Eremias rafiqi, Masroor & Khan & Nadeem & Amir & Khisroon & Jablonski, 2022
Masroor, Rafaqat, Khan, Muazzam Ali, Nadeem, Muhammad Sajid, Amir, Shabir Ali, Khisroon, Muhammad & Jablonski, Daniel, 2022, Appearances often deceive in racerunners: integrative approach reveals two new species of Eremias (Squamata: Lacertidae) from Pakistan, Zootaxa 5175 (1), pp. 55-87 : 70-75
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Eremias rafiqi sp. nov.
Suggested vernacular name: Rafiq’s Racerunner
Holotype. PMNH 856 View Materials (cyt b: n/a; Rag1: n/a), an adult male, collected from Tanishpa village, Torghar Mountains, Killa Saifullah district , Balochistan (31.1869º N, 68.4126º E; Fig. 1 View FIGURE 1 ), elevation 2,506 m a. s. l., May 25, 1997, leg. Khalid Javed Baig ( Fig. 4 View FIGURE 4 ). GoogleMaps
Paratypes. Males: PMNH 855 View Materials (cyt b: n/a; Rag 1: n/a), 857 (cyt b: n/a; Rag 1: n/a) , PMNH 859 View Materials (cyt b: n/a; Rag 1: n/a) , PMNH 861 View Materials (cyt b: n/a; Rag 1: n/a) , PMNH 4056 View Materials (cyt b: n/a; Rag 1: MT 554487 View Materials ). Females : PMNH 3724 View Materials (cyt b: n/a; Rag 1: MT 554476 View Materials ) , PMNH 3735 View Materials (cyt b: MT 554461 View Materials ; Rag 1: MT 554477 View Materials ) , PMNH 4058 View Materials (cyt b: n/a; Rag 1: n/a). Juveniles : PMNH 837 View Materials (cyt b: n/a; Rag 1: n/a) , PMNH 3723 View Materials (cyt b: MT 554470 View Materials ; Rag 1: MT 554496 View Materials ) , PMNH 4053 View Materials (cyt b: MT 554457 View Materials ; Rag 1: MT 554485 View Materials ) , PMNH 4054 View Materials (cyt b: MT 554454 View Materials ; Rag 1: MT 554480 View Materials ) . PMNH 857 View Materials , collected along with the holotype ; PMNH 855 View Materials and 861, May 26, 1997, Ashewat, Qamar Din Karez, Zhob district (31.3448ºN, 68.6307ºE), leg. Khalid Javed Baig GoogleMaps ; PMNH 859 View Materials , May 24,1997, Tanishpa village, Torghar, Killa Saifullah district , leg. Khalid Javed Baig ; PMNH 3723–24 View Materials , October 09, 2017, Khar, Nushki district , Balochistan (29.5879ºN, 65.6609ºE), leg. Muazzam Ali Khan GoogleMaps ; PMNH 3735 View Materials , October 21, 2017, Khar, Nushki district , leg. Muazzam Ali Khan ; PMNH 4053 View Materials , September 05, 2018, Zamkai Nala, Tanishpa village, Torghar , Killa Saifullah district (31.1930ºN, 68.4111ºE), leg. Rafaqat Masroor GoogleMaps ; PMNH 4054 View Materials , September 01, 2018, Ashewat, Qamar Din Karez, Zhob district , leg. Rafaqat Masroor ; PMNH 4056 View Materials and 4058, August 30, 2018, Kunder, Torghar, Killa Saifullah district , leg. Ibad ur Rehman ( Figs. 5 View FIGURE 5 & 6 View FIGURE 6 ) .
Morphological diagnosis. A large-sized lacertid lizard, maximum snout-vent length ( SVL) = 99.3 mm, tail 1.67 to 1.89 times longer than body length ( SVL), hindlimbs relatively long ( HLL / SVL ratio 0.6–0.8); subocular scale reaching to the edge of the mouth, 5–7 (mainly 6, rarely 5) anterior to subocular; dorsals 56–67; ventrals in 14–17 oblique longitudinal series; frontal separated from supraoculars; the height of the first two to three transverse rows of ventral scales in the pectoral region more than its breadth; 17–21 femoral pores on each side, separated medially by 1–4 scales (mainly 3, rarely 1), the space between the femoral pores less than one–fourth length of each row; toes without fringe, encircled by three scales in a single series of 22–27 unicarinate and bicarinate scales underneath; the tip of the fourth toe reaches to the forelimb and extends to just behind the collar. The adult specimens are grayish in life with four series of longitudinal black ocelli on the dorsum originating from behind the parietals and extending onto the tail; on each lateral side, a broader dark stripe originates from behind the eye and continues onto the tail with disconnected white round ocelli at the margins as well as white ocelli inside the stripe.
Molecular data. Eremias rafiqi sp. nov. represents the so-called Zabol clade sensu Rastegar-Pouyani et al. (2010) from eastern Iran and the clade E identified by Khan et al. (2021) from northeastern Balochistan, Pakistan. Whereas Rastegar-Pouyani et al. (2010) identified this clade solely on mtDNA (cyt b, 12S), Khan et al. (2021) used mitochondrial (16S, COI, cyt b) as well as nuclear data (Rag 1) that clearly showed deep differentiation of this new species from other available sequences of the genus. The distinction of Eremias rafiqi sp. nov. is supported by its phylogenetic position (monophyletic clade among other species of E. persica complex, sister to E. fahimii but with weak statistical support; Fig. 1 View FIGURE 1 ), the differentiation on solely Rag1 dataset ( Fig. 2 View FIGURE 2 ), and the value of the uncorrected p distances reaching from 8.5% ( E. fahimii ) to 20.7% ( E. strauchi ) ( Table 3 View TABLE 3 ). The intraclade genetic diversity (cyt b) is 2 % with six detected haplotypes found in Iran and Pakistan ( Fig. 2 View FIGURE 2 , Table 3 View TABLE 3 ).
Etymology. The species epithet “ rafiqi ” is taken from the first name of late Rafique Ahmed Rajput (1968– 2008) to whom the new species is dedicated. The deceased Rajput served in Sindh Wildlife Department from 1986 till his demise. With no formal education in wildlife research, conservation and management, his passion for the conservation of wildlife in Pakistan remain unparalleled. From collecting the first-ever data of Ursus arctos isabellinus in the high-altitude Deosai Plateau, Gilgit-Baltistan, to the faunistic studies in the Indus River Delta and desert areas, he was a symbol of hard work. He also has very sound techniques for the collection of lizards and snakes. Unfortunately, during one such endeavor for gathering faunistic data, he collected a juvenile venomous krait, Bungarus sp. (possibly B. persicus ) from Jiwani town, Gwadar District, Balochistan and mistaken its identity with non-venomous Lycodon species. On the morning of October 13, 2008, when he was shifting the live snake from a container to permanently preserve it for research purposes, the snake bit him multiple times.After a couple of hours, Rajput felt severe pain, anxiety and dizziness. He was immediately taken to the hospital in Karachi, where the doctors told his relatives that anti-snake venom is not available and asked them to get it from the pharmacy market. By the time the anti-venom was arranged, Rajput breathed his last.
Description of the holotype. SVL: 99.3, TL: 168.0, HL: 26.9, HW: 14.5, HH: 15.5, TrL: 42.5, HLL: 66.8, FLL: 37.3, FrL: 6.3, FrW: 3.4.An adult male preserved in formalin in a good state of preservation ( Fig. 4 View FIGURE 4 ); head and body moderately depressed; tail long, ca. 1.7 times longer than the body, cylindrical and depressed at the base. Head relativelylong (HL/SVL, 0.27) ( Fig. 4 View FIGURE 4 ), ca. 1.7 times longer than wide (HW/HL, 0.58), head height slightly less than head width (HH/HW, 0.89). Limbs strong, hindlimbs ca. 1.8 times longer than the length of forelimbs (FLL/HLL, 0.56), hindlimbs comprise 1.4 times of the body length (HLL/SVL, 0.67).
Head slightly broader than the neck.Head shields including nasals,frontonasal,prefrontals,frontal,frontoparietals, interparietal and parietals are smooth and convex. Nasals moderately swollen, three nasals, the lower in contact with two supralabials on both right and left side and in contact with the rostral ( Fig. 4D View FIGURE 4 ). Supranasals in contact with rostral but lack such contact with first supralabial, the suture between them is 3.7 times the length of frontonasal, whose breadth is slightly more than its length; length of prefrontals 1.6 times its width, forming a median suture; length of frontal ca. 1.8 times as long as broad, its length slightly less than its distance from the tip of the snout, narrow behind; parietals smooth, slightly longer than its width; interparietal smooth, more than half the length of frontoparietals, about equal to the suture of frontoparietal; no occipital. Two large supraoculars, about equal in size, the space anterior to supraoculars filled by few small and three to five larger granules; both supraoculars in contact with frontal of their sides while separated from supraciliaries by a series of granules ( Fig. 4C View FIGURE 4 ); six supraciliaries, first longest, its length shorter than its distance from the first loreal. Rostral pentagonal, broader than high, narrower beneath than above; anterior loreal slightly higher than wide, shorter than the second loreal which is longer than high; supralabials 10 on the right side, 9 on the left side; subocular keeled just below the eye, bordering the mouth, wedged between sixth and seventh supralabials on the right side and fifth and sixth supralabials on the left side ( Fig. 4D View FIGURE 4 ). Temporals smooth, a large scale above ear; auricular denticulation indistinct or three small scales forming slight denticulation anteriorly. Lower eyelid covered with numerous small semi-transparent scales.
Six infralabials on the right side, seven on the left side, gradually increasing in size posteriorly. Five pairs of chin shields; anterior three completely in contact, the fourth pair separated by 10 to 11 smaller gulars in a straight line, fifth not in contact with infralabials, separated by a single row of scales. Collar curved, free, serrated and composed of ten plates larger than adjacent gulars, the middle one quite enlarged than others. Gular fold distinct, 32 gular scales in a straight line between the symphysis of the chin shields and the collar ( Fig. 4B View FIGURE 4 ).
Dorsal scales granular, smooth, 62 across the middle of the body. Ventral plates broader than long (except for outermost series), forming oblique longitudinal series of 16 plates across mid-belly and 29 transverse rows counted from behind collar to vent; first three rows of ventral scales in the pectoral region behind collar longer than broad, the first row is twice as long as broad. Precloacal region with a pair of the enlarged median plates just above the vent, surrounded by six large scales.
Forelimb ca. 1.4 times longer than the head, the upper surface of the arm with rhombic, smooth scales. Scales on the upper surface of hindlimbs similar to dorsals, equal in size; ventral surface of hindlimbs covered by enlarged plates, the ventral surface of the tibia with one row of very large and one comparatively smaller plates; the tip of the fourth toe reaches to the forelimb and extends to just behind the collar; 21 femoral pores on the right side, most of the left side damaged, the two femoral pore series separated by two scales, length of the interfemoral space not greater than one–fourth length of each row. Toes slender, compressed, with no fringe; subdigital lamellae unicarinate, in a single row of 25 scales under the 4th toe, a total of three scales around the 4 th toe. Upper caudal scales oblique, truncate, strongly and diagonally keeled, 30 scales in the 9 th –10 th annulus behind the postcloacal granules.
Coloration in life. The adult specimens are grayish in life with four more or less regular rows of black spots on the light dorsum, originating from behind the parietals, smaller on the nape, larger on the middle of the dorsum, disappearing on proximal one-fourth of the tail. The middle two rows of black spots have white spots along each black spot of the rows. On each lateral side, a broader dorsolateral dark stripe originates from behind the eye and continues onto the tail with disconnected white ocelli at both margins as well as white ocelli inside the stripe; next to the broader dark stripe, a lateral-most stripe is composed of disconnected black ocelli, originating from behind tympanum and reaching to the hindlimb. Upper parts of both hindlimbs and forelimbs with white and black ocelli. Head gray without any markings or spots; labials white with black markings. Belly and underside of tail creamy white, tail dorsum sandy grayish.
The juveniles and subadults are nearly similar in coloration to the adults except for the following details; four longitudinal dark stripes on the body, the outermost originate from anterior parietals on the outer side and continue onto the tail, the innermost originate from the posterior of parietals and merge after running a while on the tail. A broader lateral stripe on each side originates from behind the eye and continues on the lateral side of the body and tail with interspersed white ocelli between the forelimb and hindlimb. The dorsal forelimb and hindlimb are dark gray with white ocelli.
Variations in paratypes. The paratypes of E. rafiqi sp. nov. agree with the holotype with some differences given in Table 2 View TABLE 2 ( Figs. 5 View FIGURE 5 , 6 View FIGURE 6 ). Besides sex, the specimens differ in the arrangement of supralabials i.e. subocular wedged between 6 th and 7 th in all the type series except PMNH 861 View Materials (between 7 th and 8 th) and PMNH 3724 View Materials (between 5 th and 6 th). The arrangement of postmentals has a similar pattern in the paratypes except PMNH 855 View Materials , 861 View Materials , 837 View Materials , 3723 View Materials , 3735 View Materials and 4054, where the fifth chin shield is in contact with the infralabials. The contact of postmental shield with the supralabials varies in the paratypes; PMNH 855 View Materials and 3724, the fifth postmental shield is in contact with 7 th supralabial; PMNH 837 View Materials and 3735, the fifth postmental shield is in contact with sixth supralabial; PMNH 861 View Materials , the fifth postmental shield is in contact with 7 th and 8 th supralabials. The infranasal scale in PMNH 837 View Materials , 855 View Materials , 859 View Materials , 861 View Materials , 3723 View Materials , 3735 View Materials , 4053–4054 View Materials and 4058 rests on first, second and third supralabials. The scale count of dorsals, ventrals, gulars, collars, caudals at 9 th –10 th annuli and lamellae under 4 th toe, however, show a unique value for every specimen within a certain range .
Sexual and age dimorphism. Apparently, males attain larger sizes than females in E. rafiqi sp. nov.: male SVL to 99.3 mm, female SVL 82.1 mm. Moreover, males have generally longer hindlimbs and shorter trunks as compared to females. For a larger female having SVL of 98.1 mm (PMNH 4058), the hindlimb is 56.0 mm against a smaller-sized male (PMNH 855, SVL 92.2 mm) which has a hindlimb length of 57.1 mm. Similarly, the trunk length of a smaller female (PMNH 3724, SVL 75.1 mm) is 36.8 mm against a larger male (PMNH 857, SVL 78.5 mm) which has a trunk length of 34.5 mm. The dorsal body color and pattern, however, varies in juveniles and adults of both genders ( Figs. 5 View FIGURE 5 , 6 View FIGURE 6 ).
Comparison. The new species Eremias rafiqi sp. nov. is strikingly different from species exhibiting striped and ocellate patterns in the subgenus Aspidorhinus ( E. kopetdaghica Szczerbak, 1972 , E. lalezharica Moravec, 1994 , E. papenfussi Mozaffari et al., 2011 , Eremias persica Blanford, 1874 , E. regeli Bedriaga, 1905 , E. fahimii Mozaffari et al., 2020 , E. isfahanica Rastegar-Pouyani et al., 2016 , E. montana Rastegar-Pouyani & RastegarPouyani, 2001 , E. nikolskii Bedriaga, 1905 , E. velox Pallas, 1771 ) and ocellate pattern ( E. killasaifullahi sp. nov., E. afghanistanica Böhme & Szczerbak, 1991 , E. roborowskii Bedriaga, 1912 , E. strauchi Kessler, 1878 , E. suphani Başoğlu & Hellmich, 1968 ). The new species E. rafiqi sp. nov. can also be differentiated from the geographically closely-distributed members of the subgenus Eremias having striped and ocellate pattern ( E. aria Anderson & Leviton 1967 ) and ocellate pattern ( E. nigrocellata Nikolsky 1896 ) by the arrangement of subocular scale which borders the mouth (Supplementary Tab. 1 View TABLE 1 ; see published data in Lantz 1928, Szczerbak 1974, Bischoff & Böhme 1980, Böhme & Szczerbak 1991, Anderson 1999). A brief of morphological differences is provided (the material used for a first-hand comparison is listed in parentheses at each species; see also Table 2 View TABLE 2 and S 1 View TABLE 1 ).
Besides striped and ocellate body pattern (vs. ocellate), E. rafiqi sp. nov. can be distinguished from E. afghanistanica by its larger size (SVL up to 99.3 mm vs. 67.0 mm), higher count of dorsals (56–67 vs. 44–46), gulars (30–36 vs. 25–28), femoral pores (17–22 vs. 16–18), caudal scales in the 9 th –10 th annulus (24–33 vs. 22–25), 5–7 supralabials (mainly 6, rarely 5) located anterior to subocular (vs. 5) and lower number of ventral scales in a single row from the posterior edge of collar to the vent (29–33 vs. 37–38).
From E. persica , that is partly close in dorsal coloration, pattern and size, E. rafiqi sp. nov. differs in the length of interparietal to the length of suture of parietals (longer vs. shorter), length of frontonasal to its width (longer vs. as long as wide), size of the second loreal scale to first loreal scale (more than three times vs. two times), supracaudals (strongly keeled vs. weakly keeled) and tail coloration in the juveniles (sandy grayish vs. bluish).
Besides distant distribution, Eremias rafiqi sp. nov. differs from the recently described E. fahimii by its larger size (SVL up to 99.3 mm vs. 56.0 mm), more SDLT 4 th (22–27 vs. 20–21), the greater number of scales separating the femoral pores (1–4 vs. 1) and the dorsal color and pattern in adults (dorsal stripes broken into ocelli vs. dorsal stripes persistent throughout life).
From E. isfahanica , E. rafiqi sp. nov. differs in the following morphological characters apart from its distant distribution: higher count of supralabials (8–10 vs. 6–8), 5–7 (mainly 6, rarely 5) of them located anterior to subocular (vs. 5), lower count of collars (8–12 vs. 12–15) and the dorsal color pattern in adults (dorsal stripes broken into ocelli vs. dorsal stripes persistent throughout life).
Eremias rafiqi sp. nov. differs from E. kopetdaghica in having a higher count of dorsals (56–67 vs. 48–59), gulars (30–36 vs. 19–28), caudal scales in the 9 th –10 th annulus (24–33 vs. 20–26) and collars (8–12 vs. 7) and the dorsal color and pattern in adults (presence of a broader dark stripe on each lateral side above flanks with disconnected white ocelli at the margins as well as white ocelli inside the stripe vs. no such lateral broader stripes).
Eremias rafiqi sp. nov. can be distinguished from E. lalezharica in having a higher count of dorsals (56–67 vs. 54–59), femoral pores (17–22 vs. 15–19), pair of chin shields/ submaxillary shields (5 vs. 4), lower number of collars (8–12 vs. 13–15), contact of gulars with second pair of submaxillary shields (none vs. 1-2 rows of gulars with the second pair of submaxillary shields) and dorsal color and pattern.
Apart from its peculiar distribution in the remote valley in Torghar Mountains, a part of the Palearctic region, E. rafiqi sp. nov. can be differentiated from E. montana in the following set of characters: larger size (SVL up to 99.3 mm vs. 58.5 mm), higher count of ventral scales in a row across mid-belly in the widest part (14–17 vs. 13–14), number of ventral scales in a single row from the posterior edge of collar to the vent (29–33 vs. 27–28), gulars (30–36 vs. 23–25), infralabials (6–10 vs. 4–6), number of supralabials anterior to the subocular (5–7 vs. 4–5), generally more SDLT 4 th (22–27 vs. 18–25), generally higher count of scales separating the femoral pores (1–4 vs. 2) and dorsal color and pattern.
From E. nigrocellata , E. rafiqi sp. nov. differs in dorsal body pattern (striped and ocellate vs. ocellate), higher count of dorsals (56–67 vs. 42–56), lower number of ventral scales in a row across mid-belly in the widest part (14–17 vs. 18) and the number of femoral pores on each side (17–22 vs. 11–13).
E. rafiqi sp. nov. differs from E. nikolskii by having a higher count of gulars (30–36 vs. 20–28), ventral scales in a row across mid-belly in the widest part (14–17 vs. 14) and dorsal color and pattern.
Our new species stands distinguished from E. papenfussi by its larger size (SVL up to 99.3 mm vs. 62.0 mm), higher count of gulars (30–36 vs. 24–28), number of scales anterior to subocular (5–7, mainly 6 vs. 5), generally higher count of scales separating the femoral pores (1–4 vs. 2) and dorsal color and pattern.
From E. regeli , E. rafiqi sp. nov. differs in having three scales around the penultimate phalanx of 4 th toe (vs. four scales), larger size (SVL up to 99.3 mm vs. 70.0 mm), higher count of gulars (30–36 vs. 14–24), caudal scales in the 9 th –10 th annulus (24–33 vs. 17–25), ventral scales in a row across mid-belly in the widest part (14–17 vs. 13), lower count of femoral pores (17–21 vs. 21–24) and dorsal color and pattern.
The new species E. rafiqi sp. nov. can be easily differentiated from E. strauchi by its distant distribution, color pattern (striped and ocellate vs. ocellate) and larger size (SVL up to 99.3 mm vs. 68.0 mm) besides several other characters. From E. suphani , E. rafiqi sp. nov. differs by its distant distribution, color pattern (striped and ocellate vs. ocellate), larger size (SVL up to 99.3 mm vs. 60.0 mm) and arrangement of gulars (2 rows of gulars reaching to the second pair of chin shields vs. no such arrangement). The new species E. rafiqi sp. nov. can be easily differentiated from E. velox by its distant distribution, larger size (SVL up to 99.3 mm vs. 77.0 mm), contact of infranasal to rostral (separated vs. in contact), generally higher count of gulars (30–36 vs. 19–33) and dorsal color and pattern.
From E. killasaifullahi sp. nov., E. rafiqi sp. nov. differs in the following morphological characters: color pattern (striped and ocellate vs. ocellate), larger size (SVL up to 99.3 mm vs. 70.5 mm), contact of infranasal with the rostral (in contact vs. separated), a higher number of ventral scales in a single row from posterior edge of collar to the vent (29–33 vs. 25–29) and a generally higher count of gulars (29–33 vs. 20–33).
Distribution. The evolutionary clade corresponding with the new species E. rafiqi sp. nov. was genetically confirmed from the areas of SE Iran (Zabol in the Sistan Basin, Sistan and Balochistan Province; the Zabol clade sensu Rastegar-Pouyani et al. 2010), NW Balochistan in Pakistan and south-central Afghanistan ( Rastegar-Pouyani et al. 2010, Khan et al. 2021). In Pakistan, the species is found in three Pakistani districts located along with the Afghan border including Torghar Mountains in the localities of Tanishpa village and Kunder area (Killa Saifullah district), Ashewat (Qamar Din Karez, Zhob district) and Khar in Nushki District.
Habitat and natural history. The type locality and the localities of paratypes (Tanishpa, Kunder and Ashewat) of E. rafiqi sp. nov. are located in the Torghar mountains (means “Black Mountains”), constituting the northern-most part of Toba Kakar Range which is a southern offshoot of the Sulaiman Mountains in the Hindu Kush Mountain system, lying ca. 60 km from the border with Afghanistan ( Fig. 7A View FIGURE 7 ). The Torghar mountains are very rugged semiarid sandstone ridges with an average elevation of 2,400 m and is approximately 90 km long and vary from 15 to 30 km in width. This region is characterized by having dry temperate ecology, with sparse vegetation. A great deal of information about the Torghar mountains including annual weather conditions, ecology, vegetation and sympatric fauna is provided in Masroor et al. (2020b). The species is also found in the Nushki district, ecological part of the Chagai desert (Registan desert, Afghanistan), lying ca. 340 km in aerial distance from the Torghar mountains.
All the specimens were collected between 11:00 am to 02:00 pm. Interestingly, specimens in the Tanishpa, Kunder and Ashewat were caught in the barren area of mixed loamy and sandy habitats at the foothills of Torghar mountains ( Figs. 7B, C View FIGURE 7 ). On the other hand, specimens in the Khar (Nushki district) were collected mostly from the alluvial plain, with dominant vegetation of Haloxylon persicum, Lyceum shawii, Saccharum griffithii, Chenopodium album, Periploca aphylla, Euphorbia prostrata and others.
Mus. Tinro, Vladyvostok
Queen's Gardens, College of Higher Education
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