city of Rania in the Riyadh region. Covering 2256 square kilometers, it is dominated by dark volcanic mountains, sandy desert plains, and faded granite domes. These features give rise to wadis such as Wadi Sdiri, Wadi Al-Hamal, and Wadi Al-Farsha. The granite domes in this area range in color from pink to gray and have a smooth texture. Some of these domes rise ~ 400 meters above their surroundings. Desert air circulation has created cavities in the dome facades, along with small caves that fill with water during rainfall and persist for months after the rain stops. In addition to other scorpion species, Androctonus crassicauda (Olivier, 1807) and Compsobuthus manzonii (Borelli, 1915), can also be found in these mountains.
Leiurus hadb sp. nov. differs from Leiurus quinquestriatus and other Saudi Arabian species in the following characters:
The 16S rRNA data set analysis revealed that of 307 aligned nucleotides, 100 (32.57%) bases were constant, 199 (64.82%) bases were variable, and 80 (26.05%) were parsimony informative. The data set contained 119 polymorphic segregating sites within the 307 bp. The sequence divergences among Leiurus lineages ranged from 0.00 to 0.18, with an average of 0.15 (Table
The uncorrected p-distance of the sequence divergence of 16S mtDNA sequences between Leiurus samples was included in this study (standard error shown above the diagonal).
Species | 1 | 2 | 3 | 4 | 5 | 6 | 7 | 8 | 9 | 10 | 11 | 12 | 13 | 14 | 15 | 16 | 17 | 18 |
---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
1. L. hadb sp. nov. 1 | 0.00 | 0.00 | 0.00 | 0.00 | 0.02 | 0.02 | 0.01 | 0.01 | 0.02 | 0.02 | 0.02 | 0.02 | 0.02 | 0.02 | 0.02 | 0.02 | 0.03 | |
2. L. hadb sp. nov. 2 | 0.00 | 0.00 | 0.00 | 0.00 | 0.02 | 0.02 | 0.01 | 0.01 | 0.02 | 0.02 | 0.02 | 0.02 | 0.02 | 0.02 | 0.02 | 0.02 | 0.03 | |
3. L. hadb sp. nov. 3 | 0.00 | 0.00 | 0.00 | 0.00 | 0.02 | 0.02 | 0.01 | 0.01 | 0.02 | 0.02 | 0.02 | 0.02 | 0.02 | 0.02 | 0.02 | 0.02 | 0.03 | |
4. L. hadb sp. nov. 4 | 0.00 | 0.00 | 0.00 | 0.00 | 0.02 | 0.02 | 0.01 | 0.01 | 0.02 | 0.02 | 0.02 | 0.02 | 0.02 | 0.02 | 0.02 | 0.02 | 0.03 | |
5. L. hadb sp. nov. 5 | 0.00 | 0.00 | 0.00 | 0.00 | 0.02 | 0.02 | 0.01 | 0.01 | 0.02 | 0.02 | 0.02 | 0.02 | 0.02 | 0.02 | 0.02 | 0.02 | 0.03 | |
6. La1 L. arabicus KSA | 0.11 | 0.11 | 0.11 | 0.11 | 0.11 | 0.00 | 0.02 | 0.02 | 0.02 | 0.02 | 0.02 | 0.02 | 0.02 | 0.02 | 0.02 | 0.02 | 0.03 | |
7. La2 L. arabicus KSA | 0.11 | 0.11 | 0.11 | 0.11 | 0.11 | 0.00 | 0.02 | 0.02 | 0.02 | 0.02 | 0.02 | 0.02 | 0.02 | 0.02 | 0.02 | 0.02 | 0.03 | |
8. Lh1 L. haenggii KSA | 0.06 | 0.06 | 0.06 | 0.06 | 0.06 | 0.11 | 0.11 | 0.00 | 0.02 | 0.02 | 0.02 | 0.02 | 0.02 | 0.02 | 0.02 | 0.02 | 0.03 | |
9. Lh2 L. haenggii KSA | 0.06 | 0.06 | 0.06 | 0.06 | 0.06 | 0.11 | 0.11 | 0.00 | 0.02 | 0.02 | 0.02 | 0.02 | 0.02 | 0.02 | 0.02 | 0.02 | 0.03 | |
10. MT111845 L. quinquestriatus Egypt | 0.17 | 0.17 | 0.17 | 0.17 | 0.17 | 0.15 | 0.14 | 0.19 | 0.19 | 0.01 | 0.00 | 0.01 | 0.02 | 0.02 | 0.02 | 0.02 | 0.03 | |
11. MT111856 L. quinquestriatus Egypt | 0.16 | 0.16 | 0.16 | 0.16 | 0.16 | 0.16 | 0.16 | 0.18 | 0.18 | 0.02 | 0.01 | 0.00 | 0.02 | 0.02 | 0.02 | 0.02 | 0.03 | |
12. MT111862 L. quinquestriatus Egypt | 0.17 | 0.17 | 0.17 | 0.17 | 0.17 | 0.15 | 0.15 | 0.19 | 0.19 | 0.00 | 0.03 | 0.01 | 0.02 | 0.02 | 0.02 | 0.02 | 0.03 | |
13. MT111864 L. quinquestriatus Egypt | 0.17 | 0.16 | 0.17 | 0.16 | 0.16 | 0.16 | 0.15 | 0.18 | 0.18 | 0.02 | 0.00 | 0.02 | 0.02 | 0.02 | 0.02 | 0.02 | 0.03 | |
14. MT111865 L. quinquestriatus S. Saini Egypt | 0.18 | 0.17 | 0.18 | 0.17 | 0.17 | 0.14 | 0.15 | 0.18 | 0.18 | 0.10 | 0.09 | 0.11 | 0.09 | 0.00 | 0.02 | 0.02 | 0.03 | |
15. MT111866 L. quinquestriatus S. Saini Egypt | 0.18 | 0.17 | 0.18 | 0.17 | 0.17 | 0.14 | 0.15 | 0.18 | 0.18 | 0.10 | 0.09 | 0.11 | 0.09 | 0.00 | 0.02 | 0.02 | 0.03 | |
16. AY226174.2 L. macrocentrus Oman | 0.11 | 0.11 | 0.11 | 0.11 | 0.11 | 0.13 | 0.14 | 0.12 | 0.12 | 0.18 | 0.18 | 0.17 | 0.18 | 0.18 | 0.18 | 0.02 | 0.03 | |
17. KU318423.1 L. abdullahbayrami Turkey | 0.10 | 0.10 | 0.10 | 0.10 | 0.10 | 0.14 | 0.15 | 0.10 | 0.10 | 0.15 | 0.15 | 0.15 | 0.15 | 0.14 | 0.14 | 0.14 | 0.03 | |
18. AJ277598.1 Androctonus crassicauda (Outgroup) | 0.57 | 0.57 | 0.57 | 0.57 | 0.57 | 0.55 | 0.54 | 0.57 | 0.57 | 0.52 | 0.53 | 0.52 | 0.52 | 0.52 | 0.52 | 0.56 | 0.57 |
The phylogenetic analyses resulted in an identical topology between the maximum-parsimony and the neighbor-joining tree (Fig.
Table
1 | Medial intercarinal surfaces of tergites II–III smooth or sparsely, lightly shagreened | 2 |
– | Medial intercarinal surfaces of tergites II–III heavily or densely, finely shagreened | 4 |
2 | Females with pedipalp patella L/W < 3.20, Fs < 23 | L . haenggii |
– | Females with pedipalp patella L/W > 3.20, Fs > 23 | 3 |
3 | Metasomal segment IV fuscous except anteriorly; Smooth or weakly granulated median carinae on sternites III–V of females; lateral inframedian carinae represented in its posterior zone by 1/2 of the length on II and III; metasoma V with enlarged subtriangular or lobate denticles on ventrolateral carinae | L. hadb sp. nov. |
– | Metasomal segment IV yellow; Moderate to strongly granulated median carinae on sternites III–V of females; median lateral carinae restricted to posterior 0.28 of II, posterior 0.23 of III; metasoma V with enlarged, triangular, or subtriangular denticles on ventrolateral carinae | L . arabicus |
4 | Metasoma III ventromedian carinae with > 30 denticles; metasoma I–IV uniformly fuscous | L. jordanensis |
– | Metasoma III ventromedian carinae with < 30 denticles; metasoma I–IV yellow to light | L. brachycentrus |
Our results indicate morphological differences between L. hadb sp. nov., L. arabicus, and L. haenggii. L. hadb is closely related to L. arabicus, as they share slender leg, pedipalp, and metasomal segments (
Also, we conducted a molecular phylogenetic analysis using the mRNA 16S mitochondrial gene. Our analysis revealed a genetic divergence between L. hadb sp. nov. and samples of L. arabicus and L. haenggii from Saudi Arabia (p-distance = 0.06–0.012; Table
The authors extend their appreciation to the National Center for Wildlife in the Kingdom of Saudi Arabia for supporting this research work.
The authors have declared that no competing interests exist.
No ethical statement was reported.
National Center for Wildlife in the Kingdom of Saudi Arabia.
Conceptualization: AHAQ. Data curation: AB. Formal analysis: FM, AMAS, WSA, MSAB, AB. Investigation: AB, AHAQ, ARA. Methodology: FM, MSAB, AB, WSA, ARA, AHAQ, AMAS. Supervision: AHAQ. Visualization: AB. Writing - original draft: AB, ARA. Writing - review and editing: AHAQ, AB.
Abdulaziz R. Alqahtani https://orcid.org/0000-0003-2682-7965
Ahmed Badry https://orcid.org/0000-0002-2610-2019
All of the data that support the findings of this study are available in the main text.