Acanthodactylus pardalis ( Lichtenstein, 1823: 99 )

Bauer, Aaron M., DeBoer, Jonathan C. & Taylor, Dylan J., 2017, Atlas of the Reptiles of Libya, Proceedings of the California Academy of Sciences 64 (8), pp. 155-318 : 215-219

Acanthodactylus pardalis ( Lichtenstein, 1823: 99)

1823 L [acerta] Pardalis Lichtenstein, Verzeichniss der Doubletten des zoologischen Museums der Königl. Universität zu Berlin nebst Beschreibung vieler bisher unbekannter Arten von Säugethieren, Vögeln, Amphibien und Fischen. T. Trautwein, Berlin. x + 118 pp., 1 pl .

LECTOTYPE.— ZMB 1077, “Aegyptus”designated by Moravec et al. (1999). The paralectotypes are ZMB 1072–1076 and ZMB 55905 (formerly part of ZMB 1077 ) (Bauer and Günther 1995).

Acanthodactylus pardalis, Le Berre 1989:188 .

Acanthodactylus pardalis, Schleich, Kästle, and Kabisch 1996:388 .

Acanthodactylus pardalis, Sindaco and Jeremčenko 2008:218 .

DISTRIBUTION.— Mediterranean coast from the Gulf of Sirte east to the Nile Valley of Egypt (Sindaco and Jeremčenko 2008; Tamar et al. 2016b), a single record from east of the Nile is discussed by Moravec et al. (1999). In Libya they are found along the coast from the eastern Gulf of Sirte to the Egyptian border, but are absent from the elevated regions of the Jabal al Akhdar.

Libyan Records (Map 30): CYRENAICA: BENGHAZI: 342: MZUT R2289; Calabresi 1923; Zavattari 1929, 1930, 1934. 355: BMNH 1954.1.6. 5–7, Salvador 1982. 357: BMNH 1988.189, 1988.191–92; MZUF 852, 12688; NHMW 8161; von Martens 1883; Werner 1909; Ghigi 1920; Calabresi 1923; Umani 1923; Zavattari 1929, 1930, 1934; Moravec et al. 1999. 358: MZUF 853; Calabresi 1923; Zavattari 1929, 1930, 1934; Moravec et al. 1999. 364: Cornalia in Haimann 1882, 1886; Zavattari 1934. 367: MZUF 770; MZUT R2412; Calabresi 1923; Zavattari 1929, 1930, 1934; Moravec et al. 1999. 376: BMNH 1965.1259; Salvador 1982. MARJ: 385: MCSN 2079; Zavattari 1937. 399: Fonseca et al. 2008. 402: MZUT R2297; Calabresi 1923; Zavattari 1929, 1930. 403: NHMC 80.3.73.1. JABAL AL AKHDAR: 417: Zavattari 1929, 1930. 452: Ghigi 1920; Calabresi 1923; Zavattari 1929, 1930, 1934.

457cq: KNP 1983 /513[2 specimens]; Schleich

1987. DARNAH: 466: Zavattari 1922, 1929,

1930, 1934; Calabresi 1923. 472: NHMC

80.3.73.2; Fonseca et al. 2008. 474: MSNG

28431; MZUF 669; Vinceguerra 1927; Zavattari 1929, 1930, 1934. 475: BMNH 1965.1261;

MCZ R 46799–800; Salvador 1982. 480: ZSM

131/1983. 484: ZSM 132/1983. BUTNAN: 507:

BMNH 1965.1260; Salvador 1982. 512: ZSM

190/1979. AL WAHAT: 526: MSNG 31574*;

Vinciguerra 1931. 527: BMNH 1965.1265;

Salvador 1982; Moravec et al. 1999. 528:

BMNH 1965.1266; Salvador 1982; Moravec et al. 1999. 529: BMNH 1965.1262–64; MSNG

31574*; NMBA-REPT 15321; Vinciguerra

1931; Zavattari 1934; Scortecci 1935b; Salvador 1982; Moravec et al. 1999. 544: BMNH MAP 30. Distribution of Acanthodactylus pardalis in Libya.

1932.3.6. 7; MSNG 31574 *; Vinciguerra 1931 ;

Zavattari 1934. “ Cyrenaica ”: ZMB 10503. “ between Darnah and Marsa al Brega ”: BMNH 1965.1267; Salvador 1982. “ Marmarica ”: Zavattari 1937. “ Nordost-Libyen ”: ZMB 62964-67. “ deserto di Sahara ”: Rizzardi 1896. “ Sirtica ”: Zavattari 1937.

COMMENTS.— Some early citations to A. vulgaris (e.g., Cornalia from “Uadi Ahmar”) or A. lineomaculatus (von Martens from Benghazi) have been interpreted as referring to A. pardalis . However, the similarity of Acanthodactylus species to one another has probably resulted in the misidentifications of many specimens and unless confirmed by later specialists (e.g., Salvador 1982; Moravec et al. 1999) identifications provided in early citations are tentative at best. Records from Tripoli cited by Ghigi (1913), Werner (1909) and Zavattari (1934) are all derived from Rizzardi (1896), but this author actually reports the species (as Acantodactylus [sic] lineatomaculatus ) from the “deserto di Sahara.” Moravec et al. (1999) reviewed the A. pardalis group and described A. beershebensis for the taxon inhabiting the Negev, restricting true A. pardalis to North Africa. Tamar et al. (2016b) generated a multilocus phylogeny for the genus Acanthodactylus . Acanthodactylus pardalis was represented by Egyptian material only in their study and was embedded within A. maculatus , but basal support values within their entire “ pardalis group” are too low to infer meaningful support for this pattern. Salvador (1982) and Arnold (1983) recognized essentially the same Libyan taxa and boundaries as later molecular works, although Arnold (1983) included what are now A. bedriagai and A. busacki as subspecies within A. pardalis . Werner (1909) referred a juvenile specimen from Gherran (Garyān) to this species and this record was subsequently repeated by Ghigi (1913) and Zavattari (1934). Boulenger (1914, 1918, 1921) believed that this species extended to Tripoli and identified material from Al Khums and Misratah (e.g., MSNG 52517) as A. pardalis . These records, as well as the miscitations of Rizzardi (1896) noted above, are also the basis for Zavattari’s (1937) inclusion of the species in the Tripolitanian fauna. Most recently Essghaier et al. (2015) noted a record from Murzuq. We regard these as misidentifications referable to some congener and have not included them in the accompanying map.

IUCN THREAT STATUS.— Vulnerable A2c; B1ab(I,ii,iii).

Acanthodactylus scutellatus audouini Boulenger, 1918:154 (FIG. 23)

1918 Acanthodactylus scutellatus Var. Audouini Boulenger , Sur les lézards du genre Acanthodactylus Wiegm. Bulletin de la Société Zoologique de France 43:143–155.

LECTOTYPE.— BMNH 1920.1.20. 3006, “Wed El Kreil, Tunisia ” designated by Crochet et al. (2003), [ Tunisia]. Original type locality: “Egypte, Nubie, Tripoli, sud de la Tunisie.”

Acanthodactylus scutellatus [part], Le Berre 1989:190.

Acanthodactylus scutellatus, Schleich, Kästle, and Kabisch 1996:397 .

Acanthodactylus scutellatus audouini, Sindaco and Jeremčenko 2008:221 .

Acanthodactylus scutellatus audouini, Trape, Trape, and Chirio 2012:312 .

DISTRIBUTION.— Senegal,

Mauritania and Western across

Africa to Egypt (although absent from much of the Eastern Desert ,

Baha El Din 2006a) and Sudan, continuing through Sinai Israel, Jordan,

Iraq, Kuwait and far northwestern,

as well as eastern Saudi Arabia In

Libya distributed across the country.

Libyan Records (Map 31):

CYRENAICA: NUQAT AL KHAMS: 3:

NHMC 80.3.110.10–17. 5: BMNH

1955.1.8. 61–62, 1965.1130, 1965. FIGURE 23. Acanthodactylus scutellatus audouini from Awiss area , 1282; Salvador 1982. 5a: AIC (3 Tadrart Akakus , Ghat, Fezzan , Libya, 2008. Photo © Roberto Sindaco. specimens) no numbers provided; Ibrahim and

Ineich 2005. 7: Brito et al. 2008. ZAWIYAH: 12:

ZMB 39061. 17: BMNH 1955.1.8. 63; Salvador

1982. 16: BMNH 1982.372–76. 23: MZUT

R3262. JAFARA: 26: MZUT R3264. TRIPOLI:

35: BMNH 1954.1.6. 10–30, 1954.1.6. 45–60;

Salvador 1982. 36: BMNH 1955.1.6. 10–30,

45–60. 38: MCZ R 22348; USNM 148485. 39:

FMNH 82967, 82970; MCZ R 22350–51;

MNHN 1976.369–70; UMNH 4238. 43:

MZUT R3263; NHMW 31120. 45: BMNH

1955.1.8. 58–60; Crotchet et al. 2003; NHMW

8228; SMF 13648–51; ZMB 13781, 14031,

15304, 15307, 15322, 15329–30; CondorelliFrancaviglia 1896; König 1888; Werner 1909;

Ghigi 1913; Boulenger 1918, 1921; Zavatari

1934. 50: BMNH 1954.1.6. 43–44; Salvador

1982. 51: NMBA-REPT 15393. 53: BMNH MAP 31 . Distribution of Acanthodactylus scutellatus 1955.1.6. 43–44; MSNG 31589 *; NMBA- audouini in Libya .

REPT 15323. 56: BMNH 1955.1.8. 65, 1965.1283; Salvador 1982. MURQUB: 62: BMNH 1982.369–71. 64: FMNH 82924; ZMB 18015–17. 68: BMNH 1913.12.30. 6–10, 1954.1.6. 38–40, 1965.1129; CUP R 047–050; MSNG 28258, 52520, 52522; Boulenger 1914, 1921; Zavattari 1934;

Bons and Girot 1964; Salvador 1982; Frynta et al. 2000; Crochet et al. 2003. 69: MZUF 671. MISRATAH: 72: MCZ R 22349. 80: Boulenger 1914; Zavattari 1934. 81: Boulenger 1914. 87: NHMC 80.3.110.4. NALUT: 101: BMNH 1975.1208–09; Salvador 1982. 105: MNHN 2004.0089–91; Ibrahim and Ineich 2005. 108: FMNH 82936. 110: de Witte 1930. 114: MCCI R1468(1); Tamar et al. 2016. 116: MCCI R1465; Sindaco, pers. obs. 5/2/2008; Tamar et al. 2016. 118: NHMC 80.3.110.119–120. 119: BMNH 1954.1.6. 31–37; Salvador 1982. 124: MZUF 39095. JABAL AL GHARBI: 132: BMNH 1965.1279–81; Salvador 1982. 154: FMNH 82941. SIRTE: 174: ZMB 18012. 194: BMNH 1899.9.5. 6; Salvador 1982. 199: EPHE/CNHM 82965; Scortecci 1946; Bons and Girot 1964; Crochet et al. 2003. “ Tripolitania ”: FMNH 82924; FMNH 82925. “ Tripolitania settentrionale ”: Zavattari 1937. “ 14 km S El Habila ”: BMNH 1955.1.8. 64; Salvador 1982. “ Sirtica ”: Zavattari 1937. FEZZAN: WADI AL SHATII: 205: BMNH 1965.1285; Salvador 1982. 206: BMNH 1965.1286; Salvador 1982. 207: MCSN 2096; Scortecci 1946. JUFRA: 216: ZMB 18013– 14; Peters 1880, 1881; Werner 1909; Ghigi 1913; Zavattari 1934. 217: Zavattari 1937. 218: MCSN 2084; Scortecci 1946. 219: Scortecci 1946. 221: MCSN 2108. GHAT: 231: Scortecci 1946. 232: Scortecci 1946. 237: Sindaco, pers. obs. 4/20/2008. 239: MCSN 2090; Scortecci 1946. 242: MCCI- R1463–64; Frynta et al. 2000; Tamar et al. 2016b. 243: Scortecci 1946. 244: Scortecci 1946. 245: MCSN 2099; Scortecci 1935b. 248: MCSN 2097; Scortecci 1946. 249: MCSN 2092, 2101, 2115; Scortecci 1946. 253: MCSN 2089, 2100, 2105, 2109; Scortecci 1946. 255: Scortecci 1946. 256: MCSN 2098; Scortecci 1946. 258: CUP R 013, 014; Frynta et al. 2000. 260: Sindaco, pers. obs. 4/22/2008. WADI AL HAYAA: 267: CUP R 028; Frynta et al. 2000. 269: Scortecci 1946; Essghaier et al. 2015. 274: CUP R 026–27; Frynta et al. 2000. SABHA: 280: MCSN 2104. 281: ZCT 2006.55; Ibrahim 2008a; Werner and Ibrahim 2012. 286: BMNH 1954.1.6. 41; MSNG 50657; NMBA-REPT 17808, 17811; Salvador 1982. MURZUQ: 293: Sindaco, pers. obs. 4/23/2008. 294: Sindaco, pers. obs. 4/23/2008. 295: Sindaco, pers. obs. 4/23/2008. 298: BMNH 1945.11.9. 8–9, 1956.1.1. 8; MCSN 2103; ZCT 2005.41; Scortecci 1946; Salvador 1982; Ibrahim 2008a; Werner and Ibrahim 2012; Essghaier et al. 2015. 303: Essghaier et al. 2015. 325: MCSN 2088. Fezzan: “ Surual (Tasili) ”: Scortecci 1946. “ Between Erg Ouan Kasa and Erg Murzuq ”: MCCI R1466–67; Tamar et al. 2016b. CYRENAICA: BENGHAZI: 357: BMNH 1965.1278; Umani 1923; Zavattari 1930, 1934; Salvador 1982. 358: MCSN 2093. 371: CUP R 024, 029, 033, 035; Frynta et al. 2000. MARJ: 385: MCSN 2091; Zavattari 1937. JABAL AL AKHDAR: 448: SMF 55409. DARNAH: 475: FMNH 82926. 479: MCSN 2086; Scortecci 1935b. BUTNAN: 520: Vinciguerra 1931; Zavattari 1934. 521: MSNG 28434, 37520; Vinciguerra 1927; Zavattari 1929, 1930, 1934, 1937. 524: Vinciguerra 1927; Zavattari 1929, 1930, 1934. AL WAHAT: 529: BMNH 1955.1.1. 46–47; MCSN 2110–11; NMP 34933; MSNG 31588*; NMP 34933; Salvador 1982; Moravec 1995. 538: NMBA-REPT 15322. 540: BMNH 1965.1268–73, 1965.1277; Salvador 1982. 541: NHMC 80.3.110.18. 544: BMNH 1965.1274–76; Vinciguerra 1931; Zavattari 1934; Salvador 1982. 553: Zavattari 1937. 554: MCSN 1841. 564: BMNH 1932.3.6. 9–10; MSNG 31588*; Zavattari 1929, 1930, 1934, 1937; Vinciguerra 1931; Salvador 1982. KUFRAH: 571: MZUF 771; Scortecci 1935c; Zavattari 1937. 571a: MSNG 31589*; Vinciguerra 1931. 572: BMNH 1932.3.6. 8; Vinciguerra 1931; Salvador 1982. 573: Vinciguerra 1931; Zavattari 1937. 575: BMNH 1932.3.6. 12; BMNH 1975.1229–36; BMNH 1982.381–82; MSNG 31589*; ZMB 18010–11; Peters 1880, 1881; Werner 1909; Ghigi 1913; Zavattari 1929, 1930, 1934; Salvador 1982. 575a: MSNG 31589*; Vinciguerra 1931. 577: MSNG 31589*; Vinciguerra 1931. “ Kufrah: 1000 km South of Benghazi ”: BMNH 1987.1030–32. “ Cyrenaica ”: MCSN 2094; MZUF 772; MSNG 52499. “Between Agedabia and Gialo”: MSNG 31588*–89*; Vinceguerra 1931. LIBYA: RMNH RENA 13917. “ Libysche Wüste ”: ZMB 27456. “dunes south of Ettoni ”: MZUT R3266. [* = two different specimens with apparently the same number but different localities]

COMMENTS.— Acanthodactylus species have proved particularly difficult for many authors to identify (Vinceguerra 1927, 1931; Zavattari 1930; Scortecci 1946) and the identity of A. scutellatus and its relatives has been considered in detail by Scortecci (1946) and Bons and Girot (1964). The latter authors plotted localities across Libya for A. s. audouini, which they considered as clearly distinct form the nonminotypical form occurring in Egypt and Sudan. Arnold (1983) considered the names audouini, inornatus and dumerilii as conspecific with scutellatus and declined to try to identify subspecific divisions among them, whereas Salvador (1982) separated dumerilii (see account above) as a different species. Three subspecies have typically been recognized: A. s. scutellatus ( Egypt, Sinai, and southern Israel), A. s. audouini Boulenger, 1918 (Saharan), and A. s. hardyi Haas, 1957 ( Iraq, Saudi Arabia, Kuwait) ( Schleich et al. 1996; Sindaco and Jeremčenko 2008). Crochet et al. (2003) recently revised the A. scutellatus group and recognized six constituent taxa. Tamar et al. (2016b) found deep divergences between the reciprocally monophyletic A. s. scutellatus and A. s. audouini, with robust substructure in the latter. Libyan samples fall out in two clades, one allied to Tunisian populations and the other Egyptian + Mauritanian samples. The degree of divergence from the nominate form raises the question if specific status is warranted for A. s. audouini. Brygoo (1988) claimed that Boulenger had designated a specimen figured by Audouin in the Déscription de l’Egypte as the type of this taxon, although this does not appear to be the case, although this specimen may be considered as part of the original syntype series, which also includes various specimens in the BMNH collection.

IUCN THREAT STATUS.— Not assessed, but assumed to be Least Concern.