Ictitherium wongii, Özkurt & Güleç & Erkman, 2015

Özkurt, Şakir Önder, Güleç, Erksin & Erkman, Ahmet Cem, 2015, Carnivores from the Late Miocene locality of Hayranlı (Hayranlı, Sivas, Turkey), Turkish Journal of Zoology 39 (5), pp. 842-867 : 845-853

publication ID	https://doi.org/ 10.3906/zoo-1407-38
persistent identifier	https://treatment.plazi.org/id/EB384D54-5826-5E5B-FFD0-6D99E195AE35
treatment provided by	Felipe
scientific name	Ictitherium wongii
status

Treatment

Species: Hyaenictitherium wongii ( Zdansky, 1924)

1862–1867 Ictitherium hipparionum (Gervais) - p. 68, Pl. 12: 1–2; 1887 Palhyaena hipparionum (Gervais) - Kittl, pp. 333–335, Pl. 18: 2–7; 1924 Ictitherium wongii sp. nov. - Zdansky, pp. 73–84, Figs. 4–6, Pls. 14: 3–6, 15: 1–4, and 16: 1–2; 1925 Ictitherium hipparionum (Gervais) - De Mecquenem, p. 50, Pl. 9: 3; 0 1938 Ictitherium wongii Zdansky - Kretzoi, p. 113; 1938 Palhyaena hipparionum (Gervais) - Kretzoi, p. 113; 1939 Ictitherium hipparionum (Gervais) - Krokos, p. 160; 1952 Hyaenalopex atticus gen. et sp. nov. - Kretzoi, p. 21; 0 1980 Palhyaena wongii (Zdansky) - Howell and Petter, pp. 584 and 588; 1980 Ictitherium hipparionum (Gervais) - Koufos, pp. 56–65,

Figs. 18–19, Pls. 7: 2–3, 8: 1; 1981 Thalassictis wongii (Zdansky) - Solounias pp. 71–74, Fig. 17; 1985 Thalassictis mesotes sp. nov. - Kurten pp. 81–82, Figs. 1–2; 1988 Thalassictis wongii (Zdansky) - Werdelin, pp. 223–230, Fig. 9 View Figure 9 ; 1989 Hyaenotherium magnun gen. et sp. nov. - Semenov, pp. 94–105, Figs. 28–31; 1989 Hyaenotherium wongii (Zdansky) - Semenov, pp. 105–118; Werdelin & Solounias 1991: 33; and Bonis 1994: 21.

Locality: Hayranlı.

Age: MN11–12; Late Miocene, 9–7 Ma.

Material: 58- HAY - 1996 Sivas Yüzey. Complete skull with right I1–I3, P2–P4, and M1–M2, and left I1–I3, P2– P4, and M1–M2; 58-HAY-2/187, maxillary fragment with left I1–I2; 58-HAY-2/51, mandibular fragment with left I1, c, and p2–p4, and right m1–c and p1–p3; 58-HAY-2/243, right mandibular fragment with c and p2–m2; 58-HAY- 2/223, right mandibular fragment with c and p2–p3; and 58-HAY-2/102, right mandibular fragment with p3 and p4.

Description

Skull: 58 HAY 1996 Sivas Yüzey ( Figure 2 View Figure 2 , 1a and 1b). The skull is clear but it has been displaced from its original position because of the sediment pressure of the nasal bones, from cranial to caudal. Due to the pressure applied from the top, the frontal area, which connects to the dorsal nasal bones, is crushed through to the inside, while the sagittal crest on the caudal skull is completely intact. The zygomatic arches are broken. The posterior part of the skull is protected; mastoids and occipitals are in their original position. The right P1 and C are missing and I3 is displaced from its enamel in the tooth arrangement. On the left, the carnassial composition is broken, and P1 and C are missing. The nasal aperture is in an elliptic and vertical position. The nasal cavity starting from the prosthion remains vertical up to the middle of the canine. The snout is narrow and ends at the anterior or the orbits. The front part of the skull bones ends in the middle of the orbits. The infraorbital foramen lies on the contact points of P3 and P4. The orbits are oval, the front parts end between P3 and P4, and there is visible postorbital narrowing. The zygomatic arches are strong-thick and have a tubercle through the buccal. The cranial part is relatively small and round. The sagittal suture starts at the front of the postorbital bulges and forms the crest by connecting in the middle of the neurocranium and ending in the acrocranion. The occipitals are strong, and the occipital line has a bulge through the posterior and connects with the sagittal crest coming from the dorsal at the hind of the occipital condyle. The condyles are relatively weak and are laterally connected to the occipital bones. The foramen magnum is robust and round. The postglenoid tubercles are large. The bullae are relatively small and elliptical shaped, and the front part is a little behind that of the front surfaces of the postglenoid region. The palate is short and relatively narrow. The width of the palate is parallel until P2, where it then narrows through the buccal and hits its largest point in the distal part of P4. The incisive are lined up and touch each other, but do not overlap. There is a small diastema between the canine and I3, and a smaller diastema between the canine and P1. Specimen 58-HAY-2/187 ( Figure 2, 2 View Figure 2 ) is a nasal piece of bone formed by I1 and I2. The cranial dimensions of H. wongii are given Table 2.

Upper teeth: In the right tooth sequence, P1 and C are missing, and I1 is displaced from its enamel. In the left tooth sequence, the carnassial is broken, and P1 and C are missing. The rest are in good condition.

I1, 2: Incisives are based on a curvy line, and there is a cingulum torus in the elliptic cusp and buccal.

I3: Looks like a canine and there is a clear tuber cingulum bending to the lingual.

P2: Short and large. There is no supporting cusp in the mesial. There is a small cingular tuber in the mesiolingual corner of the anterior of the tooth, connected by a crest, starting from the top of the main tubercle. The cingular is based in the distal of the torus and there is a well-raised cusp in the distal. There is a well-raised cingulum around the tooth.

P3: Relatively short and large. There is a large cavity in the lingual half of the tooth at the distal end, caused by a cingular eminence. On the mesiolingual corner of the tooth, there is a small anterior cusp. In the posterior, there is a large small cusp.

P4: The carnassial is in a long and narrow structure. There is a protocone formed along and apart from the parastyle. The mesial edge of the protocone is a little behind the anterior edge of the parastyle. The parastyle is long and a little smaller than the paracone. The sharp part of the metacone is relatively long and curves toward the buccal. There is a certain strong lingual cingulum.

M1: Long, triangular, and there is a paracone tubercle through the labial. The metacone is small, while the protocone is long and high.

M2: Is a very small tooth with an elliptic cusp.

The upper tooth measurements of the H. wongii specimens are given in Table 3.

From the Greco-Iranian province, comparisons of the teeth of the most common hyaenid H. wongii with L. dubia , L. chaeretis , and T. hyaenoides are given in Figure 3.

Mandible: 4 mandible remains belonging to the H. wongii species from Hayranlı localities. 58-HAY-2/51, (left) I1, c, and p2–p4, and (right) m1, c, and p1–p3 ( Figure 2 View Figure 2 , 3); 58-HAY-2/243, (right) c and m2–M2 ( Figure 2 View Figure 2 , 4); 58-HAY-2/223, (right) c and p2–p3 ( Figure 2 View Figure 2 , 5 View Figure 5 ); and 58- HAY-2/102, (right) p3 and p4 ( Figure 2 View Figure 2 , 6 View Figure 6 ) are in the list. The symphysis is relatively lengthened, narrow, and has a certain lean toward the back. Many of the mandibular rami are missing, broken, or lost. The ramus is round in shape and the anterior part starts beneath m2. The mandibular corpus is quite thin. It evolves from anterior to posterior and m1 leans to the interior. There is only 1 fossa beneath p2. The mandibular dimensions of H. wongii are given in Tables 4 and 5.

0.08

0.06

0.04

0.02

–0.02

–0.04

–0.06

–0.08

–0.1

–0.12 LP1 LP2 LP3 LP4

LM1 BP1 BP2 BP3

Lycyaena dub a, 58-HAY-2/151, Turkey, Present Study

Lycyaena dub a, Ch na, Werdel n, 1988

Lycyaena dub a, Ch na, Zdansky, 1924

Lycyaena chaeret s, NHMW-A.4744, Samos, 2009 (or g. meas.)

Lycyaena chaeret s, Greece, P lgr m, 1931

hvalass ct s hyoeno des, Ch na, Werdel n 1988

Hyaen ct ther um wong, 58-HAY-1996, Turkey, Present Study

Hyaen ct ther um wong, Turkey, Gerva s, 1859

Hyaen ct ther um wong, Greece, Koufos, 2009

Figure 3. Logarithmic ratio diagram comparing the teeth of L. dubia , L. chaeretis , T. hyaenoides , and H.

wongii from various localities.

Mandibular teeth: Four different mandibles are used in the study and all of the specimens are in good condition, except for the incisive.

c: The canine is strong and the root is larger than the petal. The root is round at the connection point in between the root and the petal. This roundness extends to the distal crest, starting from the top of the mesiolingual and canine.

p1: Has a very small, single-rooted, and rounded petal.

p2: There is no secondary mesial tubercle and it is lengthened through p3. The posterior secondary tubercle is strong and based in the distal part of the cingulum. The posterior secondary has a powerful lingual and a weak buccal cingulum.

p3: There is an anterior cuspid that is relatively short and large, and is not totally evolved. The posterior cuspid is small and there is a large distal cingular bulge that looks like a talonid. There is a well-developed buccal cingulum.

p4: Short and large, with a strong anterior cuspid and a very strong posterior cuspid based in the distal cingular position. The lingual level of the distal cingular structure is high and it has a crest form similar to the entoconid. There is a well-evolved buccal cingulum.

m1: Has a relatively small, narrow, and short talonid. The metaconid is relatively half the size of the protoconid. The trigonid length is two-thirds the length of the tooth. The protoconid and paraconid have a blade-like edge and are separated by a deep carnassial notch. The hypoconid is strong and based in the center of the talonid. The entoconid is smaller and combined with the hypoconulid by a crest covering the distal limit of the talonid. A crest starting from the top of the hypoconid increases in size and faces the metaconid and protoconid where they join. A well-evolved cingulum makes the tooth prominent. The lower tooth measurements of the H. wongii specimens are given in Table 6.

Comparisons

The taxonomy of small- to medium-sized Late Miocene Hyaenidae is complex. These hyaenids are known as ‘ictitheres’ and have a long taxonomic past ( Kurten, 1982; Semenov, 1989; Werdelin and Solounias, 1991; Koufos, 2000; Bonis, 2005). In Greece, in the Late Miocene, ictitheres have been found in various localities, such as Pikermi, Samos, Axios Valley, Perivolaki (Thessaly), and Kerassia (Evia Island) ( Koufos, 2006). Among the Pikermi ictitheres, there is a species that is smaller than Adrocuta eximia and bigger than Ictitherium viverrinum , which was for a long time called Ictitherium hipparionum . The generic name of this species was changed to Palhyaena , Thalassictis , Ictitherium , and Hyaenotherium . The name Palhyaena hipparionum was given to a specimen by Gervais (1859) in Mt. Luberon, France, and the type sample of this taxon was placed in the Avion museum, but the type specimen has since been lost ( Kurten 1982). This hyaenid is nomen dubium and can only be included in the taxon defined by the type specimen of Mt. Luberon until the Mt. Luberon type specimen is found ( Werdelin, 1988; Werdelin and Solonuias, 1991).

Similar hyaenids were defined as Ictitherium wongii , which has characteristics between those of Adrocuta eximia and Ictitherium viverrinum from the Late Miocene and from China ( Zdansky, 1924). After these, similar European materials were included with these species. As a result, these hyaenids were included in the genus Hyaenotherium , with Hyaenotherium wongii as the type species of the genus ( Semenov, 1989). Thus, another medium-size hyaenid, called Ictitherium hyaenoides in China by Zdansky, was transferred to the Hyaenictitherium by Kretzoi (1938). These 2 hyaenids should be included in the same genus even though they are different at the species level, and because Hyaenictitherium has priority, these 2 species were included in the genus Hyaenictitherium ( Zhang et al., 2002; Bonis, 2004, 2005). Therefore, Late Miocene hyaenids between Adcrocuta eximia and Ictitherium viverrinum are known as Hyaenictitherium wongii ( Zdansky, 1924) .

Similar Hyaenidae from Turkey were recorded from Elmadağ - I. robustum and I. hipparionum ( Şenyürek, 1960) ; Çobanpınar - I. hipparionum and H. eximia ( Ozansoy, 1965) ; Kemiklitepe - H. wongii , A. examina , and L. chaeretis (Bonis, 1994) ; Sinap - H. intuberculatum and H. wongii ( Viranta and Werdelin, 2003) ; and Akkaşdağı - H. wongii , A. eximia , and I. viverrinum ( Bonis, 2005) . All of these I. hipparionum records from Turkey are considered as H. wongii because it is a valid taxon.

58-HAY - 1996 Sivas Yüzey. Complete skull with right I1–I3, P2–P4, and M1–M2, and left I1–I3, P2–P4, and M1– M2; 58-HAY-2/187, maxillary fragment with left I1–I2 compared with Greek specimens in the Aristotle University of Thessaloniki Mytilini-1A: MTLA-200 and MTLA-1; Maragheh specimens in the Natural History Museum of Paris: MAR 3353, MAR 3354, MAR 3355, MAR 3356, and MAR 3357; Samos specimens in the Natural History Museum of Vienna: NHMW-SAM-A.4743, NHMW-SAM-A.4745, NHMW-SAM-A.4746, and NHMW-SAM-A.4749 Mar-0019 and Mar-0019/1 (A4803). Comparisons of the cranial dimensions and upper tooth measurements of H. wongii from Hayranlı-Sivas and various localities showed that the specimens belong to the H. wongii taxon ( Tables 2 and 3; Figures 4 and 5).

58-HAY-2/ 51 specimens, mandible fragment with left I1, c, and p2–p4, and right m1–c, p1–p3; 58-HAY-2/243, right mandible fragment with c and p2–m2; 58-HAY- 2/223, right mandible fragment with c and p2–p3; and 58-HAY-2/102, right mandible fragment with p3 and p4. Fifty-one specimens compared with Greek specimens in the Aristotle University of Thessaloniki: MTLA-2, MTLA-237, MTLA-266, MTLA-468, MTLB-1, MTLB-100 ,

300

250

200

150

100

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29

MAX MAX MAX MAX MAX MAX MAX MAX MAX MAX MAX MAX MAX MAX MAX MAX MAX MAX MAX MAX MAX MAX MAX MAX MAX MAX MAX MAX MAX

Figure 4. Comparison of the cranial dimensions of Hyaenictitherium wongii from

Hayranlı-Sivas and various localities.

MTLB-171, MTLC-18, LGPUTRZ1-2, LGPUTRZO-10, and LGPUTVAT-100; Thessaloniki specimens in the Natural History Museum of Paris: MNHNSLQ-929 and MNHNSLQ-930; and Samos specimens in the Natural History Museum of Vienna: NHMWSAMA.4745, MHNWSAMA.4743, NHMWSAMnn-1, and NHMWSAMnn-2. This showed that these mandible specimens belong to the H. wongii taxon. Comparisons of the mandibular dimension and lower tooth measurements of H. wongii from Hayranlı-Sivas and various localities are given Table 4 and Figures 3, 6, and 7.