Lycyaena dubia, Özkurt & Güleç & Erkman, 2015
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publication ID |
https://doi.org/ 10.3906/zoo-1407-38 |
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persistent identifier |
https://treatment.plazi.org/id/EB384D54-582E-5E43-FCAF-6801E535AD0F |
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treatment provided by |
Felipe |
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scientific name |
Lycyaena dubia |
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Species: Hyaenictitherium intuberculatum ( Ozansoy, 1965)
Locality: Hayranlı.
Age: MN11–12; Late Miocene, 9–7 Ma.
Material: 58-HAY-2/222, maxillary fragment with right I1–I3, P1–P4, and M1–M2, and left I2–I3, P1–P4, and M1–M2; 58-HAY-2/75, maxillary fragment with right I1–I3, C, and P1; 58-HAY-2/168, maxillary fragment with right P3 and P4; 58-HAY-2/101, maxillary fragment with right P4 and M1; and 58-HAY-2/57, maxillary fragment with left I–1–3.
Lİ1 B2 L 2 B2 L 3 B3 Lc Bc Lp1 Bp1 Lp2 Bp2 Lp3 Bp3 Lp4 Bp4 Lm1Bm1Lm1Lm2Bm2
tal
Figure 7. Comparison of the lower tooth measurements of Hyaenictitherium wongii from Hayranlı-Sivas and various localities.
Description
Maxilla: 58-HAY-2/75, maxillary fragment (right) I1–I3, C, and P1 ( Figure 8 View Figure 8 , 2c View Figure 2 ); 58-HAY-2/168, maxillary fragment (right) P3–P4 ( Figure 8 View Figure 8 , 2d View Figure 2 ); 58-HAY-2/101, maxillary fragment (right) P4–M1 ( Figure 8 View Figure 8 , 2e View Figure 2 ); 58- HAY-2/57, maxillary fragment (left) I–1–3 ( Figure 8 View Figure 8 , 2f View Figure 2 ); and 58-HAY-2/222, maxillary fragment (right) I1–I3, P1–P4, and M1–M2, and (left) I2–3, P1–P4, and M1–M2 ( Figure 8 View Figure 8 , 2a and 2b View Figure 2 ). The maxillary tooth specimens are in good condition, except for the left canine. The palate was originally well fossilized and the skull’s anterior orbits are missing, but the frontal part is intact. There is only a small depression because of the pressure applied to the nasal bones. The nasal space-hole is elliptic and vertical. The nasal space starting from the prosthion is in a vertical position up to the area behind the canine. The nasals are narrow and go to the anterior of the orbits. The infraorbital foramen lays on the contact point of P3 and P4. The orbits are round, extroversive front points that end between P3 and P4, and there is a postorbital narrowing ( Figure 8 View Figure 8 , 2a and 2b View Figure 2 ). The palate is short. From the prosthion to P2’s mesial has the same width, gets larger from the middle of P2 to the nuchal, and has its largest point in P4’s distal. M1 is settled through the buccal at a 90° angle in the distal of p4. The incisives are settled straight without overlapping; only the coronal parts are in contact. There is a small diastema between the canine and I3, and a smaller diastema between the canine and P1. The I3s have higher coronals than the other incisives. The cranial dimensions of H. intuberculatum are given in Figure 8 View Figure 8 .
Upper teeth: In the right tooth sequence, P1 and C are missing, and I3 is detached from its enamel. In the left tooth sequence, the carnassial is broken, and P1 and C are missing. Apart from these, the rest are in good condition.
C: The canine is strong, located through the tooth sequence, and is leaning slightly. It has the same position as the incisive. It is pressed up against by the buccal lingual, is strong, and there is a larger root than cusp and a round root coronal pass. This roundness is mesiobuccal. The distal crest, starting from the top of the canine and the crest in the mesial lingual, extends to the root coronal pass.
I1, 2: The incisive are located on a curvy line.
I3: Looks like a canine and there is certain cingulum process bending through the lingual.
P1: Round and small. There is very certain cingulum in the buccal. The coronal part of the lingual is flattened and there is a small crest starting at the mesial of the cingulum ending on the top.
P2: Is a short and large tooth. There is no supporting cusp in the anterior. It is based at the distal of the cingular eminence and there is a cusp in the posterior. There is a well-developed cingulum around the tooth.
P3: Is a relatively short and large tooth. In the distolingual, there is a cavity caused by a certain-strong cingular eminence. On the mesiolingual corner of the tooth, there is a small anterior cusp. There is a crest starting from this cusp going to the top. There is a little cusp in the posterior and a weak crest extends to the top from this cusp.
P4: The carnassial is lengthened and the metacone in the distal is sharpened. The metacone has a 2-sided shape in the buccal. The distal part of the paracone forms the highest point of the carnassial and it is higher than its part in the mesial, and in between there is a buccalingual sulcus. The protocone is lower than the metacone and paracone, and it is located through the lingual with an angle to the paracone on the top. The posterior edge of the protocone begins one-third of the way behind the paracone and the whole coronal part of the carnassial leans through the distal on the cingulum. The distal parts of the paracone and metacone form the cutter. There is a certain cingulum all around the tooth.
M1: Has a large and spandrel paracone having an eminentia to the labial. The protocone is small as the paracone is long and high. There is a curve from the paracone to the protocone and the metacone is located in the middle distal of this curve. M1’s lingual edge is in the proximal when compared with its buccal edge. Moreover, it is located in the proximal with P4 at a slight angle. The dental measurements of H. intuberculatum are given in Table 7.
Comparisons
This species was described from Sinap ( Turkey) by Ozansoy (1965) as Ictitherium intuberculatum and later transferred to Hyaenictitherium ( Bonis, 2004) , in Yassıören, with 4 maxilla and 5 mandible items ground on. Ozansoy (1965) did not indicate a holotype for this species and, because of this, Werdelin and Solounias (1991) determined the mandible given by Ozansoy (1965 Pl, 2:3) to be a lectotype. However, Bonis (2004) cited Ozansoy (1965) in his Paris MNHN collection, and by examining specimens TRQ 1192 maxilla (P1–M1), TRQ 1191 maxilla (P3–M2), TRQ 1012 maxilla (P2–M1), TRQ 1013 (C, P3–M2), and hemimandibular TRQ 1014 (C, P2–M1), he was able to evaluate the TRQ 1012, TRQ 1013, and TRQ 1014 specimens as holotype. Ozansoy (1965) defined H. intuberculatum ; the upper carnassial protocone is clear, is higher than that of Pikermi, Marageh I. hipparionum , and the protocone is in the anterior of the paracone. The first tuberculum of Ictitherium carnassials found in Sinap shows a bigger character. This type is more evolved than other Ictitherium . H. intuberculatum is bigger than I. robustum and smaller than I. hipparionum . Because of these data, H. intuberculatum shows a great difference from I. hipparionum of China, Greece, Iran, and Turkey. Werdelin and Solounias (1991) stated that this species’ dimensions and metric characters are very close to those of Polgardi I. pannocinum and might be conspecific to the Hungarian form, but there are some differences in terms of the rates of M2 and M1. Hence, the 2 species are not synonymous. This species’ approximate phylogenetic position is next to I. viverium Werdelin and Solounias (1991) .
Viranta and Werdelin (2003) recorded H. intuberculatum from Sinap when they found specimens AS.92.223, AS.93.63, S.91.653, S.91.654, S.91, Sc.1, and S.91.854. The comparison of the maxillary tooth sizes of Ozansoy (1965), mid Sinap and Viranta and Werdelin (2003), and Sinap with the 58-HAY-2-222, 58HAY- 2/75, 58-HAY-2/168, 58-HAY-2/10, and 58-HAY-2/ 57 specimens of Sivas-Hayranlı showed similarities. After the Paris MNHN collection, TRQ 1192 maxilla (P1-M1), TRQ 1191 maxilla (P3-M2), and TRQ 1012 maxilla (P2-M1) were seen by Özkurt; 58-HAY-2/222, maxillary fragment with right I1–I3, P1–P4, and M1–M2, and left I2–I3, P1– P4, and M1-M2; 58-HAY-2/75, maxillary fragment with right I1–I3, C, and P1; 58-HAY-2/168, maxillary fragment with right P3 and P4; 58-HAY-2/101, maxillary fragment with right P4, M1; and 58-HAY-2/57, maxillary fragment with left I-1-3 were included with the H. intuberculatum taxon. Comparisons of the upper tooth measurements of H. intuberculatum from Hayranlı-Sivas and various localities are given in Table 7 and Figure 9 View Figure 9 .
Family: Hyaenidae Gray, 1821
Genus: Ictitherium Wagner, 1848
Species: Ictitherium sp.
Locality: Hayranlı.
Age: MN11–12; Late Miocene, 9–7 Ma.
Material: 58-HAY-2/132, canine (lower right); 58- HAY-2/138, canine (lower right);
58-HAY-2/53, canine (lower left); and 58-HAY-2/252, part of isolated m1.
Description
Canine: Is strong, the posterior is curvy from the top of the corona, pressed from the sides, has a powerful root that is larger than the coronal part, and a round root coronal pass. There is a posterior crest that ends in the bulb of the distal part, which is not very clear. There is a certain mesial crest. The corona has a sharp and pointed top ( Figure 10 View Figure 10 , 5a, 5b, 5c View Figure 5 ). The canine measurements of Ictitherium sp. are given in Table 8.
m 1: Is considered as belonging to the Ictitherium species ( Figure 10 View Figure 10 , 5f View Figure 5 ).
Comparisons
58-HAY-2/132, 58-HAY-2/138, and 58-HAY-2/53 left canine samples are evaluated as Ictitherium sp. after being matched with Ictitherium specimens in morphological and morphometric comparisons. 58-HAY-2/252 isolated (right); it is clear that m1 belongs to Hyaenidae .
Genus: Lycyaena Schaub, 1941
Species Lycyaena dubia Zdansky, 1924
Lycyaena dubia sp. nov. Zdansky, 1924; 1981 Thalassictis (Lycyaena) sp. nov. - Solounias & De Beaumont, 1981; Thalassictis (Lycyaena) dubia (Zdansky) - Qiu, 1985; and Thalassictis (Lycyaena) dubia (Zdansky) - Werdelin, 1988.
Locality: Hayranlı.
Age: MN11–12; Late Miocene, 9–7 Ma.
Material: 58-HAY-2/151, maxillary fragment with left P1–P4 and M1; 58-HAY-2/152, maxillary fragment with left P3–P4 and M1.
Description
Maxilla: 58-HAY-2/152, maxilla left P3–P4 and M1 ( Figure 8 View Figure 8 , 3c); 58-HAY-2/151, maxilla left P1–P4 and M1 ( Figure 8 View Figure 8 , 3a and 3b). The right part of the specimen, except for the left part carrying canines P1–P 4 in the frontal, incisive, and dorsal parts of the skull, is lost. Has a higher skull than H. intuberculatum and H. wongii . The frontal part of the skull is relatively short ( Figure 8 View Figure 8 , 3a, 3b, and 3c).
Upper teeth: On the left tooth sequence, the metacones and paracones of P1–P4 are broken near the buccal.
P1: Round, small, very certain cingulum in the buccal; the coronal part in the lingual is flattened, and there is a small crest starting from the mesial side of the cingulum, ending on the top.
P2: Is a short and large tooth. There is no supporting cusp in the anterior. The coronal height is quite large. On the mesiolingual corner of the tooth’s anterior, there is a cingular eminentia, and there is a weak crest starting from the top of the cusp that becomes more defined near the cingulum. There is a cingulum all around the tooth, but it is stronger in the lingual.
P3: Has larger characteristics compared to the other P3s in the study. There is a large cavity in the distal of the tooth caused by a certain strong cingular eminence. On the mesiolingual corner of the tooth, there is a small cingular eminentia. A weak crest starting from the top of the cusp in the posterior goes through the cingulum. The cingulum is quite clear, especially in the labial.
P4: The carnassial is curved toward the dorsal side, and there is a certain cingulum, especially in the labial of the metacone. The protocone is located on the same plaque as the paracone in the buccalingual position, which means that the anteriors of the protocone and paracone start from the same point. The protocone is smaller, located in the paracone, and the paracone is smaller and located in the posterior part of metacone. There is a certain hole in the diagonal palate that was formed from the combination of the P4’s dorsal edge and m1. The location of m1 has an angle larger than 90° through the dorsal when compared to P4. The dental measurements of L. dubia are given in Table 9.
Comparisons
A comparison of the measurements of the specimens of 58-HAY-2/151 with L. dubia from China and L. chaeretis from Samos, which are in between those of L. dubia and L. chaeretis , shows that they are generally bigger than those of H. wongii ( Table 9). According to the size of the idems in the posterior ends of P4 and BP4 (anterior), L. dubia is a little larger than H. wongii and H. intuberculatum . L. dubia has a wider palate in between P4 but a shorter diastema, and comparatively has a bigger P4 according to 58-HAY- 2/ 151 L. dubia than the H. wongii 58-HAY-1996 Sivas Yüzey and H. intuberculatum 58-HAY-2/ 222 specimens.
In the structure of the P4 carnassial, the protocone is settled lower compared to the paracone, the metacone has 2 parts and is located at approximately a 90° angle compared to M1 and P4, and the enamel in the lingual of P3 is very clear. 58-HAY-2/151 P4 has a higher paracone than H. wongii and H. intuberculatum , M1 is wider and larger, and P3 is bigger and has a strong and recognizable crest from the bottom to the tip at the mesial side.
According to Werdelin (1988), the upper dentitions are fewer and less informative. The most marked feature is a tendency for the specimens of Lycyaena to have a narrow P4 blade, and 58-HAY-2/151 has this feature.
Specimens 58-HAY-2/151 and HAY-2/152 were compared with Miohyaena montadai , from Przeworno MF/1991/91 ( Heizmann and Kubiak, 1992) in the Polish National Academy of Science in Krakow, and, as a result, they were found to be different from M. montadai .
Werdelin (1988) showed that L. dubia is close to L. chaeretis from Samos and Pikermi, so close that they may be conspecific. However, until sufficient material has been recovered of the latter species, it is better to consider L. dubia a distinct species of Lycyaena , since L. dubia is the better known of these species.
Werdelin (1988) mentioned that L. dubia is close to L. chaeretis ; however, a comparison of the 58-HAY-2/ 151 specimens with Samos A. 4744- Lycyaena chaeretis and Zdansky, 1924 M. 3856- Lycyaena dubia shows that it more closely resembles L. dubia , especially because the skull height from a lateral view is the same. The most recognizable feature of Sivas 58-HAY-2/151 is the height of the skull. 58-HAY-2/151 has a higher skull than H. intuberculatum and H. wongii , and also higher than Samos A. 4744 L. chaeretis , and the frontal part of the skull is relatively short.
As a result of the morphological and morphometric comparisons of the study findings, it was decided that the specimens 58-HAY-2/151, a maxillary fragment with left P1–P4 and M1, and 58-HAY-2/152, a maxillary fragment with left P3–P4 and M1, should be included in the Lycyaena aff. (conf.) dubia taxon.
Family: Felidae Gray, 1821
Genus: Machairodus Kaup, 1833
Species: Machairodus giganteus (Wagner, 1848)
Locality: Hayranlı.
Age: MN11–12; Late Miocene, 9–7 Ma.
Material: 58-HAY-91/5, metatarsal III (right); 58-HAY-91/8, metatarsal II (right); 58 HAY-91/7, astragalus (right).
Description
Metatarsal III (right) ( Figure 10 View Figure 10 , 4a): The proximal joint face is spandrel and has a concave shape, leaving a groove in the middle. There is a very clear tubercle at the end of the surface where it connects with metacarpal 2, which has a sagittal orientation. The body has a slight curve, just like metacarpal 2. The trochlea, which is on the distal joint surface, is divided into 2 equal pieces, with a clear crest in the middle. The measurements of metatarsal III are given in Table 10.
Metatarsal II (right) ( Figure 10 View Figure 10 , 4b): The joint part of the proximal edge in the medial is broken, hence making an anatomic determination from this specimens not possible. The axle is slightly curved. The medial and lateral parts of the trochlea that are on distal edge are the same size and the crest between them is quite clear. The measurements of metatarsal II are given in Table 10.
Astragalus (right) ( Figure 10 View Figure 10 , 4c): Covers a larger area than the tibia surface of the lateral medialin. The additional surface of the lateral is quite smooth and is in a half-moon shape. The lateral surface is concave, while the medial surface is convex. The tuber calcanium is quite certain. Measurements of the astragalus are given in Table 10.
Comparisons
Bonis (1994) mentioned that a Machairodus aphanistus was found in the lower level localities of Kemiklitepe, which would indicate that it was from the Late Vallesian or Early Turolian. A skull and associate mandible from the upper level of Kemiklitepe is one of the most complete known specimens of M. giganteus UEK-124, recorded in the Late Miocene fauna of Kemiklitepe by Geraads et al. (2004). According to previous findings, M. giganteus was distributed in Anatolia in Late Miocene carnivore fauna.
The third metatarsal III of 58-HAY-91/5 is almost the same in dimensions to that of M. giganteus (AMPG PA 3256/91) from Pikermi (maximum length: 119.7 mm, distal breadth: 22.3 mm) ( Roussiakis, 2002).
In the Paris MNHN collection of PIKERMI, for Greek specimens of M. giganteus PIK 3278 metatarsal III of the proximal tip is broken, PIK 3279 metatarsal III of the proximal part, the PIK 3244 metatarsal III proximal tip is broken, the PIK 3240 metatarsal II proximal part is broken, the PIK 3280 metatarsal II proximal tip is broken, and they all have the same features as the 58-HAY-91/8 metatarsal II and 58-HAY-91/5 metatarsal III, and as the unpublished specimens of metatarsal III and metatarsal II of M. giganteus of Dr Koufos of the University of Thessaloniki, Greece.
In the Madrid MNCN (Madrid Natural History Museum) collection of Los Valles De Fuentidueña specimens of metatarsal II MNCN 65889, 68890, and 65891 of Machairodus alberdiae are almost the same in terms of their anatomy but they are quite small. A comparison of the maximum length shows the M. giganteus 58 HAY-91/8 metatarsal II as 133.7 mm and the MNCN M. alberdiae 65891 metatarsal II as 108.05 mm.
On the other hand, Machairodus aphanistus specimens from Batallones (Madrid) of B-1894 show metatarsal II (max length: 138.31 mm, distal breadth: 22.41 mm), metatarsal III (max length: 128.34 mm, distal breadth: 19.84 mm), B-707 metatarsal II (max length: 135.80 mm, distal breadth: 20.13 mm), metatarsal III (max length: 131.50 mm, distal breadth: 18.82 mm), B-7157 metatarsal II (max length: 142.70 mm, distal breadth: 26.40 mm), metatarsal III (max length: 141.80 mm, distal breadth: 19.57 mm), B-2695 metatarsal II (max length: 130.10 mm, distal breadth: 23.67 mm), and metatarsal III (max length: 128.00 mm, distal breadth: 19.23 mm). The average Batallones (Madrid) specimen has a metatarsal II maximum length of 136.72 mm, metatarsal III maximum length of 132.41 mm, metatarsal II distal breadth of 23.15 mm, and metatarsal III distal breadth of 19.36 mm. According to these measurements, M. giganteus is smaller than M. aphanistus but larger than M. alberdiae .
All of these comparisons show that 58-HAY-91/8 metatarsal II and 58-HAY-91/5 metatarsal III belong to the M. giganteus taxon. The 58-HAY-91/8 metatarsal II, 58-HAY-91/5 metatarsal III, and 58 HAY-91/7 astragalus were found together and were associated with each other, which is most probably because they belong to the same specimen and taxon.
The Madrid MNCN collection of Concud (Tervel) Amphimachairodus giganteus specimen of astragalus MHCN 14.669 has a maximum length of 51.90 mm, and the Batallones (Madrid) Machairodus aphanistus specimens B-6029, B-633, and B-2024 of the astragalus have maximum lengths of 66.38 mm, 61.00 mm, and 57.85 mm, respectively.
58 HAY-91/7 astragalus has the same dimensions as Concud Astragalus MHCN 14.669, but is smaller than Batallones (Madrid) Machairodus aphanistus B-6029, B-633, and B-2024.
The specimens of 58-HAY-91/7 astragalus were deemed to include the M. giganteus taxon according to the comparisons of the Concud (MN12) (Tervel) specimens of the astragalus MHCN 14.669 and unpublished specimens of the astragalus of M. giganteus of Dr Koufos of the University of Thessaloniki, Greece.
Order: Carnivora Bowditch, 1821
Locality: Hayranlı.
Age: MN11–12; Late Miocene, 9–7 Ma.
Material: 58-HAY-91/33, canine (upper left); 58-HAY- 2/130, canine (upper right); 58-HAY-2/55, distal part of the humerus; 58-HAY-2/244, distal phalanx; and 58-HAY- 2/54, medial phalanx.
Description
Canine: Is very strong, there is a certain and blade mesial ridge ( Figure 10 View Figure 10 , 5d, 5e View Figure 5 ). Has a larger and shorter corona compared to the other canines.
Humerus: Is a part of the humerus trochlea ( Figure 11 View Figure 11 , 6 View Figure 6 ). In the morphology of the trochlea, it is defined as carnivore indet.
Distal phalanx (claw) ( Figure 11 View Figure 11 , 7): The joint surface in the distal has a subversion. A quite clear sulcus can be seen from the plantar.
Medial phalanx ( Figure 11 View Figure 11 , 8 View Figure 8 ): The proximal edge of the medial phalanx is stable but the distal part is broken. The crista in the trochlea divides the joint surface into 2 pieces.
Comparisons
58-HAY-91/33, left upper canine; 58-HAY-2/130, right upper canine; 58-HAY-2/157, cranium; 58-HAY-2/55, humerus (distal piece); 58-HAY-2/244, distal phalanx; and 58-HAY-2/54, the medial phalanx specimen shows that these are carnivore, but they do not have a diagnostic characteristic to make a detailed taxonomic evaluation.
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