New rodents from the late Oligocene site of Gözükızıllı in Anatolia (Turkey)

Figures & data

Figure 1. Satellite image of the area near Gözükızıllı village with the two sites where fossil rodents have been collected. Image from Google Earth, date 19/06/2020, copyright 2020 CNES/Airbus, 2020 Google, 2020 Basarsoft

Figure 2. Nomenclature of tooth parts of Daxneria nov. gen. described in this paper (adapted after Flynn et al. 1986).

Figure 3. Upper cheek teeth of Daxneria fragilis nov. gen. nov. sp. (a-j) and Sayimys sp. (k) from Gözükızıllı-1, occlusal, lingual or antero-lingual view. Daxneria fragilis: a & e GOZ1b-237, b GOZ1a-195, c GOZ1b-233, d GOZ1b-277, f & g GOZ1b-228, h upper incisor, no number, i & j GOZ1a-199. Sayimys sp.: k GOZ1b-251

Figure 4. Lower cheek teeth Daxneria fragilis nov. gen. nov. sp. and Sayimys sp. (lower row) from Gözükızıllı-1, occlusal and labial views. Daxneria fragilis: a & e GOZ1a-219, b GOZ1b-247, c & f GOZ-1b-248, d & g GOZ1b-241, h & i GOZ1a-220, j & k GOZ1a-202, l lower incisor no number. Sayimys sp.: m & n GOZ1b-253, o & p GOZ1b-252

Figure 5. Sections of the lower incisor of Daxneria fragilis nov. gen. nov. sp. a-c transverse sections; b enlargement of square in a, c enlargement of square in b. d-f sagittal sections: e enlargement of square in d, f enlargement of square in e

Table 1. Measurements of Daxneria fragilis nov. gen. nov. sp. from Gözükızıllı-1a and 1b; shown are minimum, maximum, mean, number of observations and standard deviation

D. fragilis	Length					Width
GOZ1a & 1b	min	max	mean	N	SD	min	max	mean	N	SD
D3	0.5	0.70	0.63	10	0.059	0.71	0.88	0.806	10	0.067
DP4	1.76	1.90	1.843	4	0.060	1.93	2.15	2.045	4	0.090
M1-2	2.35	2.71	2.518	4	0.152	2.47	2.63	2.550	5	0.060
M3	2.98	3.54	3.260	2	–	3.12	3.20	3.160	2	–
dp4	2.05	2.30	2.160	7	0.094	1.45	1.77	1.600	11	0.093
m1-2	2.45	2.85	2.608	5	0.147	1.99	2.69	2.401	10	0.235
m3	–	–	3.430	1	–	–	–	3.190	1	–

Figure 6. Upper and lower cheek teeth of Baluchimys ganeshapher (a, b, c, d, m, n, o, p), Baluchimys barryi (l, f, g, h, q, r, s, t) and Lindsaya derabugtiensis, all three from site Y-GSP417 (Flynn et al., 1986). The bar represents 1 mm; the approx. enlargement has been added

Figure 7. Comparison of upper cheek teeth of Daxneria fragilis (v-y) with Bugtimys Marivaux et al., 2002 site Paali nala C2 (a-e), Hodsahibia Marivaux & Welcomme 2003 site Paali nala C2 (f-h), H. gracilis Paali nala C2 (j-m), Lophibaluchia Flynn et al., 1986 site Y-GSP417 (n-q) and Ottomania de Bruijn et al., 2003 site Süngülü. Daxneria, Ottomania and Lophibaluchia retained their DP4 and have no P4. The bar represents 1 mm; the approx. enlargements have been added, these are not the same

Figure 8. Comparison of lower cheek teeth of Daxneria fragilis with Bugtimys Marivaux et al. (2002) site Paali nala C2, (a-e); Hodsahibia beamshaiensis Marivaux and Welcomme (2003) site Paali nala C2, (f-h); H. gracilis Marivaux and Welcomme (2003) site Paali nala C2, (i-k); Lophibaluchia Flynn et al. (1986) site Y-GSP417, (l-o) and Ottomania de Bruijn et al. (2003), site Süngülü, (p-s). Daxneria, Ottomania and Lophibaluchia retained their DP4 and have no P4. The bar represents 1 mm; the approx. enlargements have been added, these are not the same

Figure 9. Comparison of Sayimys sp. from Gözükızıllı with Sayimys aff. obliquidens from the middle member of the Chul’adyr Formation Aktau Mountains, Kazakhstan (e-i) and S. flynni from Z113, Zinda Pir, Pakistan (j-m)

Table 2. List of species of Eucricetodon from Europe. The allocation into groups is after Freudenthal & Martin-Suarez (2016)

Species	Type locality; age; group; remarks
E. atavus (Misonne, 1957)	Hoogbutsel, Belgium; early Oligocene, MP21; Atavocricetodon grp
E. atavoides Freudenthal, 1996	Ollalla 4, Spain; early Oligocene, MP22; Atavocricetodon grp
E. nanoides Freudenthal, 1996	Ollalla 4, Spain; early Oligocene, MP22; Atavocricetodon grp
E. minusculus Freudenthal, 1996	Ollalla 4, Spain; early Oligocene, MP22; Atavocricetodon grp
E. nanus Peláez-Campomanes, 1995	Valdecollares, Spain; early Oligocene, MP23; Atavocricetodon grp
E. hugueneyae Freudenthal, 1996	Montalbán 1D, Spain; early Oligocene, MP23; Atavocricetodon grp
E. huberi (Schaub, 1925)	Mümliswil, Switzerland; late Oligocene, MP26; huerzeleri grp
E. martinensis Freudenthal, 1994	Mirambueno 4D, Spain; late Oligocene, MP26; huerzeleri grp
E. margaritae Daams et al. 1989	Pareja, Spain; late Oligocene, MP26; huerzeleri grp
E. huerzeleri Vianey-Liaud, 1972	Oensingen, Switzerland; late Oligocene, MP26; huerzeleri grp
E. robustus Augusti & Arbiol, 1989	Fraga 4, Spain; late Oligocene, MP27; dubius grp
E. quercyi Vianey-Liaud, 1972	Pech-du-Fraysse, France; late Oligocene, MP28; dubius grp
E. liber Ziegler 1994	Herlingen 9, Germany; late Oligocene, MP29; dubius grp
E. thezelensis Comte, 1985	Thézels, France; late Oligocene, MP30; dubius grp
E. collatus (Schaub, 1925)	Küttigen, Switserland; latest Oligocene, MP30; collatus grp; holotype by Thaler 1966
E. longidens Hugueney, 1969	Coderet, France; latest Oligocene, MP30; collatus grp; originally subspecies of E. collatus
E. hesperius Engesser, 1985	Paulhiac, France; earliest Miocene, MN1; collatus grp
E. cetinensis Daams, 1976	Cetina de Aragon, Spain; early Miocene, MN2; collatus grp
E. gerandianus (Gervais, 1848-1852)	Langy, France; early Miocene; collatus grp; see Daams 1976
E. aquitanicus Baudelot & de Bonis, 1968	Laugnac, France; early Miocene, MN2; collatus grp
E. hochheimensis (Schaub, 1925)	Hochheim, Germany; early Mioc.; collatus grp; known from a single m2 only (Hugueney 1999)
E. haslachensis (Schaub, 1925)	Haslach/Ulm, Germany; early Mioc.; MN2?; collatus grp; holotype by Dienemann 1987
E. infralactorensis (Viret, 1930)	Estrepouy, France; early Miocene, MN3; collatus grp; see Hugueney & Bulot 2011
E. quadratus (Viret, 1930)	Estrepouy, France; early Miocene, MN3; collatus grp; see Hugueney & Bulot 2011

Table 3. Eucricetodon species based on limited type material from unknown type localities and of uncertain age

Species	Type locality
E. gergovianum (Schaub, 1925)	Quercy, France; Oligocene
E. murinus (Schlosser, 1884)	Quercy, France; Oligocene
E. praecursor (Schaub, 1925)	Quercy, France; Oligocene
E. incertus (Schlosser, 1884)	Quercy, France; Oligocene
E. dubius (Schaub, 1925) *	Quercy, France; Oligocene

*included in Heterocricetodon by Hugueney (1980) and Engesser (1987)

Table 4. List of Asian Eucricetodon species

Species	locality; age
E. kurthi de Bruijn et al., 2003	Süngülü, Turkey; latest Eocene
E. wangae Li et al., 2016	Red Beds, Erden Obo, China; latest Eocene
E. caducus (Shevyreva, 1967)	Zayzan depression, Kazakhstan; early Oligocene
E. deploratus (Shevyreva, 1967)	Zayzan depression, Kazakhstan; early Oligocene
E. paaliensis (Marivaux et al., 1999)	Paali Nala C2, Pakistan; early Oligocene
E. asiaticus (Matthew and Granger, 1925)	Hsanda Gol, China; middle Oligocene
E. bagus Gomes Rodrigues et al., 2012	Ulantatal UTL4, China; late Oligocene
E. jilantaiensis Gomes Rodrigues et al., 2012	Ulantatal UTL4, China; late Oligocene
E. occasionalis Lopatin, 1996	Altyn Shokysu bone bed 4, Kazakhstan; late Oligocene
E. sajakensis Bendukidze, 1993	Sayaken, Kazakhstan; late Oligocene
E. youngi Li & Qiu, 1980	site IVPP#78027,near Tianjiazhai, Xining basin, China; early Mioc.

Figure 10. Length-width scatter diagrams of the molars of the two species of Eucricetodon from Gözükızıllı-1 (red points) and E. margaritae (black crosses) from Pareja. The line separates the Gözükızıllı points into E. oculatus nov. sp. and E. ruber nov. sp. The measurements of E. margaritae from Pareja have been digitised from Daams et al. (1989, Figure 3)

Figure 11. The length of M1 (range and average) of Eucricetodon from Gözükızıllı compared with that of late Oligocene and early Miocene species of Eucricetodon from Europe. For Bouzigues see Aguilar (1974); for Cetina (de Aragon) see Daams (1976); for Estrepouy see Hugueney & Bulot: for Fraga 4 see Augusti & Arbiol 1989; for Laugnac see Daams (1976); for Mirambueno 4D see Freudenthal (1994). for Oensingen, St-Victor-la-Coste and Pech-du-Fraysse see Vianey-Liaud (1972); for Paulhiac, Boudry and Küttigen see Engesser 1985; for Pareja see Daams et al. (1989); for Ulm & Weiss-6 see Dienemann (1987)

Figure 12. Nomenclature of parts of Eucricetodon molars (adapted after Mein and Freudenthal 1971b)

Figure 13. Upper molars of Eucricetodon oculatus, E. ruber from Gözükızıllı-1 and E. margaritae from Pareja (Spain). E. occulatus: a GOZ1b-151, b GOZ1b-131, c GOZ1b-102 holotype, d GOZ1b-141, e GOZ1b-121, f GOZ1b-111; E. ruber: g GOZ1b-156, h GOZ1b-126, i GOZ1b-106, j GOZ1b-144, k GOZ1b-128, l GOZ1b-105 holotype; E. margaritae: m-o

Figure 14. Lower molars of Eucricetodon oculatus,Eucricetodon oculatus, E. ruber from Gözükızıllı-1 E. ruber from Gözükızıllı-1 and E. margaritae from Pareja (Spain). E. oculatus: a GOZ1b-202, b GOZ1b-183, c GOZ1b-162, d GOZ1b-211, e GOZ1b-182, f GOZ1b-161; E. ruber: g GOZ1b-216, h GOZ1b-186, i GOZ1b-166, j GOZ1b-204, k GOZ1b-188, l GOZ1b-167; E. margaritae: m-o

Table 5. Measurements of Eucricetodon ruber nov. sp. from Gözükızıllı-1 and Güvendik

E. ruber	Length					Width
Gözükızıllı-1	min	max	mean	N	SD	min	max	mean	N	SD
M1	1.92	2.07	2.016	16	0.044	1.30	1.45	1.388	17	0.044
M2	1.37	1.58	1.479	20	0.062	1.33	1.50	1.419	20	0.049
M3	0.97	1.22	1.124	18	0.069	1.18	1.30	1.244	18	0.037
m1	1.60	1.87	1.720	15	0.072	1.08	1.24	1.157	15	0.054
m2	1.42	1.69	1.561	12	0.078	1.24	1.42	1.335	12	0.052
m3	1.37	1.51	1.456	8	0.045	1.17	1.31	1.230	8	0.047
Güvendik
m1	–	–	–	–	–	–	–	1.140	1	–
m3	–	–	1.600	1	–	–	–	1.220	1	–

Table 6. Measurements of Eucricetodon oculatus nov. sp. from Gözükızıllı-1 and Güvendik

E. oculatus	Length					Width
Gözükızıllı-1	min	max	mean	N	SD	min	max	mean	N	SD
M1	2.10	2.45	2.228	4	0.155	1.48	1.66	1.546	8	0.062
M2	1.52	1.76	1.647	6	0.090	1.57	1.71	1.638	5	0.050
M3	1.32	1.42	1.374	5	0.046	1.38	1.55	1.442	5	0.066
m1	1.87	1.90	1.885	2	0.021	1.30	1.31	1.305	2	0.007
m2	1.68	1.87	1.774	5	0.07	1.37	1.5	1.45	5	0.051
m3	1.74	1.93	1.812	5	0.073	1.36	1.48	1.430	5	0.046
Güvendik
M1	–	–	2.240	1	–	–	–	1.430	2	–
m3	1.66	1.67	1.635	2	–	1.22	1.38	1.300	2	–

Figure 15. Transverse sections (a-e) and sagittal sections (f-h) of the lower incisor of Eucricetodon from Gözükızıllı-1

Table 7. Character states in upper molars of Eucricetdon ruber nov. sp. and E. oculatus nov. sp. from Gözükızıllı-1

	M1		M2		M3
	E. ruber	E. oculatus	E. ruber	E. oculatus	E. ruber	E. oculatus
Anterior arm protocone to paracone	5/19	2/6	–	–	–	–
Anterior arm protocone free ending	14/19	4/6	–	–	–	–
Mesoloph absent to very small	9/19	2/7	7/18	1/6	5/20	0/2
Mesoloph medium long	8/19	4/7	12/8	2/6	13/20	1/2
Mesoloph long	2/19	1/7	1/19	3/6	2/20	1/2
Para- and metacone spurs absent	4/17	2/8	7/19	4/6	–	–
Para- and metacone spurs present	5/17	2/8	11/19	2/6	–	–
Ectolophs present	8/17	4/8	2/19	0/6	–	–
Metaloph interrupted	4/17	1/7	–	–	–	–
Metaloph complete	13/17	6/7	–	–	–	–
Sinus incompletely closed	–	–	–	–	11/18	4/5
Sinus closed	–	–	–	–	7/18	1/5

Table 8. Schmelzmuster types of lower incisors in Eucricetodon samples. The allocation to group (grp) is after Freudenthal and Martín-Suárez (2016)

	Species, locality, (group)	Type	Age	References
	E. infralactorensis, Estrepouy, (collatus grp)	Type 4	MN3, early Mioc.	Kalthoff 2000
	E. gerandianus, La Chaux 7, (collatus grp)	Type 4	MN2a, early Mioc.	Kalthoff 2000
	E. longidens, Coderet, (collatus grp)	Type 4	MP30, late Oligoc.	Kalthoff 2000
	E. liber, Herrlingen 9, (dubius grp)	Type 4	MP29, late Oligoc.	Kalthoff 2000
	E. dubius, Herrlingen 8, (dubius grp)	Type 4	MP28, late Oligoc.	Kalthoff 2000
	Eucricetodon sp., Ugljevik	Type 10	MP 27–28, late Oligoc.	van de Weerd et al. in prep.
	E. huerzeleri, Mas de Pauffié, (huerz. grp)	Type 1	MP26, late Oligoc.	Gomes Rodrigues et al. 2013
	Eucricetodon, Gözükızıllı-1	Type 10	MP25-26, late Oligoc.	This paper
	E. cf. huberi, Belgaric-1, (huerz. grp)	Type 8	MP25, late Oligoc.	Gomes Rodrigues et al. 2013
	E. atavus, Itardies, (Atavocr. grp)	Type 1	MP23, early Oligoc.	Kalthoff 2000, Gomes Rodrigues et al. 2013
	E. aff. nanus, Cavalé, (Atavocr. grp)	Type 1	MP22, early Oligoc.	Gomes Rodrigues et al. 2013

Figure 16. The Dipodidae and Gliridae from Gözükızıllı: a Dipodidae gen. et sp. indet. GOZ1b-260; Bransatoglis cf. sjeni: b GOZ1b-255, c GOZ1b-258; Peridyromys or Microdyromys: d GOZ1b-257

Figure 17. Chronostratigraphic scheme of Palaeogene faunas from Anatolia and the wider region of the Balkans and Southern Asia. Broken lines indicate uncertainty ranges, duration of these are arbitrarily set at 8% of the estimated age. Locations with a magneto-stratigraphic or chronostratigraphic age estimate have a solid range and names are underlined. References on the sites are: Aral Fm (Lucas et al. 1998; Bendukidze et al. 2009), Banovice (de Bruijn et al. 2013), Benara (Métais et al. 2016), Buštranje, Strelac, Valniš, Zvonce (de Bruijn et al. 2018ab), Harami, Inkonak, Kargi, Yeniköy, Keseköy (Krijgsman et al. 1996; Ünay et al. 2003a), Kala Chitta Range/upper Subathu Grp. (Hussain et al. 1978; Gingerich 2003), Kavakdere, Kocayarma (Ünay-Bayraktar 1989), Kyprinos (Doukas and Theocharopoulos 1999), Paragovo, Ugljevik (Marković et al. 2019; van de Weerd et al. in prep), Sabuncubeli (de Bruijn et al. 2006), Süngülu (de Bruijn et al. 2003), Y-GSP417, DBC2, Z108 (Métais et al. 2009), Y-GSP116 (Hartman et al. 2019), Z113 (Lindsay and Flynn 2016)

Table 9. Composition of the faunas found in Gözükızıllı-1a, −1b and Güvendik. Shown are the total number of first and second molars (M1/2) and the minimum number of individuals (N) as determined from the largest number of one of the dental elements. The cross indicates that no first or second molar has been found, but that the taxon is represented by another dental element

	GOZ-1a		GOZ-1b		Güvendik
	M1/2	N	M1/2	N	M1/2	N
Daxneria fragilis	9	4	7	3
Sayimys sp.			2	1
Eucricetodon	28	7	60	10	3	2
Bransatoglis cf. sjeni			1	1
Peridyromys / Microdyromys sp.			1	1
cf. Dipodidae gen. et sp. indet.			X	1
totals	37	11	71	17	3	2

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