catalogue of the holotypes of fossil land mammals in the...

Georgian National Museum

Catalogue of the Holotypes of Fossil Land Mammals

in the Georgian National Museum, Tbilisi

By

Maia Bukhsianidze

March 2015

ISBN 978-9941-9380-5-4

2015 Georgian National Museum

Introduction

Since 2005 efforts have been made to consolidate and organize fossil collections kept in the National Museum of Georgia and the Institute of Palaeobiology (now part of the GNM) through different international and national projects, with different extent of curation tasks: RHOI project - Upper Miocene of Georgia (2005-2007); GNSF/ST 1-5/23 (2010-2012); Pleistocene NSF project #BCS-1019408 (2011-2013); Volkswagen project #85 820 2 (2010-2014), SNF IZ73Z0 152380 (2012-2014), Rustaveli foundation project #11/05 (2012-2015); SNF IZ73Z0 127940 (2014-2016). This catalogue is one of the outcomes of these conducted works.

Some explanations for the information included in the catalogue:

New and old collection numbers are indicated for most of the specimens: the first is the current accession number and the second is the original number (in parenthesis), e.g.: 20-2013/128 (# 35) for the holotype of Crocuta abessalomi Gabunia, 1958.

Only figures from the first publication of the type specimen of each taxon are indicated.

Derivation of name is indicated if name is given to honor a place of discovery or individual.

Diagnosis published in languages other than English, French and German are translated. The latest version of a diagnosis is given a preference.

Synonyms are not listed.

All the photos are by David Tsvariani, except of holotypes of Udabnopithecus garedziensis and Homo georgicus that are photgraphed by Malkhaz Machavariani.

This is a dynamic document, and will be updated if the some type specimen which was not found yet will be found. Also it is quite possible that some of the holotypes are not listed here. If you notice this the author will be deeply thankful if you let her know about it.

The catalogue is freely available on the Georgian National Museum web-site.

The author is thankful to all the GNM team members who helped during the collection organization process. Special thanks to Prof. Dr. R.-D. Kahlke for advice to make this catalogue.

Order Hyracoidea Huxley, 1869

Family Pliohyracidae Osborn, 1899

Genus Kvabebihyrax Gabunia & Vekua, 1966

Type species of the genus Kvabebihyrax kachethicus Gabunia & Vekua, 1966

Derivatio nominis: “Kvabebihyrax” - The site Kvabebi is located on the slope of the Kvabebi Mountain in South East Georgia.

Diagnosis (Vekua, 1972, p. 98): Skull relatively high with a short muzzle. Orbits very small, significantly raised above the frontal surface and directed laterallys. Sagittal crest moderately developed. Zygomatic arches high, widely diverging, with two disto-lateral projections. Lower jaw deep, with shortened premolar part of the mandible body and deep symphyseal fossa. First upper incisors possibly are permanently growing, other incisors and canines morphologically similar to premolars. Premolars are almost completely molarised. First lower incisor with a wide shovel-like crown; second - canine like, permanently growing; third - completely reduced. Canine similar to premolars, Cheek teeth starting from the third premolar are mesohypsodont or even hypsodont.

Species Kvabebihyrax kachethicus Gabunia & Vekua, 1966

Pl. I figs. 1, 2.

Holotype: 29-2013/299 (K-10), lower jaw with complete tooth row.

First publication: Gabunia & Vekua, 1966; pp. 643-647, figs. 1-3.

Type locality and age: Kvabebi, Georgia; Late Pliocene, ca. 3 Ma.

Derivatio nominis: “kachethicus” - Kakheti is a region in the East Georgia. The site Kvabebi is in Kakheti.

Diagnosis (Vekua, 1972, p. 98): the same as for the genus.

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Order Rodentia Bowdich, 1821 Family Castoridae Hemprich, 1820

Genus Dipoides Jaeger, 1835

Species Dipoides eldaricus Bendukidze & Burchak-Abramovich, 1991

Holotype: E-9, upper P4/ and M1/ of the same individual.

First publication: Bendukidze & Burchak-Abramovich, 1991; pp. 148-151, fig. 3.

Type locality and age: Eldari, Azerbaijan; Late Miocene, Upper Sarmatian.

Derivatio nominis: “eldaricus” - Eldari is the site where fossil was found.

Diagnosis (Bendukidze & Burchak-Abramovich, 1991, p. 148): Small size representative of the genus with parastria on the P4/, rudimentary para- and metastrias in a form of weak

grooves on M1/. Mesoflexia on the M1/ closes into mesofacet approx. at the half of the unworn tooth height. Among upper teeth the “S” shape pattern is present only on molars. Hypostriae as well as mesostria on P4/ don’t reach the crown base. Crown bases closed, yet roots are hardly noticeable. Hypostriid reaches crown base on lower teeth, while parastriid gradually disappears at about 1/3 of the tooth height.

* Specimen not found.

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Genus Stenofiber Geoffroy Saint-Hilaire, 1833

Species Stenofiber caucasicus Burchak-Abramovich & Gabashvili, 1980

Pl. II figs. 1-3.

Holotype: 32-2013/1075 (269-53), fragment of a hemi-mandible dex. with canine (basal part), P/4-M/2.

First publication: Burchak-Abramovich & Gabashvili, 1980; pp. 20-35, figs. 1-4.

Type locality and age: Udabno, Georgia; Late Miocene, Upper Sarmatian-Meotian.

Derivatio nominis: “caucasicus” - after Caucasus - a region between the Black and the Caspian seas, Udabno site is in the Caucasus region.

Diagnosis (Burchak-Abramovich & Gabashvili, 1980, p. 26): mesial part of the P/4 crown is rounded and constricted; its apex is slightly turned mesially. Buccal sinus on the occlusal surface directed abruptly downwards and distally. The same synus on the buccal wall is observable along the whole height of the crown, it divides the tooth intwo two lobes: on larger mesial and smaller distal. On the occlusal surface arc-like mesial and central enamel folds with distally directed convexities. Distal enamel fold almost straight. On the M/1 and M/2 all enamel folds perpendicular to mesio-distal axis of the crown, almost parallel to each other. Mesial enamel fold turned mesially in its buccal half. Buccal synuses on the occlusal surface of M/1 and M/2 are more centrally located i.e. close to the coronal plane of the crown compared to P/4.

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Order Proboscidea Illiger, 1811 Family Gomphotheriidae Hay, 1922

Genus Platybelodon Borissiak, 1928

Species Platybelodon jamandzhalgensis Beljaeva & Gabunia, 1960

Pl. III fig. 1.

Holotype: 20-2013/221 (5/23-5/24) incomplete hemi-mandible dex. with DI/2, DP/2-DP/4.

First publication: Belyaeva & Gabunia, 1960; pp. 88-92, fig. 12; pl. II, figs. 1, 2.

Type locality and age: Belomechatskaia, Russian Federation; Middle Miocene, Chokrak.

Derivatio nominis: “jamandzhalgensis”- Jaman-Dzhalga is a name of a gully near the village Belomechetskaia.

Diagnosis (Beljaeva & Gabunia, 1960, p. 47): lower deciduous tusks abruptly widen at the crown base, with a crenulated lateral edge. Oblique ridges on deciduous premolars are strongly developed.

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Family Elephantidae Hay, 1922

Genus Mammuthus Brookes, 1828

Species Mammuthus meridionalis (Nesti, 1825)

Subspecies Mammuthus meridionalis taribanensis Gabunia & Vekua, 1963

Pl. IV figs. 1-3.

Holotype: incomplete skeleton, K/13.

First publication: Gabunia & Vekua, 1963; p. 68, figs. 6-7, 9-13; pl. I-VI.

Type locality and age: Taribana, Georgia; Early Pleistocene, Apsheronian Stage, Olduvai normal polarity subchron.

Derivatio nominis: “taribanensis”- Taribana steppe, place where the fossil was found.

Diagnosis (Gabunia & Vekua, 1963, pp. 21-22): Taribana elephant is characterized by rather large size (by the size it is second only to the Nogaisk elephant). Skull low, top of the head noticeably inclined caudally; edge of the top of the head in a frontal view semi circular shape; occipital convex with well developed protuberances; frontals wide, less constricted above the nasal cavity than in other elephants from the group (ratio of the frontal width at the constriction to the max width of occipital is 45); supraorbital processes well developed, bent downwards and backwards; nasal cavity of a moderate height with slightly sloping edges; nasal process turned upwards and abruptly delimited from the frontal plane; zygomatic relatively low and thin (the min. depth of the zygomatic to the length from the sinciput to the mesial edge of the alveolus is 0.5); dorsally the surface between alveolus has a shape of conus; articular condyle of the mandible is clearly separated from the mandibular body by the narrow neck.; mandibular body surface is tangibly convex. Last upper molar (M3/) is composed by 12 plates, excluding talon. Plate folding - insignificant. Plate frequency - 4, enamel thickness varies between 3.7-5.0 mm. M/3 is composed by 11 plates, excluding talonid. Plate frequency and enamel thickness is the same as for the M3/.

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Order Primates Linnaeus, 1758. Family Hominidae Gray, 1825

Genus Udabnopithecus Burchak-Abramovich & Gabashvili, 1945

Type species of the genus: Udabnopithecus garedziensis Burchak-Abramovich & Gabashvili, 1945.

Derivatio nominis: “Udabnopithecus” –from the name of the site Udabno.

Diagnosis: no formal diagnosis.

Species Udabnopithecus garedziensis Burchak-Abramovich & Gabashvili, 1945

Pl. V figs. 1-10.

Holotype: 156-192, fragment of a maxilla dex. with P4/ and M1/

First publication: Burchak-Abramovich & Gabashvili, 1945; pp. 451-464, figs. 1-7.

Type locality and age: Udabno, Georgia; Late Miocene, Upper Sarmatian-Meotian.

Derivatio nominis: “garedziensis” - Garedjis udabno (literal translation from Georgian - desert of Gareji) is the name of the area where Udabno fossil site is located.

Diagnosis: no formal diagnosis.

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Genus Homo Linnaeus, 1758

Species Homo georgicus Gabunia, Lumley, Vekua, Lordkipanidze & de Lumley, 2002

Pl. VI figs. 1-4.

Holotype: D2600, almost complete mandible.

First publication: Gabunia et al., 2002, pp. 243-253. fig. 1.

Type locality and age: Dmanisi, Georgia; Early Pleistocene, 1.76 Ma.

Derivatio nominis: “georgicus” – Georgia – a name of the country in the Southern Caucasus.

Diagnosis (Gabunia et al., 2002, pp. 251-252): diagnosis in French.

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Order Carnivora Bowdich, 1821 Family Felidae Waldheim, 1817

Genus Machairodus Kaup, 1833

Species Machairodus davitashvili Vekua, 1972

Pl. VII fig. 1.

Holotype: 29-2013/453 (K-14), skull.

First publication: Vekua, 1972, pp. 80-92, figs. 11-12, pl. IV figs. 1-3.


Derivatio nominis: “davitashvili” – in honor of the palaeontologist Prof. Dr. Leo Davitashvili, director of the Institute of palaeobiology, Tbilisi.

Diagnosis (Vekua, 1972, p. 80): Large saber-toothed cat, with rather long, narrow and relatively low skull (basal length of the skull is 257 mm); zygomatic arches massive, weakly diverging; sagittal crest strongly developed, lower jaw relatively deep and massive; symphysis deep; diastema long and strongly concave; upper canine long, flat, curved, with crenulated cutting edges; upper and lower second premolars absent, premolars significantly reduced; M/1 without a metaconid.

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Family Hyaenidae Gray, 1821

Genus Crocuta Erxleben, 1777

Species Crocuta abessalomi Gabunia, 1958

Pl. VIII figs. 1-3.

Holotype: 20-2013/128 (# 35), incomplete hemi-mandible sin. with P/2, P/3 and M/1.

First publication: Gabunia, 1958; pp. 249-252, fig. 1.

Type locality and age: Belomechatskaia, Russian Federation; Middle Miocene, Chokrak, MN6.

Derivatio nominis: “abessalomi” – in honor of the palaeontologist Dr. Abesalom Vekua, who being a young student accompanied Leo Gabunia to the excavations in Belomechetskaia.

Diagnosis (Gabunia, 1973, p. 27): one of the smallest representatives of the genus (length of M/1 varies between 19-22 mm). Skull of a moderate height, muzzle relatively short, abruptly narrows mesially. Raising branch of the lower jaw strongly inclined posteriorly. Premolars narrow; upper and lower first premolars are completely absent; accessory cuspules on the premolars are weakly developed; upper carnassial (P4/) with a very low, rudimentary protocone, lower carnassial (M/1) without or with insignificant rudiment of a metaconid.

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Species Crocuta miriani Meladze, 1967

Pl. IX figs. 1-3.

Holotype: 21-2013/1201 (B-40), skull.

First publication: Meladze, 1967; pp. 31-34, pl. III figs. 1,2.

Type locality and age: Bazaleti, Georgia; Late Miocene, Pontian.

Derivatio nominis: “miriani”– in honor of the geologist Dr. Mirian Dzvelaia, deputy director of the Institute of Palaeobiology, Tbilisi.

Diagnosis (Meladze, 1967, p. 31): medium size Crocuta, upper P4/ is long (MD length) with strongly reduced, mesially directed protocone and a very long metastyle, Upper P1/ and M1/ absent.

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Genus Ictitherium Wagner 1848

Species Ictitherium ibericum Meladze, 1967

Holotype: B-22, skull.

First publication: Meladze, 1967; pp. 25-31, pl. I figs. 5, 6.


Derivatio nominis: “ibericum” - Iberia a name of an ancient kingdom in the Southern Caucasus, centered on present-day Eastern Georgia.

Diagnosis (Meladze, 1967, p. 25): Ictitherium similar in size to I. robustum. Sagittal crest strongly developed; orbuts round. I3/ much larger than I2/; P1/ with one root, without accessory cuspules, width of M1/ is almost twice as large as its length, M2/ is of a medium size, it has an almost triangular shape, similar to M1/ its width is almost twice as large as its length. Talonid length of the lower M1/ is ¼ of the length of the tooth. Second phalanges are asymmetrical. Third phalanges partially retractile type.


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Family Ursidae Gray 1825

Genus CephalogaleJourdan 1862

Species Cephalogale meschethense Gabunia, 1964

Pl. X figs. 1-3.

Holotype: 3-2013/-65 (7/411), M/1 sin.

First publication: Gabunia, 1964; pp. 38-42, fig. 13, pl. I figs. 4-5.

Type locality and age: Benara, Georgia; II fossiliferous layer, Late Oligocene.

Derivatio nominis: “meschethense” - Meskheti is a region in south of Georgia where Benara site is located.

Diagnosis (Gabunia, 1964, p. 38): small size canid (L. Gabunia considered it to belong to the Family Canidae, MB). Lower carnassial has combination of traits characteristic to archaic cynodonts on the one hand and to more specialised cephalogales on the other hand: trigonid is relatively law, yet cusps are still rather closely placed to one another; paraconid with an accessory inner cuspid, metaconid almost entirely fused with protoconid and slightly shifted to the distal edge. Hypoconid is rather high, of a cutting type; entoconid low, crenulated; talonid wide and relatively elongated (index of its length 42).

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Family Canidae Fischer, 1817

Genus Canis Linnaeus, 1758

Species Canis tengisii Vekua, 1962

Pl. XI figs. 1-6.

Holotype: 18-2012/1215 & 18-2012/1216 (Akh-214, 664, 681, 684, 883, 907), skull fragments and a hemi-mandible dex.

First publication: Vekua, 1962; pp. 30-37, pl. II figs. 1-3.

Type locality and age: Akhalkalaki, Georgia; end of the Early Pleistocene.

Derivatio nominis: “tengisii” – in honor of the geologist Dr. Tengiz Lazarashvili, discoverer of the Akhalkalaki site.

Diagnosis (Vekua, 1962, p. 30): Relatively very small size Canis with primitive upper and lower premolars (with one cone/conid). No diastema between premolars. Length of molars to length of premolars ratio for the upper cheek teeth is relatively low. Zygomatic arches diverge relatively weakly. This form is characterized by constricted premolar section of the muzzle, as a result premolars are placed along one straight line. Diastema between cheek teeth and canine is relatively short. Mandible relatively not deep and thin; premolars with one conid, placed immediately after each other. Limb bones very small and gracile with some characteristic morphological features.

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Family Mustelidae G. Fischer de Waldheim, 1817

Genus Promephitis Gaudry, 1861

Species Promephitis brevirostris Meladze, 1967

Holotype: B-1207, complete mandible.

First publication: Meladze, 1967; pp. 22-25, fig. 11, pl. I fig. 4.


Diagnosis (Meladze, 1967, p. 22): Promephitis with extremely shortened muzzle – length of lower molars (L M/1-M/2) exceeds half of the length of the entire tooth row (Length I-M/2). Two incisors on each side on the mandible. Distal edge of the mandible is not straight, condyle above the tooth row level.


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Family Amphicyonidae Haeckel, 1886

Genus Amphicyon Lartet, 1836

Species Amphicyon caucasicus Gabunia, 1973

Pl. XII figs. 1-3.

Holotype: 20-2013/110 (#77), hemi-mandible dex. with I/2, I/3, /C, P/3-M/2.

First publication: Gabunia, 1973; pp. 19-26, fig. 3, pl. I fig. 1.

Type locality and age: Belomechatskaia, Russian Federation; Middle Miocene, Chokrak, MN6.

Derivatio nominis: “caucasicus” - after Caucasus - a region between the Black and the Caspian seas, Benara site is in the Caucasus region.

Diagnosis (Gabunia, 1973, p. 19): Large amphicyon with completely reduced P/1 and P/2. Lower carnassial moderately elongated, with relatively weakly detached metaconid. M/2 with noticeably narrowed talonid.

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Order Perissodactyla Owen, 1848 Family Hyracodontidae Cope, 1879

Genus Ardynia Matthew and Granger, 1923

Species Ardynia plicidentata Gabunia, 1964

Pl. XIV figs. 1-5.

Holotype: 3-2013/45 (7/78), M2/ sin.

First publication: Gabunia, 1964; pp. 74-77, fig. 36, pl. II fig. 5.

Type locality and age: Benara, Georgia; Late Oligocene

Diagnosis (Gabunia, 1964, p. 74): M2/ small (crown length 18.7 mm, width 18.9 mm). Middle valley deep, framed by 5-6 small folds buccally and distally; distal valley is completely closed by the distal wall of the crown connecting ends of ectoloph and metaloph. Ectoloph

strongly concave buccally, yet gets noticeably bent labially near its distal edge. Cingulum distinctly expressed only mesially and lingually, yet its clear traces are visible distally as well.

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Genus Benaratherium Gabunia, 1955

Type species of the genus Benaratherium callistrati Gabunia, 1955

Derivatio nominis: “Benaratherium” – Benara is a village where site is located.

Diagnosis (Gabunia, 1964, p. 83): The smallest representative of indricotheriids (M/3 MD length - 60 mm, P/3-M/3 length - 249.4 mm) with a combination of Indricotherium and Paraceratherium traits. Mandibular body deep, molars relatively law crowned. Upper molars, especially M3/ characterised by strong development of parastyle and weakly shaped paracone and metacone. Distal branch of hypolophid on P/3 and P/4 reaches lingual edge of the tooth. Mesial part of lower premolars is significantly narrowed (mesial edge of P/3 is slightly sharpened as in Paraceratherium). Metalophid on premolars is significantly raised above hypolophid. Distal branch of metalophid on molars forms open, almost straight angle with its labial branch. All teeth with strongly developed cingulum. Limb bones relatively gracile (index max. width to length for the lunate bone – 70; index of proximal width to max. length for mc III - 26.3) and straight, with a distinct inclination towards intensification of the function of the third digit.

Species Benaratherium callistrati Gabunia, 1955

Pl. XIV figs. 1-3.

Holotype: 3-2013/1 (7/15), hemi-mandible fragment dex. with P/3-M/3.

First publication: Gabunia, 1955; pp. 177-182. fig. 1, 2.

Type locality and age: Benara, Georgia; Late Oligocene.

Derivatio nominis: “callistrati” – in honor of the geologist Dr. Calistrate Gabunia, father of Leo Gabunia.

Diagnosis (Gabunia, 1964, p. 83): the same as for the genus.

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Family Rhinocerotidae Gray, 1820

Type species of the genus: Meschotherium meschethicum Gabunia, 1964

Derivatio nominis: “Meschotherium” – Meskhi (Lat. Meschi) is the name of people living in Meskheti, region of Georgia.

Diagnosis (Gabunia, 1964, p. 67): medium size rhinocerotid, characterised by rather weakly developed anterior transverse ridge and significant inside sloping of outer walls of trigonid and talonid on lower molars. Transverse ridges on molars obliquely placed towards the tooth axis, their lingual edges incline mesially.

Genus Meschotherium Gabunia, 1964

Species Meschotherium meschethicum Gabunia, 1964

Holotype: 7/42, M/1 sin.

First publication: Gabunia, 1964; pp. 67-70, fig. 29, pl. II fig. 2.


Derivatio nominis: “mescheticum” – Meskheti (Lat. Meschethi) is a region in south of Georgia where Benara site is located.


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Genus Diceros Gray, 1821

Species Diceros gabuniai Tsiskarishvili, 1987

Pl. XV fig. 1-4.

Holotype: 32-2013/1115 (352-1), almost complete skull.

First publication: Tsiskarishvili, 1987; pp. 30-38, pl. I fig. 1, pl. II fig 1, pl. III fig. 1.

Type locality and age: Eldari-2, Azerbaijan; Late Miocene, Upper Sarmatian.

Derivatio nominis: “gabuniai” - after the Georgian palaeontologist Prof. Dr. Leo Gabunia.

Diagnosis (Tsiskarishvili, 1987, p. 30): Large rhino (max. length of a skull - 620 mm), skull is low (skull height in the occipital region - 132 mm) and wide (max. width in the region of zygomatic arches - 333 mm), caudal edge of the naso-maxillary notch - at the level of the limit between P3/ and P4/; anterior edge of the orbit - at the level of the limit between M2/ and M3/; caudal edge of chaonas - at the level of posterior edge of M2/. Teeth brachyodont, with weak layer of cement, croche present on all the teeth, anticroche is absent, crista present only on P2/ and P3/. DP2/ and P2/ are weakly molarised. On the P/3 anterior closed valley is formed. Strongly reduced P/1 is still present. Temporal ridges are not developed.

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Genus Chilotherium Ringström, 1924

Species Chilotherium eldaricum Tsiskarishvili, 1987

Pl. XVI figs. 1, 2.

Holotype: 32-2013/1109 (352-15), mandible.

First publication: Tsiskarishvili, 1987; pp. 52-59, figs. 3-4, pl. X fig. 2.

Type locality and age: Eldari-2, Azerbaijan; Late Miocene, Upper Sarmatian.

Derivatio nominis: “eldaricum” - Eldari steppe is the name of the area were the Eldari site is located and where the specimen comes from.

Diagnosis (Tsiskarishvili, 1987, p. 52): mandible of medium size rhino, with well developed mandibular angle projecting downwards; distal edge of the body of the mandible convex, anteriorly bending upwards; symphysis rather massive, mesially almost unfused and not widened, turned somewhat upwards; incisors (I/2) powerful, strongly curved upwards (almost unworn regardless the fact that the mandible belongs to a rather adult individual), completely covered by enamel, cross-section is triangular across entire length, teeth subhypsodont with shallow mesial grooves, and with weakly developed cingulum.

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Genus Dicerorhinus Gloger, 1841

Species Dicerorhinus vekuai Tsiskarishvili, 1987

Pl. XVII figs. 1-5.

Holotype: 29-2013/298 & 29-2013/299, skull fragment and hemi-mandible dex.

First publication: Tsiskarishvili, 1987; pp. 81-91, pl. XXVI figs. 1 - 2, pl. XXVII figs. 1 - 3.


Derivatio nominis: “vekuai”– in honor of the palaeontologist Dr. Abesalom Vekua.

Diagnosis (Tsiskarishvili, 1987, p. 81): Medium size rhino (smaller then D. megarhinusand D. jeanvireti, close to D. etruscus) with a rather narrow skull. Lower jaw relativelywide, body of the mandible not deep; ramus of the mandible is low, short, significantly inclined backwards. Mandibular angle – talon is completely undeveloped. Upper teeth hypsodont (P4/ - 118), relatively stretched in width, croche on premolars with multiple blades, accessory valley is present; on upper P2/ lingual cingulum is absent, on P3/ and P4/ it’s present only on the posterior half, lingually; crista absent on M/1 and M2/, croche is large, sometimes with two blades; metacone rib absent on maxillary teeth.

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Family Chalicotheriidae Gill, 1872

Genus Schizotherium Gervais, 1879

Species Schizotherium chucuae Gabunia, 1951

Pl. XVIII figs. 1-3.

Holotype: 3-2013/81 (7/7), incomplete hemi-mandible dex.

First publication: Gabunia, 1951; pp. 279-284, figs. 1-3.


Derivatio nominis: “chucuae” – in honor of the geologist Dr. Mariam (Maro) Khuchua, the discoverer of the Benara site, Schizotherium remains were the first fossils found there.

Diagnosis (Gabunia, 1964): medium size Schizotherium, characterised by weak development of metastylid, incomplete fusion of metastylid and hypoconid, noticeable reduction of M/3 talonid and complete absence of cingulum on molars, relatively gracile limb bones.

* Specimen damaged.

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Family Equidae Gray, 1821

Genus Hipparionde Christol, 1832

Species Hipparion eldaricum Gabunia, 1959

Pl. XIX figs. 1, 2.

Holotype: 32-2013/47 & 32-2013/6 (139-85), fragmentary skull.

First publication: Gabunia, 1959; pp. 121-133, pl. V fig. 4.


Derivatio nominis: “eldaricus” - Eldari is the site where fossil was found.

Diagnosis (Gabunia, 1959, pp. 121-122): Large size hipparion: skull basal premolar length (P2/ – basion) – 307 mm, length P2/-M3/ - 145-152 mm. Skull possibly narrow, ratio of linguo-buccal width of a maxilla to skull basal premolar length - 34.2. Tooth row short: ratio of mesio-distal length of tooth row to skull basal premolar length - 47.2 mm. Upper molars are large relative to premolars: molar/premolar ratio is 86.7; one infraorbital fossa, very short and shallow: index of its location to the orbit is ca. 150 and to the facial crest ca. 285.7. Swelling of diastema is well observable; its ratio is ca. 70. Protocone short and wide: index of its length - 25.2-39.5 on weakly worn P3/-M2/, while for relatively heavily worn teeth 30.1-35.7; correspondingly protocone shape ratios are 43-80.2 and 64.2-90.1. Enamel folding on upper teeth is moderate: on the distal wall of a mesial fosset and on the mesial wall of the distal fosset on moderately and heavily worn P3/, P4/ and M1/ there are in average 4-6.5 and 9 folds. Premolars and molars are low crowned: index of their height to MD length on P3/, P4/ - 162, on M1/-M3/ 180.2 - 191, and on P/3, P/4 - 200. Double loop of a hipparion type. Buccal groove of lower molars is deep, accessory elements are weakly developed. Calyx on I/3 occupies only mesial part of the crown. Limb bones moderately massive: ratio - distal width to length for mc III is 17-18.1; for mt III - 15.7. Metapodials relatively short: index mt/t 67.1. lateral digits are well developed: average ratio values of dorso-palmar depth of distal ends of mc II, IV to mc III is 83.4; of mt II, IV to mt III – 77.2.; index of average length of first phalanges of lateral digits to dorso-palmar length is 57.6. Limb bones evidently were strongly bent in joints. Middle hooves are narrow: index of its width to dorso-palmar length is 88.3.

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Species Hipparion garedzicum Gabunia, 1959

Pl. XX figs. 1-3.

Holotype: 32-2013/1003 (156/13), skull.

First publication: Gabunia, 1959; pp. 137-145, pl. VI fig. 1.

Type locality and age: Udabno, Georgia; Late Miocene, Upper Sarmatian/Meotian.

Derivatio nominis: “garedzicum” – Garedjis udabno (literal translation from Georgian is: desert of Gareji) is the toponym of the area where Udabno fossil site is located.

Diagnosis (Gabunia, 1959): Large size hipparion: skull basal lenght is about 457 mm., length P2/-M3/ – 145 mm. Muzzle long: ratio of orbital width to muzzle length - 67. Frontal region narrow: index of its width - ca. 39.5, fronto-basal indicator is about 253. Tooth row short: index of its length to basal length - ca. 30.8; diastema/tooth row index - 72.5. Upper molars large relative to premolars: molar/premolar ratio -84.5. Praeorbital fossa is one, deep, short and low: ratio of its depth to length - 52, ratio of its location to the orbit is ca. 119 and to the facial crest ca. 97. Nasal notch small: its caudal edge does not reach and falls far from the P2/. Protocone is short and wide: ratio of its length - 30-31.8 on weakly and somewhat worn M1/-M2/; correspondingly index of the protocone shape is 42.3-46. Enamel folding on upper teeth is moderate: on the distal wall of a mesial fosset and on the mesial wall of the distal fosset on moderately worn P2/-P4/ and M1/, M2/ is 7-8 folds. Teeth are low crowned: index of height to MD length on P3/ is ca. 198. Double loop of a hipparion type. Buccal grove of lower teeth is deep. Calyx on lower i3 occupies almost entire width of the crown. Lateral digits are moderately developed: index of dorso-palmar depth of distal end of mc II to mc III is 73. Limb bones possibly were strongly bent in joints: distal phalanxes of third digit are narrow: index of its width to dorso-palmar length 88.

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Species Hipparion urmiense Gabunia, 1959

Pl. XXI figs. 1, 2.

Holotype: 148/191, maxilla dex.

First publication: Gabunia, 1959; pp. 166-176, pl. VI figs. 2, 3.

Type locality and age: Urmia - Surroundings of the village Kurtevul 50 Km east of the LakeUrmia, Iran;Early Pliocene.

Derivatio nominis: “urmiense” – the Lake Urmia, the fossil comes form the surroundings of the lake.

Diagnosis (Gabunia, 1959, p. 166): Medium size hipparions: although the length of upper tooth row indicates rather large size (length P2/-M3/ – 147 - 155 mm) other skull dimensions contradict this and doubtlessly indicate medium size hipparion. Skull possibly narrow: frontal width at the outer edges of orbits - ca. 130 mm; braincase width at the postorbital constriction - ca. 61 mm; width of nasal bones (together) ca. - 85 mm. Teeth

apparently relatively large. Upper molars relatively small: ratio - molar row length to premolar row length - 82.1. Praeorbital fossa either is completely absent or is present in a form of a hardly noticeable rudimentary lacrymal fossa, which is very much shifted posteriorly and upwards. Protocone belongs to the group of short and wide ones, yet, it is distinguished by somewhat more elongated (elliptical) shape: index of its MD length on P3/-M2/ with weak and medium wear is 26 - 36.9 (mean 30.1); on those with heavy wear 26.2 - 42 (mean 31.5); protocone shape index 41.1 - 66.5 (mean ca. 50) and 56 - 71.5 (mean 64.4) correspondingly. Folding of the enamel on upper teeth is weak: on the posterior wall of mesial fosset and on the mesial wall of the posterior fosset on moderately and heavily worn P3/, P4/ and M1/, M2/ - in average 2 - 3.5 to 5 folds. Teeth moderately hypsodont: index of tooth height to MD length for P3/, P4/ is 230 - 246.1: for P/3, P/4 and M/1, M/2 it varies between 234 - 258. Double loop in general is of a hipparion type, however it reveals tendency to the cabbaloid type in some cases. Buccal groove of lower teeth is shallow. Accessory elements are weakly developed. Calyx on I/3 occupies entire width of the crown. Limb bones gracile: ratio of the distal width to length for mc III is ca. 12.8; the same ratio for mt III is 11.7. Lateral digits are moderately developed: index of the dorso-palmar depth of the distal end of the mc lateral (mc IV) to the mean of the distal dorso-palmar depth of the mc III is 64.1. The same for the distal end of the mt II to mt III is 61.4. Ratio of the length of the first phalanx of the lateral digit to the mean length of the first phalanx of the middle digit is for the fore limb - 50, and for the hind limb - 47.4. Limb bones were weakly bent in joints: dorsal surface of tibia distally is weakly bent; dorsal fossae above the distal guiding roller are shallow. Distal phalanges of the third digit are wide: ratio of the width to its length (DP length) for fore limbs is about 123, for hind limbs 96.1 and 111.8.

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Gnus Equus Linnaeus, 1758

Species Equus hipparionoides Vekua, 1960

Pl. XXII figs. 1-3.

Holotype: 18-2012/1733 (Akh-99), lower tooth row P/2-M/2 sin.

First publication: Vekua, 1960; pp. 1417-1420, fig. 2.


Diagnosis (Vekua, 1962, p. 79): Relatively small and rather specialised representative of equids. Upper cheek teeth are distinguished by very short protocone (index of its length is 23.3 - 36.5); lower permanent cheek teeth are distinguished by development of parastylid, hypostylid and ectostylid, the latter is relatively weakly developed. Metapodiums are very gracile (ratio of the proximal width of mc III is 19.6, the same ration for MT III has the following values 16, 10.2, 14.7).

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Order Artiodactyla Owen, 1848 Family Anthracotheriidae Leidy, 1869

Genus Anthracotherium Cuvier, 1822

Species Anthracotherium kwablianicum Gabunia, 1964

Pl. XXIII figs. 1-6.

Holotype: 3-2013/43 (7/310) – M2/ dex. and 3-2013/63 (7/317) – M/3 dex.

First publication: Gabunia, 1964; pp. 137-148, figs. 62, 63b, pl. X figs. 2, 3.


Derivatio nominis: “kwablianicum” – riv. Kvabliani near the Benara site.

Diagnosis (Gabunia, 1964, p. 137): A very large anthracothere (length of the M/3 is 66.6 mm). It is characterised by presence of strongly developed mesial and disto-buccal projections of the cingulum on the P4/ and clear pillar on the mesial projection of the cingulum. Mesial crest of a protoconule of the upper molars joins the mesial crest of a paracone. Metastyle relatively strongly developed, mesostyle not divided, accessory cuspules present in the middle of mesial and distal edges of upper molars and in front of metaconide of the lower third molar. Talonid of the lower third molar has a complex structure.

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Family Eltelodontidae Lydekker, 1883

Genus Paraentelodon Gabunia, 1964

Type species for the genus Paraentelodon intermedium Gabunia, 1964

Diagnosis (Gabunia, 1964, pp. 109-110): A very large entelodon (molar row length ca. 325 mm) with rather powerful canines and robust metapodiums. Apex of incisors is somewhat pointed, with hardly noticeable lingual cingulum. Upper canines relatively very large, with two roots, and weakly expressed mesial and distal ridges. First upper premolar is conical, noticeably enlarged with clear mesial and hardly noticeable distal cingulum. Third upper premolar is very wide and relatively short with weakly pronounced ribs. Fourth – rounded square shaped, on the lingual side the tooth is somewhat compressed mesio-distally. First upper molar semi-bunodont with 6 main cusps and one accessory cusp located behind the middle cusp of the distal cusp row. Third upper molar with significantly narrowed distal part, which possess weakly developed disto-lingual cusp (hypocone). Third lower premolar relatively very high, crown apex is clearly shifted distally, and possess clearly distinct accessory cusplet at the base of the distal edge of the tooth crown, above the cingulum. Lower molars with relatively high distal pairs of cusps, and weakly developed entoconid. Crown apexes are rounded, non-faceted. Cingulum is weakly or moderately developed on premolars as well as on molars. Third metacarpal is very robust, with divided facet on the proximo-medial protuberance and significantly concave palmar surface of the

bone. Phalanges are robust as well, with pronounced rugosities in the upper part on the dorsal surface and with deep medial fossae for the attachment of ligaments.

Species Paraentelodon intermedium Gabunia, 1964

Pl. XXIV figs. 1-29, Pl. XXV figs. 1-14.

Holotype: incomplete upper tooth row: 3-2013/22 (7/181) - P1/ dex., 3-2013/19 (7/188) – M2/ dex., 3-2013/27 (7/189) – M3/ dex., 3-2013/23 (7/184) - P1/ sin., 3-2013/24 (7/185) – P3/ sin., 3-2013/25 (7/186) – P4/ sin., incomplete lower tooth row: 3-2013/28 (7/194) – P/3 sin., 3-2013/29 (7/192) – M/2 sin., 3-2013/30 (7/195) – M/3 sin., 3-2013/31 (7/193) – M/3 dex. and manus dex.: 3-2013/32 (7/184) – mc III, 3-2013/33 (7/186) – mc IV, 3-2013/34 (7/181) - phalanx prox., 3-2013/35 (7/185) – phalanx prox. of the same individual.

First publication: Gabunia, 1964; pp. 109-133, figs. 56, 58, pl. VIII, 1-6, pl. IX, 1-5.


Diagnosis (Gabunia, 1964, pp. 109-110): the same as for the genus.

* P3/ dex., M2/ sin. of the figured type tooth row (Gabunia, 1964, fig. 56) not found.

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Family Suidae Gray, 1821

Subfamily Kubanochoerinae Gabunia, 1958

Genus Kubanochoerus Gabunia, 1955

Type species for the subfamily and genus Kubanochoerus robustus Gabunia, 1955

Derivatio nominis: “Kubanochoerus” – riv. Kuban – vil. BelomechetskaIa is located at the bank of the river.

Diagnosis (Gabunia, 1973, pp. 76-77): Largest among ever known suids. Skull relatively law, snout long. Males, at least, possess large unpaired hornlike outgrowth on the frontal bone and pair of small outgrowths above postorbital appendages of frontals. Neurocranium is strongly bent downwards. Parietal surface is very narrow. Temporal fossae deep. Zygomatic arches apparently very wide, mesially they continue into strongly developed facial lateral plate-like crests. Incisor part of the praemaxilla relatively wide. Mandible long with abruptly concave symphyseal part and rather important diastema between canine and second premolar. Dental formula I3/3; C1/1; P4/3; M3/3. Lower canines are of a “verrucosus” type. Premolars except of upper P1 are relatively large. Second and third premolars, especially the lower ones, visibly preserve cutting character. P/4 has one major cuspid with one apex. Molars are low-crowned, with four major, well developed accessory cusps/cuspids.

Species Kubanochoerus robustus Gabunia, 1955

Pl. XXVI fig. 1.

Holotype: 20-2013/213 (# 33), mandible.

First publication: Gabunia, 1955; p. 1263, fig. 1.

Type locality and age: Belomechatskaia, Russian Federation; lower fossiliferous horizon, Middle Miocene, Chokrak.

Diagnosis (Gabunia, 1973, pp. 76-77): the same as for the genus.

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Family Hippopotamidae Gray, 1821

Genus Hippopotamus Linnaeus, 1758

Species Hippopotamus georgicus Vekua, 1959

Pl. XXVII figs. 1-29, Pl. XXVIII figs. 1-35

Holotype: articulated manus dex. - 18-2012/585 (Akh. 1), radius dex.; 18-2012/586 (Akh. 2), piramidale dex.; 18-2012/587 (Akh. 3), lunatum dex.; 18-2012/588 (Akh. 4), scaphoideum dex.; 18-2012/589 (Akh. 5), trapezoideum dex.; 18-2012/590 (Akh. 14) mc V dex.; 18-2012/591 (Akh. 12) mc IV dex.; 18-2012/592 (Akh. 9), mc IIIdex.; 18-2012/593 (Akh. 6), mc II dex.; 18-2012/594 (Akh. 15), I phalanx (v digit) dex.; 18-2012/595 (Akh. 13), I phalanx (iv digit) dex.; 18-2012/596 (Akh. 10), I phalanx (iii digit) dex.; 18-2012/597 (Akh. 7), I phalanx (ii digit) dex.; 18-2012/598 (Akh. 16), II phalanx (v digit) dex.; 18-2012/599 (Akh. 11), II phalanx (iii digit) dex.; 18-2012/600 (Akh. 8), II phalanx (ii digit) dex.; 18-2012/601 (Akh. 18) astragalus sin.

First publication: Vekua, 1959; pp. 561-656, figs. 1, 2.


Derivatio nominis: “georgicus”- from the country name Georgia.

Diagnosis: a very large and peculiarly specialized representative of hippos. Characterized by relative elongation of the third digit and corresponding changes in carpal and possibly in tarsal bones as well, confirming intensification of the function of this digit.

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Family Leptomerycidae Zittel, 1893

Genus Lophiomeryx Pomel, 1853

Species Lophiomeryx benarensis Gabunia, 1951

Pl. XXIX figs. 1-3.

Holotype: 3-2013/54 (7/10), hemi-mandible dex. fragment with P/4-M/3.

First publication: Gabunia, 1951; p. 142.

Type locality and age: Benara, Georgia; III fossiliferous layer, Late Oligocene.

Derivatio nominis: “benarensis” – vil. Benara in the southern Georgia, where Benara site is located and where the fossil was found.

Diagnosis (Gabunia, 1964, pp. 167-168): Relatively small representative of the genus with a well developed cingulum on the upper molars (cingulum borders with the entire lingual side of the crown) and with clear accessory cuspules on the buccal groove of the lower molars. On the buccal side of the metacone of the upper molars rib is weakly expressed, skewness of crescents is moderate. Lower molars are relatively narrow; talonid mesio-distally elongated, with a shape of almost a crescent. Lower premolars with rather well developed middle ridge, but with open disto-lingual valley. Mesial ridge of the premolars is rather well developed. Distally the middle metapodials with a dorsal ridge.

* Specimen damaged, only P/4 with a fragment of mandibular body is preserved.

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Genus Prodremotherium Filhol, 1877 (synonym of Leptomeryx Leidy, 1853)

Species Prodremotherium trepidum Gabunia, 1964

Holotype: 7/11, incomplete hemi-mandible dex. with P/4-M/2.

First publication: Gabunia, 1964; pp. 175-179, figs. 80 a, b.

Type locality and age: Benara, Georgia; III fossiliferous layer, Late Oligocene.

Diagnosis (Gabunia, 1964, p. 175): The smallest representative of the genus (length of P/4-M/2 is 25.6 mm, max length of the metatarsal – 115 mm). P/4 with a moderately developed mesio-lingual ridge, but with two closed lingual valleys in the distal part of the tooth. M/1 and M/2 with closed mesio-lingual valley and accessory cuspules in the buccal valley. III and IV metatarsals are completely fused. Sagittal ridges of the distal articular surface don’t reach dorsal side.


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Family Giraffidae Gray 1821

Genus Karsimatherium Meladze, 1962

Type speceis of the genus Karsimatherium bazalethicum Meladze, 1961

Derivatio nominis: “Karsimatherium” – from the vil. Karsimaant-kari, Bazaleti site is located in this village.

Diagnosis (Meladze, 1967 pp. 76-77): Large representative of Sivatheriinae. Skull short and high, with vast pneumatic sinuses in the skull roof. Braincase rised above the facial region. Muzzle short, in the mesial part strongly shortened and raised in the mesial part. Postorbital part of the skull is narrower than the facial part. Females have a pair of ossicone protuberances, located on the posterior part of the braincase, at the limit of temporal, parietal and occipital bones. Supposedly males possessed at least one pair of ossicones located in the same area as the protuberances in females. Temporal fossa very deep, delimited by strong ridge from the above. Orbits relatively large, with elongated shape, not

projecting. Teeth, especially premolars, are very large, robust, enamel relief - rugose. Palate becomes abruptly narrow mesially, its width between P2/ is half of the width between M3/. Limb bones somewhat elongated.

Species Karsimatherium bazalethicum Meladze, 1961 (=Sivatherium bazalethicum in Meladze, 1961)

Pl. XXX figs. 1-3.

Holotype: 21-2013/1274 (B-20), almost complete skull.

First publication: Meladze, 1961; pp. 164-166.


Derivatio nominis: “bazalethicus” – from the Lake Bazaleti, which is near the site.

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Family Moschidae Gray, 1821

Genus Iberomeryx Gabunia, 1964

Type speceis of the genus Iberomeryx parvus Gabunia, 1964

Derivatio nominis: “Iberomeryx” – from Iberia, the name for a kingdom of the Southern Caucasus, centered on present-day Eastern Georgia.

Diagnosis (Gabunia, 1964, p. 179): The smallest representative among known representatives of deer (length of the M1/-M3/ 15.6 mm), with extremely brachyodont molars and with a clear palaeomeryx fold on lower molars. Upper molars with strongly developed cingulum on the mesial and especially on the mesio-lingual sides of the crown, cingulum becomes thicker at the mesio-lingual corner of the crown and is shaped as a cuspule. Parastyle and mesostyle are projecting abruptly. Mesio-buccal rib on the upper third molar is projecting in a similar way as parastyle and mesostyle. Disto-buccal rib is weak. Mesial branch of the mesio-lingual crescent completely fuses with parastyle, distal crescent joins the buccal part of the mesial branch of the distal crescent, the latter fuses with the mesostyle lingually. On the mesial crescent of the M3/ a well expressed fold extends from the apex of the protocone in the bucco-distall direction. This fold is less evident on other molars. Lingual wall is strongly inclined buccally (angle between the lingual wall and its

perpendicular plane which is passing through the crown base is 30). Lower premolars are strongly constricted laterally, with a closed posterior valley. Molars with a rudimentary, yet clearly distinct palaeomeryx fold, which joins buccally the corner of the mesio-buccal crescent and possesses signs of a cingulumon some specimens, which is bordering the almost entire buccal side of the crown. Mesio-lingual valley is widely open lingually and is divided into mesial and distal parts by a rather deep transverse groove, which is passing mesially to the protoconid. Distal valley in the apex of the tooth is lingually open as well, however after even a weak wear it closes. Metaconid in a form of a pillar; entoconid bucco-lingually constricted, which after wear becomes an elongated loop-shaped area. Radius and ulna not fused, distal end of a tibia with a relatively deep and narrow grooves for articulation with astragalus.

Species Iberomeryx parvus Gabunia, 1964

Holotype: 7/98 – fragment of a maxilla dex. with M1/-M3/ and fragment of a hemi-mandible dex. with P/4-M/2, most likely of the same individual.

First publication: Gabunia, 1964, pp. 179-187, figs. 82-83.



* Specimens not found.

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Family Cervidae Gray, 1821

Genus Paradicrocerus Gabunia, 1959

Type species for the genus Paradicrocerus flerovi Gabunia, 1959

Diagnosis (Gabunia, 1959, p. 114): small dicrocerini with antlers that possess 5 spikes. They are diverging bush-like form the triangular base of the antler. Coronet is low. Antlers are directed upwards and somewhat laterally. Antler spikes are constricted from the sides.

Species Paradicrocerus flerovi Gabunia, 1959

Pl. XXXI figs. 1, 2.

Holotype: 20-2013/50 (5/11), left antler.

First publication: Gabunia, 1959; pp. 114-117, figs. 1, 2.

Type locality and age: locality Puket, Belomechetskaia, Russian Federation; Lower fossiliferous horizon, Middle Miocene, Chokrak.

Derivatio nominis: “flerovi” – in honor of the palaeontologist Prof. Dr. Konstantin Flerov.


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Genus Dicrocerus Lartet, 1837

Species Dicrocerus salomeae Gabunia, 1955

Pl. XXXII fig. 1.

Holotype: 41-2013/140 (5/13), left antler.

First publication: Gabunia, 1955; pp. 359-360, fig. 1 on the insert of the p. 256.

Type locality and age: Arkneti, Georgia; Late Miocene, Pontian.

Derivatio nominis: “salomea” – dedicated to Salome Gabunia, the daughter of Leo Gabunia.

Diagnosis (Meladze, 1967, p. 70): Dicrocerus of medium size. Antlers with two tines. Difference among posterior and anterior tines is relatively weakly expressed. Both tines represent in a way a continuation of the coronet. They bifurcate in a rather considerable

distance from a rather well developed coronet and diverge with a small angle (ca. 25). Coronet relatively short.Teeth with low crowns. Well developed buccal accessory pillars are present of each lower molar. Cingulum is rather strongly developed on molars mesially. Palaeomeryx fold is present. Distal lobe of the lower M/3 is obliquely placed (it is directed buccally relative to the tooth axis).

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Genus Arvernoceros Heintz, 1968

Species Arvernoceros insolitus Vekua, Bendukidze & Kiladze, 2010

Pl. XXXIII fig. 1.

Holotype: D2774, almost complete left antler.

First publication: Vekua et al. 2010; pp. 43-45, figs. 3, 4.


Diagnosis (Vekua et al. 2010, p. 43): Large size deer, with massive, relatively moderately long antlers, very widened towards the apex, with several spikes. There is only one accessory first tine (brow tine, MB), placed immediately above the coronet. Antler is directed somewhat outwards, it turns abruptly upward at the level of the brow tine and widens shovel-like to the apex (palm with several spikes, MB) in a form of concave plate.

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Genus Cervus Linnaeus, 1758

Species Cervus abesalomi Kahlke, 2001 (=Cervus perrieri (Croizet & Jobert) in Vekua, 1995, p. 119, tab. 39, fig. 1 ; = Cervus (Dama) cf. nestii (Major) in Vekua, 1995, p. 124, tab. 44, fig. 3.

Pl. XXXIV fig. 1.

Holotype: D1495, complete antler.

First publication: Kahlke, 2001; pp. 43-45, figs. 3, 4.


Diagnosis (Kahlke, 2001, p. 475): diagnosis in German, see the publication.

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Family Bovidae Gray, 1821

Genus Mirabilocerus Gadzhiev, 1961

Type species of the genus: Tragocerus flerovi eldaricus Gabashvili, 1956

Diagnosis (Meladze, 1967, p. 92): Antelopes of medium to large sizes. Postcornual part is well developed; its length exceeds the length of frontals. Parietal ridges are weakly developed or absent. Frontals concave between the horn-cores. Orbits insignificantly projecting laterally. Basioccipital somewhat widened caudally; auditory bullae not large, strongly constricted from the sides. Posterior edge of the palatine is behind the posterior edge of the M3/. Praeorbital fossae present. In the region of crista pterigoidea frontals are not in contact with temporal bones. Horn cores are located above the orbits, their anterior surface somewhat in front of the anterior edge of orbit; horn cores are of a medium length or long, but always shorter than the basal length of the skull. Horn cores present both in males and females. On the anterior surface of the horn cores in males there are 2-3 demarcations (steps). Horn core cross section varies along the horn core, anteriorly it can be: sharp (one keel), flattened (two keels), or rounded (no keels), however two keel segment is always present. In females demarcations of the anterior edge is significantly less expressed or even absent; flattening of the anterior surface in females is similar to males; it is present either along the entire length or just in the apical part of the horn core. First incisors asymmetrical, stretched distally with sharpened distal corners, other incisors are noticeably smaller than the first incisors, also asymmetrical, and somewhat bent distally. Basal pillars are not always present on upper molars, while they are always present on lower ones. P2/ and P3/ with a trend to develop a pair of lingual crescents. P/4 even if it has a lingual wall, it’s formed as a result of a closure of a posterior valley.

Species Mirabilocerus eldaricus Gabashvili, 1956 (=Tragocerus flerovi eldaricus Gabashvili, 1956, = Mirabilocerus azerbajanicus Gadzhiev, 1961)

Holotype: 139-70, horn core dex.

First publication: Gabashvili, 1956; p. 20, pl. I, fig. 5.


Derivatio nominis: “eldaricus” - Eldari is the site where fossil comes from.

Diagnosis (Meladze, 1967, p. 93): parietal ridges are weakly developed. Horn cores rather long, somewhat shorter than the basal length of a skull. Antero-posterior diameter of the horn core reduces more or less gradually towards the apex. Horn cores are present in both males and females. Horn core anteriorly has three demarcations in males, that are dividing horn cores into four sections: the first basal one has a sharp edge (one keel), third section has rounded edge (no keels), and remaining sections have flattened edge (two keels); in some cases anterior edge is flattened along entire horn core length. Female horn cores are deprived of demarcations and their anterior edge is flattened along entire length. In every case keels join into one keel near the apex.


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Species Mirabilocerus brevicornis Meladze, 1967

Pl. XXXV figs. 1-4.

Holotype: 41-2013/67 (5/30), braincase with horn cores.

First publication: Meladze, 1967; pp. 93-98, fig. 21, pl. XXIX fig. 2.


Diagnosis (Meladze, 1967, p. 93): medium size mirabilicerus. Temporal crests extremely weak. Horn cores of a medium length, longer than a half of the basal length of the skull. Antero-posterior diameter of the horn core at the base is the smallest among known mirabiloceruses, gradually decreasing towards the apex. Both sexes have horns. Horn cores in males have two demarcations (steps) on the anterior surface, with sharpened (one keel) anterior edge; towards the apex anterior surface is flattened (two keels). Horn core cross-section gets circular. In females two noticeable demarcations are observable on the anterior edge of the horn cores, section below the demarcations is with one keel, and surface above is flattened, towards the apex becomes rounded. Accessory pillars on the upper molars most of the time are present, on lower ones they are always present.

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Species Mirabilocerus maius Meladze, 1967

Pl. XXXVI figs. 1-4.

Holotype: 21-2013/710 (B-555), almost complete skull.

First publication: Meladze, 1967; pp. 98-102, fig. 22, pl. XXVI figs. 1, 2; pl. XXVII figs. 1, 2.


Diagnosis (Meladze, 1967, p. 98): Large mirabilocerus. Parietal crests practically absent. Horn cores long, yet evidently somewhat shorter than the basal length of the skull. Horn-core dimensions in the basal half is very large, antero-posterior diameter abruptly diminishes towards the apex. Both, males and females possess horn cores. Male horn cores with two demarcations on the anterior surface. Horn core has one anterior keel at the base, before the first demarcation. Segment between the first and the second demarcations without keels, with a rounded anterior edge. Section above the second demarcation horn core anterior surface is wide and flat. One weak demarcation on the anterior edge in females; basal segment with a sharp anterior keel, apical one as in males. In each case apex of the horn core is rounded and possesses a very weak anterior keel, which disappears towards the base. Incisors as asymmetrical shovels, first incisor being the largest. Accessory pillars are more frequently absent on upper molars while on lower ones they are always present.

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Genus Phronetragus Gabunia, 1955

Type of the genus: Phronetragus arknethensis Gabunia, 1955

Diagnosis (Meladze, 1967, p. 103): Tragocerini of a small and medium size. Skull relatively narrow and long. Postcornual region well developed. Parietal crests present. Orbits not projecting, praeorbital fossa absent. Anterior edge of choana is behind the distal edge of M3/. Frontal bone is not in contact with temporal in the region of crista pretigoidea, alisphenoid and parietal bones are interposed between them (frontal and temporal bones, MB).Upper teeth large, placed almost along one straight line - width of the palate between M3/s equals width between P2/s. P2/, P3/ at the initial stage of molarisation, inner (lingual, MB) crescents are starting to be formed. Distal ribs on the buccal surface (metacone) on upper molars are much less developed than the mesial ones (paracone), it’s completely absent on M3/. Accessory pillars on upper and lower molars are always present; On the M/3 accessory pillar is in the mesial notch; besides, islet is present between inner crescents on M/1 and M/2. Buccal folds on the mesial surface of lower molars are weak. Both valleys are open on P/4. Horn cores attached directly behind orbits, well apart from each other at the base. Their length is always less than the half of the basal length of the skull. Horn cores have straight posterior and weakly convex anterior surface, without keels. Cross-section at the base is a stretched ellipse, its dorso-ventral diameter gradually decreases towards the apex. Horn cores are placed so that wide flat surface is directed mesially and somewhat laterally – angle between the largest diameters of the horn core cross-section is right or

open angle. Horn cores diverge with 35-40and are strongly inclined backwards, don’t rise above the frontal surface. In some cases they show weak tendency towards homonymous torsion.

Species Phronetragus arknethensis Gabunia, 1955

Pl. XXXVII figs. 1, 2.

Holotype: 41-2013/401 (5/4), skull fragment with a right horn core.

First publication: Gabunia, 1955; pp. 459-460.


Derivatio nominis: “arknethensis” – from the vil. Arkneti, where the site is located.

Diagnosis (Meladze, 1967, p. 103-104): small phronetragus. Distal ribs (metacone) on upper molars are weak, and absent on M3/; yet, buccal surface of the distal lobe is convex. On all upper molars the most feebly developed style among all the styles is the distal one (metastyle). On the lingual surface of M/1 and M/2 mesial and distal stylids are well

developed. Horn cores diverge at an angle 35.

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Species Phronetragus secundus Meladze, 1967

Pl. XXXVIII figs. 1-3.

Holotype: 21-2013/700 (B-777), fragmentary skull.

First publication: Meladze, 1967; pp. 105-107, pl. XXX figs. 1, 2.


Diagnosis (Meladze, 1967, p. 105): medium size phronetragus. Distal ribs (metacone, MB) on upper molars are very weakly developed and completely absent of the M3/, distal lobe of M3/ is flat buccally. Mesial and middle styles on upper molars strongly developed, distal style on the M1/ and M2/ are the most feeble, while on M3/ the distal style is even more developed then the other two ones (mesostyle and parastyle MB). On the lingual surface of the lower M/1 and M/2 mesial and distal stylids are weak. Horn cores diverge at

an angle not exceeding 20.

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Genus Udabnocerus Burchak-Abramovich & Gabashvili, 1969

Type species of the genus Udabnocerus georgicus Burchak-Abramovich & Gabashvili, 1969

Derivatio nominis: “Udabnocerus” – from the name of the Site Udabno.

Diagnosis (Burchak-Abramovich & Gabashvili, 1969, p. 77): skull of medium size with moderately elongated facial part; orbits weakly expressed; horn cores are with heteronymous torsion, directed backwards, up and slightly outwards. Horn cores are straight along the longitudinal plane, while in a transverse plane they are slightly convex arch-like in a lateral direction. Horn core cross-section is ellipse with larger ML diameter. horn core tissue compact (not pneumatised) in the middle of the horn core length. Horn core bases short. Postcornual part of the skull relatively shortened. Coronal suture well visible in adult state.Upper molars moderately brachyodont, without accessory elements.Milk teeth replacement of bovini type (P4/ erupts late). Premolars weakly reduced (index to Length P2/-M3/ is 45).

Horn core bases are located somewhat behind the orbits. They form 20 mm long postorbital-praecornual region. Praeorbital fossa small, with rather gentle slopes. Cornice in the postorbital and subcornual regions of frontals, above the well demarcated temporal fossa, is well visible. Anterior edge of the orbit is located at the level of distal lobe of M3/. Choana are not divided by septa. Anterior edge of choana ends much behind the distal edge of M3/. Palatine foramen is at the level of M2/-M3/ limit, distal edge of glenoid fossa for articulation of mandible is located at 2-3 mm from the anterior edge of horn core base. Layout of bones in the temporal fossa is of a bovini type. Parietal crests not sharp.

Species Udabnocerus georgicus Burchak-Abramovich & Gabashvili, 1969

Pl. XXXiX figs. 1-3.

Holotype: 32-2013/1133 (277-139), skull.

First publication: Burchak-Abramovich & Gabashvili, 1969; pp. 76-92, figs. 1-4 pl. I figs. 1, 2, pl. II, figs. 1-3.

Type locality and age: locality Ajia of the Udabno site, Georgia; Late Miocene, Upper Sarmatian-Meotian.

Derivatio nominis: “georgicus” – from the name of the country – Georgia.

Diagnosis (Burchak-Abramovich & Gabashvili, 1969, p. 77): the same as for the genus.

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Genus Urmiabos Burchak-Abramovich, 1950

Type species of the genus Urmiabos azerbaidzanicus Burchak-Abramovich, 1950

Derivatio nominis: “Urmiabos” – from the Site Urmia.

Diagnosis (Burchak-Abramovich, 1950, p. 876): Skull of a medium size. Well demarcated line between horn cores is projecting caudally from the level of horn core bases. This line divides frontal surface and occipital torus. Two concavities are expressed (The lateral one on the horn core base and medial one on the frontal surface itself) within the outline of the intercornual line on each halves of the skull. These concavities are divided by a protuberance. Caudal part of the frontals is weakly convex. Angle between frontal surface

and occipital torus is about 104. Occipital torus, which is delimited by intercornual ridge and linea nuchalis superior, is relatively narrow. Middle part of the occipital torus is concave in the region of parietal bones. Along the lateral edge of this concavity a well visible torus is extended obliquely from the linea nuchalis superior to the intercornual ridge delimiting parietal bone from frontal. Parietal region occupies more than half of the entire surface occupied by occipital torus. A flattened weakly concave area is located on the caudal surface of horn core base within the area between intercornual ridge, linea nuchalis superior, above mentioned lateral torus in the parietal region, and rough lateral surface of the horn core base. Horn cores are rounded-elliptical at the base, hollow. General direction of horn cores at the base is lateral, somewhat backwards and more distally – forwards and upwards.

Species Urmiabos azerbaidzanicus Burchak-Abramovich, 1950

Pl. XL figs. 1-3.

Holotype: 148-250, skull fragment with a right horn core.

First publication: Burchak-Abramovich, 1950; pp. 875-878, figs. 1a, 1b.

Type locality and age: Urmia, surroundings of the village Kurtevul 50 Km east of the Lake Urmia and city Meragheh, Iran; Early Pliocene.

Derivatio nominis: “azerbaidzanicus” – from the name of the region in northwestern Iran - Azerbaidjan.

Diagnosis (Burchak-Abramovich, 1950, p. 876): the same as for the genus.

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Genus Ioribos Vekua, 1972

Type species of the genus Ioribos aceros Vekua, 1972

Derivatio nominis:“Ioribos” – from the Riv. Iori (the site Kvabebi is located in the gorge of Iori).

Diagnosis (Vekua, 1972, p. 255): skull large, relatively elongated and narrow, with well developed postcornual region; frontals wide, flat, not rising above the level of orbits; parietal crests strongly developed, yet not bifurcated; orbits moderately projecting laterally; horn cores absent. Cheek teeth hypsodont, with well developed ribs, styles and accessory pillars, cement is absent.

Species Ioribos aceros Vekua, 1972

Holotype: K-1, complete skull.

First publication: Vekua, 1972; pp. 255-277, figs. 65-67, pl. XXXIV figs. 1-3.



* specimen not found.

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Genus Eosyncerus Vekua, 1972

Type species of the genus Eosyncerus ivericus Vekua, 1972

Diagnosis (Vekua, 1972, p. 277): Buffalo of a large size; horn cores of the medium length, very massive, somewhat bent upwards, outwards and backwards, weakly torsioned heteronymously; horn core bases close to each other, insignificantly constricted dorso-ventrally.

Species Eosyncerus ivericus Vekua, 1972

Pl. XLI fig. 1.

Holotype: 29-2013/534 (K-105), frontlet with horn cores.

First publication: Vekua, 1972; pp. 277-289, fig. 71, pl. XXXVI figs. 1, 2.


Derivatio nominis: “ivericus” – Iveria is one of the ways to spell the name of ancient Georgian kingdom Iberia.


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Genus Protobison Burchak-Abramovich, Gajiev & Vekua, 1980

Type species of the genus Protobison kuschkunensis Burchak-Abramovich, Gajiev & Vekua, 1980

Diagnosis (Burchak-abramovich et al., 1980, p. 486): Relatively small bison; skull of medium size, horn cores weak, with rounded-elliptical cross-section at the base (ratio - 87.9). Horn core base is weakly demarcated, burr not developed; horn cores at the base are

directed somewhat backwards and upwards, frontal surface in a transverse section more or less flat. Basioccipital narrow, elongated across the sagittal line.

Species Protobison kuschkunensis Burchak-Abramovich, Gajiev & Vekua, 1980

Pl. XLII figs. 1, 2.

Holotype: K-10, skull fragments.

First publication: Burchak-Abramovich, et al., 1980; pp. 485-487, figs. 1, 2.

Type locality and age: Kushkuna, Azerbaidjan; Early Pleistocene Middle Akchagylian.

Derivatio nominis: “kuschkunensis” – from the Mountain Kushkuna in Azerbaidjan.

Diagnosis (Burchak-abramovich et al., 1980, p. 486): the same as for the genus.

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Genus Dmanisibos Burchak-Abramovich & Vekua, 1994

Type species of the genus Dmanisibos georgicus Burchak-Abramovich & Vekua, 1994

Derivatio nominis: “Dmanisibos” – from the site Dmanisi.

Diagnosis (Burchak-Abramovich & Vekua 1994, pp. 253-254): Bos of a medium size. Frontal surface is strongly concave transversally in comparison with the praecornual part and the horn core bases. Postcornual part is inclined backwards. Cross-section of the postcornual region is undulated, with two lateral concavities and a medial convexity. Temporal fossa caudally wide and rounded. Distance between caudal edges of temporal fossae is less than the distance between lateral edges of occipital condyles. Occipital surface is perpendicular to the frontal section of the postcornual region. Length of frontal section of the postcornual region is 77.5% of the frontals. Foramen magnum is relatively narrow. Horn cores of a medium length and moderately massive (ratio between circumference at the base to the horn core to the horn core length across the curve is 82.5). Horn cores are weakly constricted (ratio – 82.2), directed in its basal part laterally, somewhat backwards and downwards, while keeping the backward direction abruptly turns upwards and in the distal half turns somewhat forwards.

Species Dmanisibos georgicus Burchak-Abramovich & Vekua, 1994

Pl. XLIII figs. 1, 2.

Holotype: D354, skull fragment.

First publication: Burchak-Abramovich & Vekua, 1994; pp. 253-261, figs. 1a, 1b.

Type locality and age: Dmanisi,Georgia; Early Pleistocene, 1.76 Ma.

Derivatio nominis: “georgicus” – from the country name Georgia.

Diagnosis (Burchak-Abramovich & Vekua 1994, pp. 253-254): the same as for the genus.

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Genus Parastrepsiceros Vekua, 1968

Type species of the genus Parastrepsiceros sokolovi Vekua, 1968

Diagnosis (Vekua, 1972, p. 238): Large antelope (orbital width is 132 mm; occipital height 70 mm). Braincase relatively long and wide; Parietal part of the supraoccipital occupies almost all the dorsal surface of the braincase; skull bending angle moderate (ca.

65). Horn cores long, massive, steeply set on the skull, coiled into open spiral (1 ½ turns), moderately torsioned. Among the existing two keels postero-lateral is more developed than the other; the antero-medial keel is well expressed only in the apical half. Teeth moderately hypsodont, upper cheek teeth without accessory tubercles, on the lower cheek teeth only anterior folds are present; P/4 with closed mesial valley.

Species Parastrepsiceros sokolovi Vekua, 1968

Pl. XLIV figs. 1-3.

Holotype: 29-2013/533 (K-601), braincase with horn cores.

First publication: Vekua, 1968; pp. 707-710, fig. 1.

Type locality and age:Kvabebi,Georgia; Late Pliocene, ca. 3 Ma.

Derivatio nominis: “sokolovi” – in honor of the palaeontologist Prof. Dr. Sokolov.


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Genus Capra Linnaeus, 1758

Species Capra dalii Bukhsianidze & Vekua, 2006

Pl. XLV figs. 1.

Holotype: D75, right horn core.

First publication: Bukhsianidze & Vekua, 2006; pp. 159-171, fig. 1.


Derivatio nominis: “dalii” – Dali is the name of the Georgian Goddess of wilderness and of hunters.

Diagnosis (Bukhsianidze & Vekua, 2006): in English, see the publication.

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Genus Gazella Blainville, 1816

Species Gazella propria Meladze, 1967

Pl. XLVI figs. 1-3.

Holotype: 41-2013/108 (5/33), skull.

First publication: Meladze, 1967; pp. 81-87, pl. XX figs. 1-3.


Diagnosis (Meladze, 1967, pp. 81-82): Small gazelle. Parietals strongly convex, Parietal crests well developed. Ethmoidal fissure is present. Females without horn cores. Length of horn cores exceeds half of the basal length of the skull; horn cores parallel to each other, somewhat bent towards the apex: apical profile almost straight, basal profile convex (in the basal part). Cross-section at the base is elliptical, largest diameters of the ellipse make less than right angle to each other. Surface of horn cores from even to strongly grooved with one sharp distal keel delimited by grooves. Teeth rather low crowned, small. Length of upper cheek teeth is ca. 1/3 of the basal length of the skull. Accessory elements of upper molars absent; on M/1 they are always present, while on M/2 and M/3 sometimes. P/4 with open anterior valley without a pillar, posterior valley closes at the medium or heavy ware stage.

__________

Species Gazella postmitilinii Vekua, 1972

Pl. XLVII figs. 1, 2.

Holotype: 29-2013/631 (K-17), skull.

First publication: Vekua, 1972; pp. 231-237, figs. 60, 61, pl. tab XXXII figs. 1, 2.


Derivatio nominis: “postmitilinii” – from the Gazella mitilinii – species of gazelle from Samos.

Diagnosis (Vekua, 1972, p. 231): relatively large gazelle. Skull wide (orbital width of the skull 103 mm), relatively low with convex frontal surface between orbits. Skull bending angle

is 70. Praeorbital fossae wide and deep; supraorbital foramina small and placed at the base of horn cores in narrow and deep pits. Orbits circular, noticeably projecting laterally. Horn cores long, bent backwards, with bases close to each other, diverging moderately from each other, cross-section almost circular. Teeth relatively hypsodont, without accessory pillars. P/4 with closed distal valley.

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Genus Paraoioceros Meladze, 1985

Type species of the genus Paraoioceros improvisus Meladze, 1985

Diagnosis (Meladze, 1985, p. 28): The largest representative of Oiocerini. Skull of medium dimensions. Postcornual region is well developed, it is straight. Frontals pneumatised, cavities enter the horn core bases as well. Horn cores diverge; they form one

complete homonymous spiral and possess several keels. Basioccipital is triangular, not flattened.

Species Paraoioceros improvisus Meladze, 1985

Pl. XLVIII fig. 1.

Holotype: R-555, horn cores.

First publication: Meladze, 1985. p. 28, pl. II fig. 1.

Type locality and age: Rustavi (Iagludja idem.), Georgia; Late Miocene, Upper Sarmatian.

Diagnosis (Meladze, 1985, p. 28): the same as for the genus.

__________

Genus Pontoceros Vereshchagin, Alexeeva, David & Baigusheva, 1971

Species Pontoceros surprine Vekua, 2012

Pl. XLIX fig. 1-4.

Holotype: D5552, skull fragment.

First publication: Vekua, 2012; pp. 139-145, figs. 1-4.


Diagnosis (Vekua, 2012, p. 140): in English, see the publication.

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Genus Protoryx Major, 1891

Species Protoryx heinrichi Vekua, 1972

Pl. L fig. 1-4.

Holotype: 29-2013/632 (K-16), skull.

First publication: Vekua, 1972. pp. 212-226, figs. 57, 58, pl. XXX fig. 1.


Derivatio nominis: “heinrichi” – in honor of the palaeobotanist, Dr. Heinrich Avakov, discoverer of the Kvabebi site.

Diagnosis (Vekua, 1972, p. 212): Antelope relatively not large (distance basion - praemolare is 150 mm; width of the skull at the orbits 75-84 mm) Skull is relatively high, with well developed postcornual region. Facial part of the skull is high, constricted from the sides.

Skull bending angle is about 50. Orbits large, circular, moderately projecting. Horn core bases close to each other, situated above orbits, long, set steeply on frontals, almost

parallel, bent arch-like backwards, cross-section elongated ellipse. Teeth mesohypsodont, without accessory elements. P/4 with open anterior and posterior valleys.

__________

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Meladze, G.K., 1967. Гиппарионовая фауна Аркнеи и Базалети [Hipparion fauna of Arkneti and Bazaleti]. "Metsniereba", Tbilisi,168 p., 28 figs., 32 pls. [in Russian, brief summary in English].

Meladze, G.K., 1985. Обзор гиппарионовых фаун Кавказа *Review of hipparion faunas of Caucasus]. "Mecniereba", Tbilisi, 93 p. 5 figs., 1 tab., 16 pls. [in Russian].

Tsiskarishvili, G.V., 1987. Позднетретичные носороги (Rhinocerotidae) Кавказа , *Late Tertiary rhinos (Rhinocerotidae) from the Caucasus]. "Mecniereba", Tbilisi 142 p. 13 tabs., 14 figs., 27 pls., [in Russian].

Vekua, A.K., 1959. hipopotamis naSTebi saqarTvelos qvedapleistocenuri naleqebidan [Remains of hippopotamus from the Early Pleistocene of Georgia]. saq. ssr mecnierebaTa akademiis moambe [Bulletin of the Academy of Sciences of the

Georgian SSR], 23(5): 561-656, 2 figs., 3 tabs. [in Georgian].

Vekua, A.K., 1960. Своеобразная ископаемая лошадь из плейстоцена Ахалкалаки (юг Грузии) *Peculiar fossil horse from the Pleistocene of Akhalkalaki+. Доклады Академии наук СССР *Proceedings of the USSR Academy of Sciences+,132(6):1417-1420, 3 figs., 2 tabs. [in Russian].

Vekua, A.K., 1962. axalqalaqis qvedapleistocenuri ZuZumwovrebis fauna [The Lower Pleistocene fauna of mammals of Akhalkalaki]. Publishing house of the GSSR Academy of Sciences, 207 p. 20 figs., 25 tabs., 19 pls. [in Georgian, with extended summary in Russian, brief summary in English].

Vekua, A.K., 1968. grexilrqiani antilopa saqarTvelos pliocenSi [Spiral

horned antelope form the Pliocene of Georgia]. saq. ssr mencierebaTa akademiis moambe [Bulletin of the Academy of Sciences of the Georgian SSR],

51(3):707-710, 2 figs. [in Georgian, summary in Russian].

Vekua, A.K., 1972. Квабебская фауна акчагыльских позвоночных *Kvabebi fauna of Akchagylian vertebrates]. 351 p. 79 figs., 72 tabs., 36 pls. "Nauka", Moskva [in Russian].

Vekua, A., Bendukidze, O., Kiladze, S., 2010. giganturi irmebi saqarTvelos plio-

pleistocenSi [Giant deers from the Plio-Pleistocene of Georgia]. saqarTvelos erovnuli muzeumis sabunebismetyvelo da preistoriuli seqciis macne [Proceedings of the Natural and Prehistoric

Section of the National Museum of Georgia], 2:38-53, 10 figs. [in Georgian, abstract in English].

Vekua, A.K., 2012. New Spiral-Horned Antelope in Dmanisi Fauna. Bulletin of the Georgian National Academy of Sciences, 6(3):139-145, 4 figs., 1 tab.

Plates

Plate I

Kvabebihyrax kachethicus Gabunia & Vekua, 1966

Holotype - 29-2013/299 (K-10), lower jaw with complete tooth row.

1. occlusal view

2. buccal veiw

Scale bar 1 cm.

Plate II

Stenofiber caucasicus Burchak-Abramovich & Gabashvili, 1980

Holotype - 32-2013/1075 (269-53), fragment of a hemi-mandible dex. with canine (basal part), P/4-M/2.

1. occlusal view

2. lingual view

3. buccal veiw

Scale bar 1 cm.

Plate III

Platybelodon jamandzhalgensis Beljaeva & Gabunia, 1960

Holotype - 20-2013/221 (5/23-5/24) incomplete hemi-mandible dex. with DI/2, DP/2-DP/4.

1. occlusal view

Scale bar 1 cm.

Plate IV

Mammuthus meridionalis taribanensis Gabunia & Vekua, 1963.

Skull of the holotype, K/13. Skull is mounted on an iron construction.

1. frontal view

2. lateral view

3. occipital view

Scale bar 20 cm.

Palte V

Udabnopithecus garedziensis Burchak-Abramovich & Gabashvili, 1945

Holotype - 156-192, fragment of a maxilla dex. with P4/ and M1/

1.occlusal P4/ dex.

2.mesial view P4/ dex.

3.lingual view P4/ dex.

4.distal view P4/ dex.

5.buccal view P4/ dex.

6. occlusal view M1/dex.

7.mesial view M1/dex.

8.lingual view M1/dex.

9.distal view M1/dex.

10. buccal view M1/dex.

Scale bar 1 cm.

Palte VI

Homo georgicus Gabunia, Lumley, Vekua, Lordkipanidze & de Lumley, 2002

Holotype D2600, almost complete mandible.

1. lateral veiw (right side)

2 frontal view

3 lateral veiw (left side)

4. superior view

Scale bar 1 cm.

Plate VII

Machairodus davitashvili Vekua, 1972

Holotype - 29-2013/453 (K-14), skull.

1. lateral view

Scale bar 5 cm.

Plate VIII

Crocuta abessalomi Gabunia, 1958

Holotype: 20-2013/128 (# 35), incomplete hemi-mandible sin. with P/2, P/3 and M/1.

1. superior view

2. lateral view

3. medial view

Scale bar 1 cm.

Plate IX

Crocuta miriani Meladze, 1967

Holotype - 21-2013/1201 (B-40), skull.

1. superior view

2. lateral view

3. basal view

Scale bar 5 cm.

Plate X

Cephalogale meschethense Gabunia, 1964

Holotype: 3-2013/-65 (7/411), M/1 sin.

1. occlusal view

2. buccal view

3 lingual view

Scale bar 5 mm.

Plate XI

Canis tengisii Vekua, 1962

Holotype- 18-2012/1215 & 18-2012/1216 (Akh-214, 664, 681, 684, 883, 907), skull fragments and a hemi-mandible dex.

1 praemaxillae & maxillae (18-2012/1215) occlusal view

2. praemaxillae & maxillae (18-2012/1215) buccal view

3. praemaxillae & maxillae (18-2012/1215) lingual view

4. mandible (18-2012/1216) occlusal view

5. mandible (18-2012/1216) mandible buccal view

6. mandible (18-2012/1216) lingual view

Scale bar 1 cm.

Plate XII

Amphicyon caucasicus Gabunia, 1973

Holotype- 20-2013/110 (#77), hemi-mandible dex. with I/2, I/3, /C, P/3-M/2.

1. occlusal view

2. buccal view

3. lingual view

Scale bar 1 cm.

Plate XIII

Ardynia plicidentata Gabunia, 1964

Holotype - 3-2013/45 (7/78), M2/ sin.

1. occlusal view

2. lingual view

3. buccal view

4. mesial view

5. distal view

Scale bar 1 cm.

Plate XIV

Benaratherium callistrati Gabunia, 1955

Holotype - 3-2013/1 (7/15), hemi-mandible dex. with P/3-M/3.

1. occlusal view

2. buccal view

3. linual view

Scale bar 5 cm.

Plate XV

Diceros gabuniai Tsiskarishvili, 1987

Holotype - 32-2013/1115 (352-1), almost complete skull.

1. lateral view

2. occipital view

3. superior view

4. basal view

Scale bar 10 cm.

Plate XVI

Chilotherium eldaricum Tsiskarishvili, 1987

Holotype - 32-2013/1109 (352-15), mandible.

1. occlusal view

2. lateral view

Scale bar 10 cm.

Plate XVII

Dicerorhinus vekuai Tsiskarishvili, 1987

Holotype - 29-2013/298 & 29-2013/299, incomplete skull and incomplete hemi-mandible dex.

1. skull 29-2013/298 superior view

2. skull 29-2013/298 basal view

3. mandible 29-2013/299 occlusal view

4. mandible 29-2013/299 buccal view

5. mandible 29-2013/299 lingual view

Scale abr 10 cm.

Plate XVIII

Schizotherium chucuae Gabunia, 1951

Holotype - 3-2013/81 (7/7), incomplete hemi-mandible dex.

1. occlusal view

2. buccal view

3. lingual view

Scale bar 1 cm.

Plate XIX

Hipparion eldaricum Gabunia, 1959

Holotype - 32-2013/47 & 32-2013/6 (139-85), fragmentary skull.

1. lateral view (tooth row - lingual side, brain case - lateral left side)

2. occlusal view of the tooth row dex.

Scale bar 2 cm.

Plate XX

Hipparion garedzicum Gabunia, 1959

Holotype - 32-2013/1003 (156/13), fragmentary skull.

1. superior view

2. lateral view

3. basal view

Scale bar 5 cm.

Plate XXI

Hipparion urmiense Gabunia, 1959

Holotype - 148/191, maxilla with P2/-M3/ dex.

1. lateral view

2. occlusal view

Scale bar2 cm.

Plate XXII

Equus hipparionoides Vekua, 1960

Holotype - 18-2012/1733 (Akh-99), lower tooth row P/2-M/2 sin.

1. occlusal view

2. buccal view

3. lingual view

Scale bar 2 cm.

Palte XXIII

Anthracotherium kwablianicum Gabunia, 1964

Holotype - 3-2013/43 (7/310) – M2/ dex. and 3-2013/63 (7/317) – M/3 dex.

1. M2/ occlusal view

2. M2/ lingual view

3. M2/ buccal view

4. M/3 occlusal view

5. M/3 buccal view

6. M/3 lingual view

Scale bar 1 cm.

Palte XXIV

Paraentelodon intermedium Gabunia, 1964

Holotype: incomplete upper tooth row: 3-2013/22 (7/181) - P1/ dex., 3-2013/19 (7/188) – M2/ dex., 3-2013/27 (7/189) – M3/ dex., 3-2013/23 (7/184), P1/ sin., 3-2013/24 (7/185) – P3/ sin., 3-2013/25 (7/186) – P4/ sin., incomplete lower tooth row: 3-2013/28 (7/194) – P/3 sin., 3-2013/29 (7/192) – M/2 sin., 3-2013/30 (7/195) – M/3 sin., 3-2013/31 (7/193) – M/3 dex. and of the same individual.

1. 3-2013/22 (7/181) - P1/ dex. occlusal view

2. 3-2013/22 (7/181) - P1/ dex. buccal view

3. 3-2013/22 (7/181) - P1/ dex. lingual view

4. 3-2013/23 (7/184) - P1/ sin. occlusal view

5. 3-2013/23 (7/184) - P1/ sin. lingual view

6. 3-2013/23 (7/184) - P1/ sin. buccal view

7. 3-2013/24 (7/185) –P3/ sin. occlusal view

8. 3-2013/24 (7/185) – P3/ sin. buccal view

9. 3-2013/24 (7/185) – P3/ sin. lingual view

10. 3-2013/25 (7/186) – P4/ sin. occlusal view

11. 3-2013/25 (7/186) – P4/ sin. distal view

12. 3-2013/25 (7/186) – P4/ sin. mesial view

13. 3-2013/19 (7/188) – M2/ dex. occlusal view

14. 3-2013/19 (7/188) – M2/ dex. mesial view

15. 3-2013/19 (7/188) – M2/ dex. distal view

16. 3-2013/27 (7/189) – M3/ dex. occlusal view

17. 3-2013/27 (7/189) – M3/ dex. buccal view

18. 3-2013/31 (7/193) – M/3 dex. occlusal view

19. 3-2013/31 (7/193) – M/3 dex. distal view

20. 3-2013/31 (7/193) – M/3 dex. mesial view

21. 3-2013/29 (7/192) – M/2 sin. occlusal view

22. 3-2013/29 (7/192) – M/2 sin. lingual view

23. 3-2013/29 (7/192) – M/2 sin. buccal view

24. 3-2013/28 (7/194) – P/3 sin occlusal view

25. 3-2013/28 (7/194) – P/3 sin buccal view

26. 3-2013/28 (7/194) – P/3 sin lingual view

27. 3-2013/30 (7/195) – M/3 sin. occlusal view

28. 3-2013/30 (7/195) – M/3 sin. lingual view

29. 3-2013/30 (7/195) – M/3 sin. buccal view

Scale 2 cm

Palte XXV

Paraentelodon intermedium Gabunia, 1964

holotype - manus dex.: 3-2013/32 (7/184) – mc IV, 3-2013/33 (7/186) – mc III, 3-2013/34 (7/181) - phalanx prox., 3-2013/35 (7/185) – phalanx prox. of the same individual.

1. 3-2013/32 (7/184) – mc IV dex. proximal view

2. 3-2013/32 (7/184) – mc IV dex. dorsal view

3. 3-2013/32 (7/184) – mc IV dex. palmar view

4. 3-2013/32 (7/184) – mc IV dex. lateral view

5. 3-2013/32 (7/184) – mc IV dex. medial view

6. 3-2013/33 (7/186) – mc III dex.dorsal view

7. 3-2013/33 (7/186) – mc III dex.palmar view

8. 3-2013/34 (7/181) - phalanx prox. (iv digit) proximal view

9. 3-2013/34 (7/181) - phalanx prox. (iv digit) dorsal view

10. 3-2013/34 (7/181) - phalanx prox. (iv digit) palmar view

11. 3-2013/35 (7/185) – phalanx prox. (iii digit) proximal view

12. 3-2013/35 (7/185) – phalanx prox. (iii digit) dorsal veiw

13. 3-2013/35 (7/185) – phalanx prox. (iii digit) palmarview

14. 3-2013/35 (7/185) – phalanx prox. (iii digit) lateral veiw

Scale 2 cm

Plate XXVI

Kubanochoerus robustus Gabunia, 1955

Holotype - 20-2013/213 (# 33), mandible.

1. occlusal view

Scale bar 5 cm.

Plate XXVII

Hippopotamus georgicus Vekua, 1959

Holotype - articulated manus dex. - 18-2012/585 (Akh. 1), radius dex.; 18-2012/586 (Akh. 2), piramidale dex.; 18-2012/587 (Akh. 3), lunatum dex.; 18-2012/588 (Akh. 4), scaphoideum dex.; 18-2012/589 (Akh. 5), trapezoideum dex.; 18-2012/590 (Akh. 14) mc V dex.; 18-2012/591 (Akh. 12) mc IV dex.; 18-2012/592 (Akh. 9), mc IIIdex.; 18-2012/593 (Akh. 6), mc II dex.; 18-2012/594 (Akh. 15), I phalanx (v digit) dex.; 18-2012/595 (Akh. 13), I phalanx (iv digit) dex.; 18-2012/596 (Akh. 10), I phalanx (iii digit) dex.; 18-2012/597 (Akh. 7), I phalanx (ii digit) dex.; 18-2012/598 (Akh. 16), II phalanx (v digit) dex.; 18-2012/599 (Akh. 11), II phalanx (iii digit) dex.; 18-2012/600 (Akh. 8), II phalanx (ii digit) dex.

1. 18-2012/585 (Akh. 1), radius dex. dorsal view

2. 18-2012/585 (Akh. 1), radius dex. palmar view

3. 18-2012/585 (Akh. 1), radius dex. distal view

4. 18-2012/586 (Akh. 2), piramidale dex. dorsal view

5. 18-2012/586 (Akh. 2), piramidale dex. lateral view

6. 18-2012/586 (Akh. 2), piramidale dex. palmar view

7. 18-2012/586 (Akh. 2), piramidale dex. medial view

8. 18-2012/586 (Akh. 2), piramidale dex. proximal view

9. 18-2012/586 (Akh. 2), piramidale dex. distal view

10. 18-2012/587 (Akh. 3), lunatum dex. proximal view

11. 18-2012/587 (Akh. 3), lunatum dex. distal view

12. 18-2012/587 (Akh. 3), lunatum dex. palmar view

13. 18-2012/587 (Akh. 3), lunatum dex. dorsal view

14. 18-2012/587 (Akh. 3), lunatum dex. medial view

15. 18-2012/587 (Akh. 3), lunatum dex. lateral view

16. 18-2012/588 (Akh. 4), scaphoideum dex. medial view

17. 18-2012/588 (Akh. 4), scaphoideum dex lateral view

18. 18-2012/588 (Akh. 4), scaphoideum dex proximal view

19. 18-2012/588 (Akh. 4), scaphoideum dex distal view

20. 18-2012/593 (Akh. 6), mc II dex. dorsal view

21 18-2012/593 (Akh. 6), mc II dex. palmar view

22. 18-2012/593 (Akh. 6), mc II dex. medial view

23. 18-2012/593 (Akh. 6), mc II dex. lateral view

24. 18-2012/593 (Akh. 6), mc II dex. proximal view

25. 18-2012/592 (Akh. 9), mc IIIdex. dorsal view

26. 18-2012/592 (Akh. 9), mc IIIdex. palmar view

27. 18-2012/592 (Akh. 9), mc IIIdex. lateral view

28. 18-2012/592 (Akh. 9), mc IIIdex. medial view

29 18-2012/592 (Akh. 9), mc IIIdex. proximal view

Scale bar 5 cm

Plate XXVIII

Hippopotamus georgicus Vekua, 1959

continues.

1. 18-2012/591 (Akh. 12) mc IV dex. dorsal view

2. 18-2012/591 (Akh. 12) mc IV dex. palmar view

3. 18-2012/591 (Akh. 12) mc IV dex. medial view

4. 18-2012/591 (Akh. 12) mc IV dex. lateral view

5. 18-2012/591 (Akh. 12) mc IV dex. proximal view

6. 18-2012/590 (Akh. 14) mc V dex. dorsal view

7. 18-2012/590 (Akh. 14) mc V dex. palmar view

8. 18-2012/590 (Akh. 14) mc V dex. medial view

9. 18-2012/590 (Akh. 14) mc V dex. lateral view

10. 18-2012/590 (Akh. 14) mc V dex. proximal view

11. 18-2012/597 (Akh. 7), I phalanx (ii digit) dex.dorsal view

12. 18-2012/597 (Akh. 7), I phalanx (ii digit) dex. palmar view

13. 18-2012/597 (Akh. 7), I phalanx (ii digit) dex. lateral view

14. 18-2012/597 (Akh. 7), I phalanx (ii digit) dex. proximal view

15. 18-2012/596 (Akh. 10), I phalanx (iii digit) dex. dorsal view

16. 18-2012/596 (Akh. 10), I phalanx (iii digit) dex.palmar view

17. 18-2012/596 (Akh. 10), I phalanx (iii digit) dex. medial view

18. 18-2012/596 (Akh. 10), I phalanx (iii digit) dex. proximnal view

19. 18-2012/595 (Akh. 13), I phalanx (iv digit) dex.dorsal view

20. 18-2012/595 (Akh. 13), I phalanx (iv digit) dex.palmar view

21. 18-2012/595 (Akh. 13), I phalanx (iv digit) dex. medial view

22. 18-2012/595 (Akh. 13), I phalanx (iv digit) dex. proximal view

23. 18-2012/594 (Akh. 15), I phalanx (v digit) dex. dorsal view

24. 18-2012/594 (Akh. 15), I phalanx (v digit) dex. palmar view

25. 18-2012/594 (Akh. 15), I phalanx (v digit) dex. medial view

26. 18-2012/594 (Akh. 15), I phalanx (v digit) dex. proximal view

27. 18-2012/600 (Akh. 8), II phalanx (ii digit) dex. dorsal view

28. 18-2012/600 (Akh. 8), II phalanx (ii digit) dex. palmar view

29. 18-2012/600 (Akh. 8), II phalanx (ii digit) dex. proximal view

30. 18-2012/599 (Akh. 11), II phalanx (iii digit) dex. dorsal view

31. 18-2012/599 (Akh. 11), II phalanx (iii digit) dex. palmar view

32. 18-2012/599 (Akh. 11), II phalanx (iii digit) dex. proximal view

33. 18-2012/598 (Akh. 16), II phalanx (v digit) dex. dorsal view

34. 18-2012/598 (Akh. 16), II phalanx (v digit) dex. aplamr view

35. 18-2012/598 (Akh. 16), II phalanx (v digit) dex. proximal view

Scale bar 5 cm

Plate XXIX

Lophiomeryx benarensis Gabunia, 1951

Holotype (3-2013/54 (7/10)) fragment - hemi-mandible dex. fragment with P/4

1. occlusal view

2. buccal view

3. lingual view

Scale bar 1 cm.

Plate XXX

Karsimatherium bazalethicum Meladze, 1961 skull.

1. superior view

2. lateral view

3. basal view

Scale bar 5 cm.

Plate XXXI

Paradicrocerus flerovi Gabunia, 1959

Holotype - 20-2013/50 (5/11), left antler.

1. frontal view

2. superior view

Scale bar 1 cm.

Plate XXXII

Dicrocerus salomeae Gabunia, 1955

Holotype - 41-2013/140 (5/13), left antler.

1. frontal view

Scale bar 1 cm.

Plate XXXIII

Arvernoceros insolitus Vekua, Bendukidze & Kiladze, 2010

Holotype- D2774, almost complete left antler.

1. frontal view

Scale bar 10 cm.

Plate XXXIV

Cervus abesalomi Kahlke, 2001

Holotype- D1495, complete antler.

1. frontal view

Scale bar 10 cm.

Plate XXXV

Mirabilocerus brevicornis Meladze, 1967

Holotype - 41-2013/67 (5/30), braincase with horn cores.

1. superior view

2. lateral view

3. occipital view

4. basal view

Scale bar 5 cm.

Plate XXXVI

Mirabilocerus maius Meladze, 1967

Holotype - 21-2013/710 (B-555), skull.

1. latral view

2. superior view

3. occipital view

4. basal view

Scale bar 5 cm.

Plate XXXVII

Phronetragus arknethensis Gabunia, 1955

Holotype - 41-2013/401 (5/4), skull fragment with a right horn core.

1. superior view

2. lateral view

Scale bar 2 cm.

Plate XXXVIII

Phronetragus secundus Meladze, 1967

Holotype - 21-2013/700 (B-777), skull.

1. superior view

2. lateral view

3. basal view

Scale bar 5 cm.

Plate XXXIX

Udabnocerus georgicus Burchak-Abramovich & Gabashvili, 1969

Holotype - 32-2013/1133 (277-139), skull.

1. superior view

2. lateral view.

3. basal view

Scale bar 5 cm.

Plate XL

Urmiabos azerbaidzanicus Burchak-Abramovich, 1950

Holotype - 148-250, skull fragment.

1. superior view

2. occipital view

3. frontal views

Scale bar 5 cm.

Plate XLI

Eosyncerus ivericus Vekua, 1972

Holotype - 29-2013/534 (K-105), frontlet with horn cores.

1. superior view

Scale bar 10 cm.

Plate XLII

Protobison kuschkunensis Burchak-Abramovich, Gajiev & Vekua, 1980

Holotype - K-10, skull fragments.

1. superior view

2. basal view

Scale barr 5 cm.

Plate XLIII

Dmanisibos georgicus Burchak-Abramovich & Vekua, 1994

Holotype - D354, skull fragment.

1. superior view

2. occipital view

Scale bar 5 cm.

Plate XLIV

Parastrepsiceros sokolovi Vekua, 1968

Holotype - 29-2013/533 (K-601), braincase with horn cores.

1. frontal view

2. basal veiw

3. occipital veiw

Scale bar 5 cm

Plate XLV

Capra dalii Bukhsianidze & Vekua, 2006

Holotype - D75, right horn core.

1. posterior view

Scale bar 5 cm.

Plate XLVI

Gazella propria Meladze, 1967

Holotype - 41-2013/108 (5/33), skull.

1. lateral view

2. superior view

3. basal view

Scale bar 2 cm.

Plate XLVII

Gazella postmitilinii Vekua, 1972

Holotype - 29-2013/631 (K-17), skull.

1. superior view

2. lateral view

Scale bar 2 cm.

Plate XLVIII

Paraoioceros improvisus Meladze, 1985

Holotype - R-555, horn cores.

1. frontal veiw

Scale bar 5 cm.

Plate XLIX

Pontoceros surprine Vekua, 2012

Holotype - D5552, skull fragment.

1. superior view

2. occipital view

3. basal view

4. lateral view

Scale bar 5 cm.

Plate L

Protoryx heinrichi Vekua, 1972

Holotype - 29-2013/632 (K-16), skull.

1. lateral view

2. uperior view

3. occipital view

4. basal view

Sale bar 2 cm.

catalogue of the holotypes of fossil land mammals in the...

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