Early and middle Miocene Sciuridae (Mammalia, Rodentia) from Anatolia, Turkey

Figures & data

FIGURE 1. Early and middle Miocene sciurid rodent tooth localities in Anatolia, Turkey: Kargı 2; Kılçak 0, 0”, 3a, and 3b; Harami 1 and 3; Keseköy; Yapıntı; Kaplangı 1 and 2; Zambal 1; Sarıçay; Bağiçi; and Yenieskihisar. All these localities are situated in lacustrine deposits.

TABLE 1. List of localities with geographical and lithological data.

Locality	District	Province	GPS coordinates	Type of deposit	References
Kargı 2	Osmancık	Çorum	40°52′58.4″N, 34°52′52.5″E	∼25-cm-thick bioturbated lignitic clay with mollusc remains and thick fossilized logs in horizontal position	Krijgsman et al., 1996; De Bruijn et al., 2013b
Kılçak 0+0"	Kalecik	Ankara	40°12′52.4″N, 33°24′20.5″E	The Kılçak sites all sit in a thin section of lacustrine deposits (silts, clays with intercalated coal seams). Kılçak 0 and 0" are in the same section. The positions of Kılçak 3a and 3b relative to one another and relative to Kılçak 0 and 0" could not be established	De Bruijn et al., 2013b
Kılçak 3a+3b	Kalecik	Ankara	40°13′03.6″N, 33°24′06.2″E	See Kılçak 0+0"	De Bruijn et al., 2013b
Harami 1	Ilgın	Konya	38°27′27.2″N, 31°49′30.8″E	∼30-cm-thick poor coal with nodules and mollusc remains, which is the top 30 cm of the coal that is mined	Krijgsman et al., 1996; De Bruijn et al., 2013b
Harami 3	Ilgın	Konya	38°27′27.2″N, 31°49′30.8″E	Thin (±5 cm) coal seam with mollusc remains some 3 m above Harami 1	Krijgsman et al., 1996
Keseköy	Kızılcahamam	Ankara	40°39′51.6″N, 32°40′51.5″E	∼10-cm-thick bioturbated gray-green clay bed intercalated between thick diatomites with plant remains	Krijgsman et al., 1996; De Bruijn et al., 2013b
Yapıntı	Mut	İçel	36°42′43.4″N, 33°22′03.9″E	∼40-cm-thick bed of mottled gray-red sandy mudstones with mollusc remains	Ünay et al., 2001
Kaplangı 1	Banaz	Uşak	38°48′18.1″N, 29°50′11.3″E	∼1-m-thick green-gray silty clay. This site does not exist anymore due to widening of the highway between Uşak and Afyon	De Bruijn et al., 2013b
Kaplangı 2	Banaz	Uşak	38°48′18.1″N, 29°50′11.3″E	∼30-cm-thick poor coal with mollusc remains that is exposed in a coal exploration test quarry	De Bruijn et al., 2013b
Zambal 1	Osmancık	Çorum	41°02′59.8″N, 34°59′19.6″E	Horizontally layered lignitic clay bed at hill top. This site has disappeared after a landslide	De Bruijn et al., 2013b
Sarıçay	Milas	Muğla	37°19′52.4″N, 27°47′39.7″E	Fluviolacustrine deposits; see Sickenberg et al., 1975:87–88	Sickenberg et al., 1975
Bağiçi	Bala	Ankara	39°31′56.8″N, 32°54′42.0″E	∼40-cm-thick horizontally layered gray clay with fragments of large bones	De Bruijn et al., 2013b
Yenieskihisar	Yatağan	Muğla	37°18′40.4″N, 28°02′44.7″E	Lacustrine deposits; see Sickenberg et al., 1975:29–30	Sickenberg et al., 1975

TABLE 2. List of localities with proposed biochronological ages.

Locality	Informal local rodent zone^a	Correlation to European MP/MN zonation^b	Epoch
Yenieskihisar	Zone H		middle Miocene
Bağiçi	Zone H	MN 7+8	middle Miocene
Sarıçay	Zone H		middle Miocene
Zambal 1	Zone G	MN 6	middle Miocene
Kaplangı 2	Zone E	MN 4	early Miocene
Kaplangı 1	Zone E	MN 4	early Miocene
Yapıntı		MN 3/MN 4^c	early Miocene
Keseköy	Zone D	MN 3	early Miocene
Harami 3	Zone C		early Miocene
Harami 1	Zone C	MN 2	early Miocene
Kılçak 3b	Zone B	MN 1	early Miocene
Kılçak 3a	Zone B	MN 1	early Miocene
Kılçak 0"	Zone B	MN 1	early Miocene
Kılçak 0	Zone B	MN 1	early Miocene
Kargı 2	Zone B	MP 30?	late Oligocene

^aÜnay et al. (2003).

^bDe Bruijn et al. (2013b)

^cAge assignment from Ünay et al. (2001).

FIGURE 2. A–K, Palaeosciurus feignouxi from Kılçak 0, 0”, and 3a. A, Ki0"-1, D4 (reversed); B, Ki0"-2, P4; C, Ki3a-5, M1/2; D, Ki0-1, M1/2 (reversed); E, Ki3a-11, p4; F, Ki3a-13, p4 (reversed); G, Ki3a-19, m1; H, Ki3a-16, m1; I, Ki0"-7, m2 (reversed); J, Ki3a-22, m2; K, Ki3a-25, m3 (reversed). L–M, Palaeosciurus cf. sutteri from Kaplangı 1. L, Kap1-101, m1; M, Kap1-103, m3. N, Dehmisciurus obtusidens from Harami 3. Har3-1, m2 (reversed). Scale bar equals 1 mm.

TABLE 3. Measurements (in mm) of teeth of Palaeosciurus feignouxi from Kılçak 0, 0", 3a, and 3b, Palaeosciurus cf. feignouxi from Kargı 2, and Palaeosciurus aff. feignouxi from Keseköy.

			Length				Width
Species	Locality	Element	Min	Mean	Max	N	Min	Mean	Max
Palaeosciurus aff. feignouxi	Keseköy	P3	—	0.83	—	1/1	—	0.86	—
Palaeosciurus feignouxi	Kılçak 0"	D4	—	1.77	—	1/1	—	1.77	—
Palaeosciurus feignouxi	Kılçak 3a	D4	—	1.72	—	1/1	—	1.76	—
Palaeosciurus aff. feignouxi	Keseköy	D4	—	1.50	—	1/1	—	1.70	—
Palaeosciurus feignouxi	Kılçak 0"	P4	1.60	1.64	1.68	2/2	1.84	1.90	1.95
Palaeosciurus feignouxi	Kılçak 3a	P4	—	1.70	—	1/1	—	1.90	—
Palaeosciurus cf. feignouxi	Kargı 2	P4	—	1.50	—	1/0	—	—	—
Palaeosciurus aff. feignouxi	Keseköy	P4	1.73	1.74	1.75	2/2	2.00	2.01	2.01
Palaeosciurus feignouxi	Kılçak 0	M1/2	—	2.02	—	1/1	—	2.29	—
Palaeosciurus feignouxi	Kılçak 0"	M1/2	—	1.76	—	1/1	—	2.24	—
Palaeosciurus feignouxi	Kılçak 3a	M1/2	1.83	1.97	2.16	5/5	2.13	2.31	2.54
Palaeosciurus feignouxi	Kılçak 3b	M1/2	—	1.92	—	1/1	—	2.23	—
Palaeosciurus cf. feignouxi	Kargı 2	M1/2	—	1.84	—	1/1	—	2.13	—
Palaeosciurus aff. feignouxi	Keseköy	M1/2	1.88	1.92	1.95	3/3	2.13	2.16	2.18
Palaeosciurus feignouxi	Kılçak 3a	M3	—	2.18	—	1/2	2.07	2.12	2.16
Palaeosciurus cf. feignouxi	Kargı 2	M3	—	—	—	0/1	—	1.91	—
Palaeosciurus aff. feignouxi	Keseköy	M3	2.18	2.19	2.20	2/2	2.12	2.13	2.13
Palaeosciurus feignouxi	Kılçak 0"	d4	—	1.54	—	1/1	—	1.38	—
Palaeosciurus feignouxi	Kılçak 3a	p4	1.58	1.76	1.91	3/3	1.66	1.73	1.79
Palaeosciurus aff. feignouxi	Keseköy	p4	1.71	1.76	1.80	2/2	1.55	1.59	1.62
Palaeosciurus feignouxi	Kılçak 3a	m1	1.89	1.99	2.08	5/5	2.00	2.10	2.18
Palaeosciurus aff. feignouxi	Keseköy	m1	—	2.17	—	1/1	—	1.99	—
Palaeosciurus feignouxi	Kılçak 0	m2	—	2.11	—	1/1	—	2.21	—
Palaeosciurus feignouxi	Kılçak 0"	m2	—	2.15	—	1/1	—	2.27	—
Palaeosciurus feignouxi	Kılçak 3a	m2	2.04	2.09	2.17	3/3	2.10	2.34	2.47
Palaeosciurus feignouxi	Kılçak 0	m3	—	2.51	—	1/1	—	2.12	—
Palaeosciurus feignouxi	Kılçak 3a	m3	2.44	2.58	2.71	2/2	2.07	2.14	2.21
Palaeosciurus aff. feignouxi	Keseköy	m3	—	2.37	—	1/1	—	2.16	—

N = number of measurements (length/width).

FIGURE 3. Palaeosciurus aff. feignouxi from Keseköy. A, Kes-101, P3 (reversed); B, Kes-102, D4 (reversed); C, Kes-103, P4; D, Kes-104, P4 (reversed); E, Kes-107, M1/2 (reversed); F, Kes-105, M1/2; G, Kes-109, M3 (reversed); H, Kes-108, M3; I, Kes-112, p4 (reversed); J, Kes-111, p4; K, Kes-114, m1; L, Kes-116, m3. Scale bar equals 1 mm.

FIGURE 4. Protospermophilus kelloggi from Split Rock, U.S.A. Comparative material. A, CM 14883, D4; B, CM 15924, P4 (reversed); C, CM 15924, M1/2 (reversed); D, CM 15924, M1/2 (reversed); E, CM 14295, M1/2; F, CM 13521, M3; G, CM14661, p4 (reversed); H, UU 475, m1; I, CM 15855, m2 (reversed); J, CM 15855, m3 (reversed); K, CM 13524, m3. Scale bar equals 1 mm.

FIGURE 5. A–H, Spermophilinus besana from Sarıçay and Yenieskihisar. A, YE-110, D4; B, SC-103, P4; C, YE-126, M1/2; D, YE-123, M1/2; E, YE-135, M3; F, YE-141, d4; G, YE-151, m1; H, YE-168, m2. I–U, Spermophilinus bredai from Zambal 1, Sarıçay, Bağiçi, and Yenieskihisar. I, YE-105, D4; J, YE-103, D4; K, SC-101, P4; L, Zam1-101, M1/2; M, Zam1-102, M1/2; N, Bag-861, M1/2; O, YE-113, M1/2; P, YE-112, M1/2; Q, YE-131, M3; R, YE-146, p4; S, YE-154, m1; T, YE-164, m2; U, YE-173, m3. All images reversed except E, J, L, and Q. Scale bar equals 1 mm.

FIGURE 6. A–G, Tamias cf. eviensis from Yapıntı. A, Yap-101, D4; B, Yap-104, M1/2 (reversed); C, Yap-103, M1/2; D, Yap-106, M3; E, Yap-108, m2; F, Yap-110, m2 (reversed); G, Yap-109, m2. H–L, Atlantoxerus adroveri from Bağiçi and Yenieskihisar. H, YE-193, P4; I, Bag-888, M1/2 (reversed); J, Bag-895, m1; K, YE-201, p4; L, Bag-890, m1 (reversed). Scale bar equals 1 mm.

FIGURE 7. A–B, Hylopetes sp. from Kılçak 0 and 0”. A, Ki0"-11, P4; B, Ki0-11, P4. C–D, Aliveria luteyni from Kaplangı 1. C, Kap1-122, D4; D, Kap1-127, m3. E, Blackia sp. from Sarıçay. SC-131, m1. F–G, Miopetaurista cf. dehmi from Harami 1. F, Har1-1732, P4; G, Har1-1736, M1/2. H–I, Albanensia albanensis from Sarıçay. H, SC-141, P4; I, SC-147, m1. All images reversed except B. Scale bar equals 1 mm.

TABLE 4. Measurements (in mm) of teeth of Spermophilinus besana from Harami 1, Kaplangı 1 (S. cf. besana), Sarıçay, Bağiçi, and Yenieskihisar.

		Length				Width
Locality	Element	Min	Mean	Max	N	Min	Mean	Max
Yenieskihisar	P3	—	0.55	—	1/1	—	0.52	—
Harami 1	D4	—	1.59	—	1/1	—	1.57	—
Kaplangı 1	D4	—	1.23	—	1/1	—	1.32	—
Bağiçi	D4	—	1.39	—	1/1	—	1.50
Yenieskihisar	D4	1.16	1.20	1.24	2/2	1.22	1.27	1.32
Sarıçay	P4	—	1.36	—	1/1	—	1.71	—
Harami 1	M1/2	1.53	1.67	1.78	7/6	1.84	1.91	1.97
Bağiçi	M1/2	—	1.56	—	1/0	—	—	—
Yenieskihisar	M1/2	1.38	1.49	1.64	9/8	1.72	1.82	1.96
Harami 1	M3	1.96	2.03	2.09	2/1	—	1.89	—
Yenieskihisar	M3	1.86	1.90	1.97	3/3	1.77	1.81	1.85
Harami 1	d4	—	1.44	—	1/1	—	1.15	—
Bağiçi	d4	—	1.30	—	1/2	1.04	1.07	1.09
Yenieskihisar	d4	—	1.11	—	1/1	—	0.97	—
Harami 1	p4	—	1.56	—	1/1	—	1.41	—
Sarıçay	p4	—	1.34	—	1/1	—	1.29	—
Harami 1	m1	1.68	1.71	1.73	2/2	1.59	1.61	1.62
Sarıçay	m1	1.60	1.62	1.63	2/2	1.57	1.66	1.74
Bağiçi	m1	1.48	1.60	1.71	2/1	—	1.41	—
Yenieskihisar	m1	1.47	1.49	1.50	2/2	1.62	1.70	1.77
Harami 1	m2	1.55	1.77	2.00	3/3	1.56	1.75	1.93
Bağiçi	m2	1.66	1.79	1.86	4/4	1.84	1.94	1.99
Yenieskihisar	m2	—	1.76	—	1/1	—	1.96	—
Harami 1	m3	—	—	—	0/2	1.92	1.97	2.02
Bağiçi	m3	—	1.93	—	1/1	—	2.01	—
Yenieskihisar	m3	—	1.84	—	1/1	—	1.74	—

Because distinction between the teeth of S. besana and S. bredai can only be made on the basis of size, small specimens of S. bredai from Sarıçay, Bağiçi, or Yenieskihisar may incidentally have been misidentified as S. besana, and large specimens of S. besana as S. bredai (see also Table 5). N = number of measurements (length/width).

FIGURE 8. Length/width scatter plots for upper (D4, P4, M1/2, and M3) and lower (d4, p4, m1, m2, and m3) tooth elements of Spermophilinus besana from Harami 1 (▴), Sarıçay (♦), Bağiçi (▾), and Yenieskihisar (●), and Spermophilinus bredai from Zambal 1 (□), Sarıçay (◊), Bağiçi (▿), and Yenieskihisar (○). Material from Paşalar (▹) is included for comparison.

TABLE 5. Measurements (in mm) of teeth of Spermophilinus bredai from Zambal 1, Sarıçay, Bağiçi, and Yenieskihisar.

		Length				Width
Locality	Element	Min	Mean	Max	N	Min	Mean	Max
Yenieskihisar	D4	1.56	1.68	1.79	3/3	1.60	1.67	1.74
Sarıçay	P4	—	1.67	—	1/1	—	2.04	—
Zambal 1	M1/2	1.74	1.78	1.82	2/2	2.11	2.16	2.20
Sarıçay	M1/2	1.69	1.72	1.75	2/2	2.09	2.10	2.10
Bağiçi	M1/2	—	1.90	—	1/1	—	2.37	—
Yenieskihisar	M1/2	1.74	1.89	2.00	5/5	2.04	2.21	2.39
Yenieskihisar	M3	—	2.14	—	1/2	1.99	2.04	2.08
Sarıçay	d4	—	1.40	—	1/1	—	1.15	—
Sarıçay	p4	—	1.71	—	1/1	—	1.64	—
Bağiçi	p4	1.51	1.57	1.62	2/1	—	1.45	—
Yenieskihisar	p4	1.48	1.54	1.57	4/4	1.30	1.38	1.42
Yenieskihisar	m1	—	1.71	—	1/1	—	1.72	—
Sarıçay	m2	—	1.90	—	1/1	—	2.13	—
Yenieskihisar	m2	1.99	2.06	2.12	3/3	2.13	2.16	2.20
Sarıçay	m3	2.22	2.26	2.29	2/2	2.13	2.20	2.26
Yenieskihisar	m3	2.30	2.31	2.32	2/2	2.35	2.36	2.36

Because distinction between the teeth of S. bredai and S. besana can only be made on the basis of size, large specimens of S. besana from Sarıçay, Bağiçi, or Yenieskihisar may occasionally have been misidentified as S. bredai, and small specimens of S. bredai as S. besana (see also Table 4). N = number of measurements (length/width).

FIGURE 9. Length/width scatter plot for M1/2 of Spermophilinus besana from Harami 1 (▴) and Yenieskihisar (●) and Spermophilinus bredai from Yenieskihisar (○). The M1/2s from Harami 1 fall within the size range of 15 randomly selected M1/2s of Spermophilinus besana from Vieux Collonges, France (+). See text for details.

FIGURE 10. Ranges and mean values of the length of M1/2 in assemblages of Spermophilinus (cf.) besana, S. (cf.) bredai, and S. (cf.) turolensis from southeastern Europe and Anatolia. Data from the literature (see Table 6 for references) and this study. The numbers of teeth on which values are based are indicated next to the horizontal lines and dots. Kaplangı 1 and Kastellios K1 did not yield M1/2s. No measurements are available for the species from Silata. For Paşalar, two sets of data are given: those provided by Peláez-Campomanes and Daams (2002) (Paşalar B) and those from this study (Paşalar A). For comparison, this figure includes measurements of M1/2 in the type collections of S. besana (Vargas 1A, Spain; MN 4) (Cuenca Bescós, 1988), and S. turolensis (Los Mansuetos, Spain; MN 12) (De Bruijn and Mein, 1968). In addition, values for M1/2 of S. besana from Vieux Collonges (France; MN 4 and MN 5) are included (cf. Fig. 9). For S. bredai, measurements of M1/2s from La Grive L7 and M (France; MN 7) are used.

TABLE 6. Localities with Spermophilinus in southeastern Europe and Anatolia.

Locality	Country	Age	Spermophilinus (cf.) besana	Spermophilinus (cf.) bredai	Spermophilinus (cf.) turolensis	Spermophilinus giganteus	Reference
Maritsa	Greece	MN 14				X	De Bruijn et al., 1970
Silata	Greece	MN 13–MN 14 boundary			X		Vasileiadou et al., 2003
Maramena	Greece	MN 13			X		De Bruijn, 1995
Mytilinii	Greece	MN 12		X			Vasileiadou and Sylvestrou, 2009
Düzyayla lignite	Turkey	MN 12		X			Bosma et al., 2013
Düzyayla clay	Turkey	MN 12		X			Bosma et al., 2013
Hayranlı 1	Turkey	MN 10-11		X			Bosma et al., 2013
Kastellios K1	Greece	MN 10		X			De Bruijn et al., 1971
Altıntaş 1	Turkey	MN 9			X		Bosma et al., 2013
Tuğlu	Turkey	MN 9	X				Joniak and De Bruijn, 2015
Yenieskihisar	Turkey	MN 7+8	X	X			This study
Bağiçi	Turkey	MN 7+8	X	X			This study
Sarıçay	Turkey	MN 7+8	X	X			This study
Zambal 1	Turkey	MN 6		X			This study
Çandır 2	Turkey	MN 5		X			De Bruijn et al., 2003
Paşalar	Turkey	MN 5		X			Peláez-Campomanes and Daams, 2002
Antonios	Greece	MN 4–MN 5 boundary	X				Vasileiadou and Koufos, 2005
Kaplangı 1	Turkey	MN 4	X				This study
Harami 1	Turkey	MN 2 or 1	X				This study

FIGURE 11. Length/width scatter plots for D4, M1/2, M3, and m2 of Tamias species from Greece and Anatolia. Tamias cf. eviensis from Yapıntı, Anatolia (●) (this study), T. eviensis from the type locality, Aliveri, Greece (○) (De Bruijn et al., 1980), T. anatoliensis from the type locality, Altıntaş 1, Anatolia (□) (Bosma et al., 2013), and T. atsali from the type locality, Maramena, Greece (◊) (De Bruijn, 1995). Teeth from Aliveri and Maramena were remeasured to ensure precise comparison.

TABLE 7. Measurements (in mm) of teeth of Atlantoxerus adroveri from Bağiçi and Yenieskihisar.

		Length				Width
Locality	Element	Min	Mean	Max	N	Min	Mean	Max
Yenieskihisar	P3	—	1.35	—	1/1	—	1.32	—
Yenieskihisar	P4	—	2.50	—	1/1	—	3.36	—
Bağiçi	M1/2	—	2.46	—	1/1	—	3.10	—
Yenieskihisar	M1/2	2.55	2.59	2.63	2/2	3.33	3.39	3.45
Yenieskihisar	M3	—	2.58	—	1/1	—	3.21	—
Bağiçi	p4	—	2.75	—	1/1	—	2.73	—
Yenieskihisar	p4	2.47	2.54	2.63	3/2	2.46	2.64	2.81
Bağiçi	m1	2.62	2.79	2.86	4/4	2.55	2.78	3.02
Yenieskihisar	m1	—	2.70	—	1/1	—	3.18	—
Yenieskihisar	m2	2.75	2.78	2.84	3/3	3.01	3.12	3.21
Yenieskihisar	m3	—	—	—	0/1	—	3.24	—

N = number of measurements (length/width).

FIGURE 12. Occurrences of Sciuridae at 15 early and middle Miocene localities in Anatolia, Turkey.

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