Corresponding author: Jiří Moravec (
Academic editor: Pavel Stoev
A recent molecular phylogeny of the Arid clade of the genus
The genus
Phylogeny of the
The discovery of a monophyletic species group consisting of one recently described and two newly recognized species calls upon a more thorough study of the nomenclatural status, evolutionary relationships, taxonomy and distribution of its members based on further genetic and morphological data. The present study focuses on this task.
In order to resolve the phylogenetic relationships between the two newly recognized
List of material used for the phylogenetic analyses. Holotype of
Species | Code | Museum number | Country | Locality | Loc. No | Lat, Long |
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Sher10660 | SMB 10660 | Egypt | Ayoun Musa | 1 |
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Hd41 | NMP6V70163/2 | Egypt | Sharm el Sheik; Sinai | 2 |
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Hd96 | NMP6V70163/1 | Egypt | Sharm el Sheik; Sinai | 2 |
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- | - | - |
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Hd97 | NMP6V70163/3 | Egypt | Sharm el Sheik; Sinai | 2 |
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- | - | - |
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HSA63 | ZFMK 94084 | Saudi Arabia | Al Wajh | 3 |
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HSA64 | ZFMK 94085 | Saudi Arabia | Al Wajh | 3 |
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HSA65 | ZFMK 94086 | Saudi Arabia | 15 km S of Al Wajh | 4 |
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HSA66 | ZFMK 94087 | Saudi Arabia | 15 km S of Al Wajh | 4 |
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HSA67 | ZFMK 94088 | Saudi Arabia | 15 km S of Al Wajh | 4 |
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HSA68 | TUZC-R8 | Saudi Arabia | 15 km S of Al Wajh | 4 |
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HSA69 | ZFMK 94089 | Saudi Arabia | 15 km S of Al Wajh | 4 |
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HSA70 | TUZC-R9 | Saudi Arabia | 72 km N of Umluj | 5 |
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HSA62 | TUZC-R10 | Saudi Arabia | 180 km W of Hail | 6 |
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HSA61 | IBES10001 | Saudi Arabia | Al Ghat | 7 |
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HSA57 | IBES10183 | Saudi Arabia | 30 km NE of Alhawiyah | 8 |
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HSA58 | ZFMK 94090 | Saudi Arabia | 30 km NE of Alhawiyah | 8 |
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HSA59 | TUZC-R11 | Saudi Arabia | 30 km NE of Alhawiyah | 8 |
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HSA60 | IBES10344 | Saudi Arabia | 30 km NE of Alhawiyah | 8 |
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HSA54 | IBES10150 | Saudi Arabia | 20 km S of Ashayrah | 9 |
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HSA55 | ZFMK 94091 | Saudi Arabia | 20 km S of Ashayrah | 9 |
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HSA56 | IBES10363 | Saudi Arabia | 20 km S of Ashayrah | 9 |
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ZFMK 87236 | ZFMK 87236 | Saudi Arabia | Taif National Wildlife Research Center | 10 |
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BJ27 | NHM-BS N41914 | Yemen | Marib | 17 |
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BJ28 | NHM-BS N41913 | Yemen | Marib | 17 |
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17 |
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JS48 | NMP6V 74834/1 | Yemen | Wadi Zabid | 11 |
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JS49 | NMP6V 74834/2 | Yemen | Wadi Zabid | 11 |
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JS45 | not collected | Yemen | Al Hababi | 12 |
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JS46 | NMP6V 74833/1 | Yemen | Al Hababi | 12 |
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12 |
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JS37 | NMP6V 74832/1 | Yemen | 3 km S of Najd an Nashamah | 13 |
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JS38 | NMP6V 74832/2 | Yemen | 3 km S of Najd an Nashamah | 13 |
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JS32 | NMP6V 74835 | Yemen | 35 km W of Lahij | 14 |
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BJ09 | NHM-BS N41916 | Yemen | Radman | 15 |
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JS17 | NMP6V 74831/1 | Yemen | Al Hadr | 16 |
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JS18 | NMP6V 74831/2 | Yemen | Al Hadr | 16 |
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JS123 | NMP6V 74845/2 | Ethiopia | Arba Minch | - |
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JS111 | not collected | Pakistan | Okara | - |
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JS113 | not collected | India | Haridwar | - |
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JS119 | not collected | Oman | Jalan Bani Bu Hassan | - |
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The final dataset consisted of 36 ingroup individuals. Specimen numbers, localities, and GenBank accession numbers of all genes sequenced are presented in maximum likelihood Bayesian inference
Maximum likelihood analyses of both datasets were performed in RAxML 7.0.3 (
The BI analyses were run in MrBayes 3.2.1 (
Heterozygous positions in nuclear genes were identified based on the presence of double peaks in chromatograms and using the Heterozygote Plugin in Geneious. For the purpose of haplotype network construction, haplotypes from sequences with more than one heterozygous position were resolved in PHASE 2.1.1 (
Material for morphological comparison included 225 specimens of 8 National Museum Prague, Czech Republic Natural History Museum in Braunschweig, Germany Senckenberg Forschungsinstitut und Naturmuseum, Frankfurt, Germany Zoologisches Forschungsmuseum Alexander Koenig, Bonn, Germany Museo Civico di Storia Naturale “Giacomo Doria”, Genova, Italy Museo Civico di Storia Naturale di Milano, Milano, Italy Museo Civico di Storia Naturale, Carmagnola, Italy Università di Firenze, Museo Zoologico “La Specola”, Firenze, Italy British Museum of Natural History, London, UK California Academy of Sciences, San Francisco, USA Taif University Zoological Collection, Taif, Saudi Arabia Institute of Evolutionary Biology Collection, Barcelona, Spain Tomas Mazuch private collection, Dříteč, Czech Republic L. Kratochvíl collection J. Šmíd collection Sherif Baha El Din private collection, Cairo, Egypt
The following measurements were taken with Powerfix digital calliper to the nearest 0.1 mm: snout-vent length head length head width head depth left eye diameter axilla-groin distance tail length
Phylogenetic analyses of both datasets resulted in trees presented in
Maximum likelihood trees of mtDNA and mtDNA + nDNA datasets of the ‘
Nuclear allele networks of the four loci analyzed (
The results of the molecular analyses, together with a unique combination of morphological features (see below) confirm the earlier conclusion that the newly recognized
SMF 8723 (lectotype, adult male), Petr. Arabica [Arabia petraea], collected by E. Rüppell in 1827 (MorphoBank
SMB 10660, Egypt, Suez governorate, Ayoun Musa (
Recent examination (by JŠ) of four specimens collected by Rüppell (SMF 8723–8726) has shown that one of them [SMF 8723 designated by
SMF 8723, adult male [erroneously determined as female by
Male lectotype of
Schematic drawing of the chin region of the lectotype and a new specimen from Sinai of
Measurements (in mm): SVL 51.5, HL 12.9, HW 9.8, HD 6.0, E 3.3, AG 23.7.
Paralectotype SMF 8724 differs from other individuals of
Morphological comparison among members of the ‘
Species / Character |
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Upper labials | 18 | 9.4 ± 0.5 | 3 | 9.3 ± 0.8 | 10 | 9.3 ± 0.8 | 27 | 9.4 ± 0.7 | 33 | 8.2 ± 0.5 | 5 | 10.8 ± 0.8 | 18 | 9.8 ± 0.7 | 51 | 10.3 ± 0.7 | 57 | 10.2 ± 0.7 |
9–11 | 8–10 | 8–10 | 8–11 | 7 - 10 | 10 - 12 | 8–12 | 9–12 | 8–12 | ||||||||||
Lower labials | 18 | 7.4 ± 0.4 | 3 | 7.7 ± 0.6 | 10 | 8.0 ± 0.6 | 27 | 7.7 ± 0.6 | 33 | 6.7 ± 0.5 | 5 | 8.1 ± 0.4 | 18 | 8.2 ± 0.6 | 51 | 7.9 ± 0.5 | 57 | 7.8 ± 0.6 |
7–9 | 7–8 | 7–9 | 6–9 | 6–8 | 7–9 | 7–10 | 6–9 | 6–10 | ||||||||||
Nasals in contact (%) | 18 | 11 | 3 | 33.3 | 10 | 40 | 27 | 22.2 | 33 | 21.2 | 5 | 0 | 18 | 5.5 | 51 | 7.8 | 57 | 5.3 |
1st postmental in contact with 2nd lower labial (%) | 18 | 100 | 3 | 33.3 | 10 | 100 | 27 | 70.3 | 33 | 12.1 | 5 | 80 | 18 | 83.3 | 51 | 98 | 57 | 89.5 |
Rows of dorsal tubercles | 18 | 14.1 ± 0.2 | 3 | 14 ± 0.0 | 10 | 14.1 ± 1.0 | 27 | 14.8 ± 1.2 | 33 | 13.8 ± 0.7 | 5 | 12.4 ± 0.9 | 15 | 14 ± 1.4 | 46 | 15.3 ± 1.1 | 53 | 15.2 ± 1.2 |
14–15 | 14–14 | 12–16 | 13–18 | 12–16 | 12–14 | 12–16 | 13–18 | 12–18 | ||||||||||
Pores | 8 | 5.6 ± 1.1 | 1 | 6 | 2 | 8 ± 0.0 | 9 | 6.1 ± 0.8 | 13 | 7.2 ± 1.4 | 1 | 4 | 9 | 7.2 ± 1.1 | 23 | 13.7 ± 2.2 | 27 | 11.2 ± 1.1 |
4–7 | 8 - 8 | 5–8 | 6–10 | 6–9 | 10–18 | 9–13 | ||||||||||||
Lamellae under 1st toe | 18 | 7.4 ± 0.5 | 3 | 8.2 ± 0.3 | 10 | 5.4 ± 0.5 | 27 | 6.1 ± 0.5 | 32 | 6.5 ± 0.5 | 5 | 6.2 ± 0.3 | 18 | 6.9 ± 0.7 | 51 | 6.7 ± 0.4 | 57 | 6.3 ± 0.4 |
7–8 | 8–9 | 5–6 | 5–8 | 6–7 | 6–7 | 6–8 | 6–8 | 5–7 | ||||||||||
Lamellae under 4th toe | 18 | 11.5 ± 0.7 | 3 | 11.2 ± 0.3 | 10 | 8.6 ± 0.5 | 27 | 10.1 ± 0.7 | 32 | 9.7 ± 0.6 | 5 | 10 ± 0.0 | 18 | 10.9 ± 0.8 | 51 | 10.4 ± 0.6 | 57 | 10.2 ± 0.5 |
10 - 13 | 11–12 | 8–9 | 8–12 | 8–11 | 10–10 | 9–12 | 9–12 | 9–11 | ||||||||||
SVL (males) | 8 | 46.8 ± 5.9 | 1 | 58.3 | 2 | 38.6 ± 2.6 | 8 | 41.8 ± 2.3 | 13 | 46.0 ± 5.8 | 1 | 49.3 | 8 | 48.4 ± 4.1 | 23 | 58.5 ± 7.1 | 25 | 56.5 ± 5.7 |
39.0–53.2 | 36.8–40.4 | 37.0–43.7 | 37.3–54.1 | 40.0–54.2 | 43.6–74.9 | 45.2–65.3 | ||||||||||||
SVL (females) | 10 | 49.0 ± 3.5 | 2 | 53.5 ± 7.9 | 2 | 40.1 ± 0.9 | 16 | 43.6 ± 4.7 | 18 | 49.2 ± 5.1 | 4 | 46.2 ± 11.4 | 8 | 48.6 ± 3.3 | 23 | 55.7 ± 5.3 | 30 | 52.6 ± 5.1 |
40.6–53.3 | 47.9–59.1 | 39.4–40.7 | 32.7–50.1 | 39.4–56.2 | 35.6–56.6 | 43.1–54.0 | 43.6–62.1 | 42.4–64.1 |
From
From
From
From
From
From
Specimens with intact tail vary in number of tail segments bearing 6 pointed tubercles (7–8). The original portion of the tail of the female NMP6V 70163/4 is very wide at the base, separated from cloacal region by a basal constriction. One specimen (IBES10212) is the only animal with 15 longitudinal rows of enlarged tubercles. Another one (IBES10284) has uppermost nasals in wide contact. Most striking is the variation in the number of preanal pores in males. Whereas the lectotype and the only male from Sinai (NMP6V 70163/2) have both 4 pores, all males from Saudi Arabia have 6–7 pores. There seems to be clinal variability in this character, males from NW of the known range (
Distribution map of
Coloration (in life) pale buff dorsally (
Live specimens of
There is a very low variation in mtDNA between specimens from Sinai and Saudi Arabia (max. 1.3% in both
Eduard Rüppell collected the original series in 1827 when he began his marine biological studies of the Red Sea and travelled from Egypt to Eritrea. There is no specific information that he went to Arabia as well (
In 1996, when the NMP specimens were collected, the locality in Sharm el-Sheikh was formed by a crop field supplied with drain water from nearby habitations. Geckos were found during the day under unused empty barrels and also inside buildings. Other species syntopic with
NMP6V 74833/2,adult male (MorphoBank
NMP6V 74833/1 (adult male, MorphoBank
NMP6V 74835 (juvenile), Yemen, Lahij governorate, wadi 35 km W of Lahij (
A small species of the ‘
From
From
From
From
From
From
From
From
NMP6V 74833/2, adult male. Body slightly depressed to cylindrical (
Holotype of
Measurements (in mm): SVL 40.4, HL 11.5, HW 8.6, HD 5.2, E 2.8, AG 16.2.
Overall dorsal coloration brownish grey. An indistinct dark horizontal stripe in loreal and temporal area. Seven dark brown transverse bands across the nape and body, the one in scapular region being the most conspicuous. Dark brown bands also on the original part of the tail. Belly whitish.
The paratypes (
Four (out of eight) paratypes of
Measurements of paratypes (in mm): NMP6V 74831/1: SVL 40.7, HL 11.5, HW 8.2, HD 4.9, E 3.0, AG 19.0; NMP6V 74831/2: SVL 32.0, HL 9.3, HW 6.6, HD 3.7, E 2.1, AG 12.7; NMP6V 74832/1: SVL 32.7, HL 9.7, HW 7.0, HD 3.4, E 2.3, AG 14.3; NMP6V 74832/2: SVL 32.9, HL 9.3, HW 6.7, HD 3.6, E 2.4, AG 13.5; NMP6V 74833/1: SVL 36.8, HL 10.7, HW 8.0, HD 4.5, E 2.4, AG 14.1, TL 42.5; NMP6V 74834/1: SVL 39.4, HL 11.1, HW 8.1, HD 4.4, E 2.7, AG 16.7; NMP6V 74834/2: SVL 32.0, HL 9.5, HW 6.7, HD 3.9, E 2.5, AG 13.8; NHM-BS N41916: juvenile, not measured.
As already mentioned (Results), the level of genetic variability within
The species epithet “
The following reptile specieswere found to occur in sympatry with
Previous phylogenetic studies of the Arid clade of
The highlands of southwestern Saudi Arabia and Yemen are known to host a high number of endemic taxa (
We thank the following curators for granting access to collections under their care: U. Joger (NHM-BS), G. Köhler and his assistant L. Acker (SMF), R. Sindaco and G. Boano (MCCI), G. Doria (MSNG), S. Scali (MSNM), A. Nistri (MZUF), J. Vindum (CAS), B. Clarke and E. N. Arnold (BMNH), and T. Mazuch. We are very indebted to R. Kovář and R. Víta for collecting the Sinai material of