LARGE MAMMALS FROM NEW LATE MIOCENE FOSSIL … medium sized hipparion species of Maragheh are...

LARGE MAMMALS FROM NEW LATE MIOCENE FOSSIL

LOCALITIES NEAR VARZEGHAN, NORTHWEST IRAN

Majid MIRZAIE ATAABADI1, Gholamreza ZAREE2, Zahra ORAK2

1 Department of Geosciences and Geography, University of Helsinki, P.O. Box 64,

FI-00014, Helsinki, Finland

2 Environment Protection Organization, MMTT, Tehran, Iran

Abstract

Large mammals from localities near Varzeghan in northwestern Iran are described here for the first

time. The fossil material consists of hipparionine horses and deinotheres. The hipparion species are

tentatively differentiated according to their size and dental morphology into a small to medium sized

species like Hipparion moldavicum and a medium to large sized species such as H. prostylum or H.

dietrichi. Deinotherium giganteum is also present in this area. A Middle Turolian age (MN 12

equivalent) is proposed for these new localities based on comparison of the fossil material to material

from other Late Miocene sites.

Key words Mammals, Miocene, Turolian, Varzeghan, Maragheh, Iran

1 Introduction Although extensive outcrops of Neogene

terrestrial deposits exist in northwest and

central Iran, knowledge of mammalian fossil

faunas in the country remains limited due to

lack of sufficient investigations. The only

exception is the richly fossiliferous beds of the

Maragheh Formation in northwest Iran. These

fossiliferous deposits are situated on the

southern and eastern slopes of the Sahand

volcano in the Maragheh district of eastern

Azarbaijan Province. For more than one and a

half centuries, this famous Late Miocene

vertebrate locality received much attention and

has been the exclusive source of information

on Neogene mammalian faunas of Iran. Based

on the latest studies the Maragheh Fm. has

been divided into three biostratigraphic

intervals (Lower, Middle and Upper

Maragheh), ranging from 9.5 to 7 million years

(Ma) in age (Bernor, 1986).

Nevertheless, recent discovery of new

localities in the northwestern parts of Iran has

demonstrated a wider distribution of mammal

bearing deposits in this area. These new

fossiliferous areas are situated north and north

east of Tabriz, the capital city of East

Azarbaijan province and are ca. 110-150 km

away from Maragheh (Fig. 1). The mammalian

fauna of Ivand, one of these new localities, has

been studied recently (Sen and Purabrishemi

2010, Mirzaie Ataabadi et al. ms). Here we

report on large mammal fossils from new

localities near Varzeghan.

MIRZAIE ATAABADI ET AL. / LATE MIOCENE LARGE MAMMALS FROM VARZEGHAN

2

Fig. 1 The geographic location of the new late Miocene mammal fossil localities (Varzeghan and

Ivand) in NW Iran (map modified after Bernor, 1986)

2 Geological setting

During the Paleogene, Azarbaijan province in

northwest Iran experienced a wide range of

volcanic activities. By the end of the early

Miocene the last Tethyan seaway disappeared

from the area, carbonate deposition terminated,

and a major landmass emerged (Aghanabati,

2004). Consequently, at the beginning of the

Neogene, this domain was mostly above sea

level and had incipient mountain ranges, basin

troughs, and a topography resembling present

day conditions (Davoudzadeh et al., 1997).

The areas north of the Tabriz fault (the

Anatolian transform of figure 1) in particular,

are segments of the Tabriz-Maku sub-zone in

the western Alborz-Azarbaijan structural zone.

During the Late Miocene and Pliocene this

zone, like other parts of Iran, underwent major

orogenic movements that established new

erosional cycles that filled the locally closed

basins with fluvial and lacustrine deposits

(Aghanabati, 2004). The new fossil localities

in the Varzeghan (and Ivand) districts and their

mammalian faunas, bounded by the Anatolian

transform to the south (Fig. 1), were formed in

such local basin troughs with different

environments compared to Maragheh

Formation.

3 Fossil localities and material

Varzeghan is situated (ca. 60 km) northeast of

Tabriz (Fig. 1). This locality is less well known

than the Ivand locality, mainly because of its

more recent discovery as a consequence of

construction activities in the area. Because of

the occurrence of fossils on the surface and

lack of sediment exposures, the nature of the

fossil bearing sequence and especially its

sedimentology and depositional environment is

not well understood. In Abkhare village (ca


3

38° 31' N, 46° 32' E), the fossil site of

deinothere material is a local concentration of

bones within a mudstone bed. The fossil

pocket is restricted to a single horizon.

Unfortunately, due to lack of outcrops

only a two-meter deep trench in Abkhare

village near the fossil site is the source of

sedimentology information. Here, fine grained

sediments are mainly present with a minor

lenticular pebbly sand layer (channel deposit)

which indicates a fluvial environment.

The deinothere material is partly

articulated. Fossils are very well preserved and

do not show any sign of weathering or

abrasion. The hipparion material is also very

well preserved. It seems that partly articulated

hipparions, as well as other fossil animals,

could be found at this locality in the future.

Therefore, the taphonomy of the fossil material

in this locality seems to be different from that

of Maragheh.

Fossil bones in Maragheh Fm. occur as

localized concentrations within the

unlaminated beds, floating in the sediments

rather than lying on bedding-planes. A single

complete articulated skeleton of the mustelid

Promeles palaeattica has been found from the

MMTT 13 quarry (Bernor et al., 1996).

Taphonomic studies of these fossil

accumulations indicate autochthonous bone

assemblages accumulated on overbank or

floodplain deposits of fluvial systems. A large

number of the bones are preserved with

articulation of distal limb elements and early

weathering stages. Pyroclastic events such as

mudflows or ash falls were not directly

responsible for mortality. On the other hand,

biologic agents were the probable cause of

death, as the bones were subaerially exposed

only long enough to allow removal of some

elements by scavengers (Morris, 1997).

The collected fossil material includes several

cranial and postcranial elements of a large

deinothere, which were unearthed while

renovating an old building in the outlying parts

of Abkhare village. These materials are stored

temporarily in the Department of Geology,

University of Tabriz, and the Department of

Environment local branch office in Maragheh.

In addition, about 100 isolated maxillary and

mandibular cheek teeth, 10 mandibular

fragments, part of a juvenile skull and a few

distal metapodials of hipparionine horses were

recovered from Varzeghan district. This

material is stored in the Department of

Environment local branch office in Maragheh.

The hipparion material has been

apparently collected from a construction site in

the city of Varzeghan (ca. 38o 30' N, 46

o 40'

E). However, we cannot exclude the possibility

of its mixture with fossils from other localities

in Varzeghan district.

The current study is based on the best

preserved mandibular fragments of hipparions

and the maxillary molar teeth of the

deinothere. Postcranial remains of the

deinothere were unavailable for this study. The

absence of other fossil groups such as bovids is

mostly due to collection bias and the limited

time available for collecting in the field.

4 Abbreviations

MMTT Muze Melli Tarikh Tabiei (National

Museum of Natural History), Tehran

MN European Mammal Neogene Faunal

Zone

M1-3 Upper Molars

p2-4 Lower premolars

m1-3 Lower molars

5 Systematic paleontology

Order Perissodactyla Owen, 1848

Family Equidae Gray, 1821

Genus Hipparion de Christol, 1832

Hipparion sp. 1 (small to medium sized)

Locality Varzeghan District

Age Middle Turolian, MN 12 equivalent

Material Right mandibular fragment with p2-

m3, MMTT-2-102-V318 (Fig 2-A, Tables 1-2)

Description The tooth row is relatively short

and the teeth are small. The premolars are wide

and robust. The length of p2-m3 is 130.8 mm

and the ratio (premolars length) ×100 / (molars

length) is 115, indicating a short molar row.

The parastylid is relatively well developed.

The metaconid and the metastylid are round to

subtriangular. The entoconid is round to

elliptical. The enamel is plicated at the flexid

borders, particularly in the premolars. The

ectoflexid is V-shaped and is moderately deep

and narrow on p4-m2, reaching the middle of

the tooth but not touching the linguaflexid. It is

deeper on m1-m2 but does not separate the

pre- and post-flexids. The linguaflexid is V-

shaped but shallower and more open on p3-p4.

The hypoconulid is well developed. A weak

plicabalinid is present on p2-p3.

Hipparion sp. 2 (medium to large sized)

Locality Varzeghan District


Material Left mandibular fragment with p3-

m3, MMTT-2-108-V324 (Fig. 2-F, Tables 1-

2); left mandibular fragment with p3-m3,

MMTT-2-105-V321 (Fig. 2-G, Tables 1-2);

right mandibular fragment with p3-m3 and the

symphysis, MMTT-2-101-V317 (Fig. 2C-D,

Tables 1-2); right mandibular fragment with

p2-m2, MMTT-2-106-V322 (Fig. 2-E, Tables

1-2); right mandibular fragment with p3-m1,

MMTT-2-103-V319 (Fig. 2-B, Tables 1-2);

right mandibular fragment with p3-m2,

MMTT-2-104-V320 (Fig. 2-H, Tables 1-2)

Description

The tooth row is relatively long and the teeth

are moderately large. The premolars are wide

and robust. The length of p2-m3 is estimated to

be 145-150 mm and the ratio (premolars

length) ×100 / (molars length) is estimated

114, indicating a short molar row. The

parastylid is well developed and closed. The

metaconid and the metastylid are round to

subtriangular. The entoconid is elliptical to

round. The enamel is plicated at the flexids and

their borders, particularly on the premolars.

The ectoflexid is U-shaped and moderately

deep and narrow on the premolars, reaching

the middle of the tooth but not touching the

linguaflexid and not separating pre- and post-

flexids. The ectoflexid is V-shaped and deep

on the molars, reaching the linguaflexid and

separating the pre- and post-flexids. The

linguaflexid is U-shaped, shallow and open. It

is deeper and V-shaped on the worn teeth. The

hypoconulid is well developed. A plicabalinid

is present especially on premolars.

TABLE 1 Measurements of Hipparion sp. 1 and sp. 2 mandibular fragments, Varzeghan District, NW

Iran

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Fig. 2 Hipparion sp. 1 and sp. 2, Varzeghan District, NW Iran. Mandibular fragment occlusal views:

A, V318: right p2-m3; B, V319: Right p3-m1; C-D, V317: right p3-m3; E, V322: right p2-m2; F, V324:

Left p3-m3; G, V321: left p3-m3; H, V320: Right p3-m2. Scale bars: 20 mm


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TABLE 2 Measurements of Hipparion sp. 1 and sp. 2 mandibular cheek teeth, Varzeghan District,

NW Iran

Comparisons

Two species of hipparionine horses from

Varzeghan are discriminated here principally

based on their size and dental morphology.

Hipparion sp. 1 is represented by a single

mandibular fragment and belongs to a small to

medium sized species while the rest of

mandibular fragments belong to a larger

species. The dental morphology of the

mandibular cheek teeth of these two species is

also somewhat different. The ectoflexid in the

larger species is deeper and the linguaflexid is

more open and shallow.

Due to the geographical proximity of the

Varzeghan and Maragheh localities,

comparison between hipparion materials of

these localities is of interest. The small and

medium sized hipparion species of Maragheh

are Hipparion matthewi and H. moldavicum.

The type material of the small hipparion H.

matthewi is from Samos Island in Greece. The

length of the lower tooth row of this species in

the type specimen is about 110mm. H.

moldavicum is the most abundant medium

sized hipparion species at Maragheh.


7

It is also very widespread in the northern

Black sea region where the type specimen was

found. The upper tooth row of this species

from Maragheh has a length of 120-140 mm

(Bernor 1985). Also, a recent comprehensive

study on the equid material from the

Akkaşdaği locality in Turkey has documented

a range of 128-142 mm for the length of the

lower tooth row (Koufos and Vlachou, 2005)

of H. moldavicum. Therefore, according to

these data Hipparion sp. 1 from Varzeghan is

in the size range of H. moldavicum and might

belong to it. Features in the mandibular dental

morphology such as deeper ectoflexids and V-

shaped linguaflexids in the molars of

Hipparion sp. 1 from Varzeghan support this

assignment.

The medium to large sized hipparionine

horses from Maragheh are H. prostylum, H.

campbelli, and H. gettyi. H. prostylum is more

abundant and smaller in size than the other

species. The upper tooth row of this species is

generally 140-150 mm long (Bernor, 1985).

Koufos and Vlachou (2005) have pointed to

the close similarity of H. dietrichi from

Akkaşdaği and some Greek localities to H.

prostylum from Maragheh. They recorded a

length of 138-145 mm for the lower cheek

tooth row of H. dietrichi from the Akkaşdaği

locality in Turkey. Thus, Hipparion sp. 2 from

Varzeghan is similar in size to H. prostylum

and H. dietrichi and might belong to either of

these species. Similarity in mandibular cheek

tooth morphology between these species, such

as deep ectoflexids and shallow and open

linguaflexids and a weak plicabalinid support

this idea. Unfortunately the muzzle is not

preserved in most of these fossils and hence

the main characteristic of H. dietrichi-H.

prostylum, which is the short and wide snout,

cannot be determined. Only one specimen

(MMTT-2-101-V317, Fig. 2-C) has the

symphysis partly preserved. In this individual

the short diastema is similar to that of H.

dietrichi, suggesting a short snout.

Order Proboscidea Illiger, 1811

Suborder Deinotherioidea Osborn, 1921

Family Deinotheriidae Bonaparte, 1845

Genus Deinotherium Kaup, 1829

Deinotherium giganteum Kaup, 1829

Locality Abkhare village, Varzeghan District


Material Upper molars including right M1,

MMTT-V16; M2, MMTT-V11 and M3,

MMTT-V14 and left M1, MMTT-V12; M2,

MMTT-V15 and M3, MMTT-V13 (Fig. 3,

Table 3).

Description

M1 (Fig. 3A-B, Table 3): The teeth are

trilophodont and have three roots. The three

transverse lophids are almost the same size. A

prominent cingulum is present on the anterior

wall. The lophids are heavily worn and partly

damaged.

M2 (Fig. 3C-D, Table 3): The teeth are

bilophodont and nearly quadratic in occlusal

outline and have three roots. The two

transverse lophids are of equal width. The teeth

are worn and damaged; nevertheless the labial

and posterior cingulae are clear. Both the

protoloph and metaloph have protocristids.

M3 (Fig. 3E-F, Table 3): The teeth are

bilophodont and have three roots. Right M3 is

very well preserved and has cingulum on the

anterior, labial, and posterior walls. The

protoloph is narrower than the metaloph and

has a less pronounced protocrista with a medial

extension.

Fig. 3 Deinotherium giganteum, Abkhare village, Varzeghan District, NW Iran. Upper molars,

occlusal views. A, V16: right M1 and B, V12: left M1. Scale bar: 40mm; C, V11: right M2 and D, V15:

left M2. Scale bar: 21 mm; E, V14: right M3 and F, V13: left M3. Scale bar: 21mm

TABLE 3 Measurements of Deinotherium giganteum upper Molars, Abkhare village, Varzeghan

District, NW Iran

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9

Comparison

The basic morphologies of the molar teeth of

species of Deinotheriidae are similar and the

main differences are in their size. Previous

studies have shown that the size of deinotheriid

dental and skeletal elements increased

gradually throughout the Miocene (Huttunen,

2002). The Deinotherium material from

Abkhare belongs to a single and very large

individual. Figure 4 shows the bivariate plots

of length and width of the molar teeth of the

Varzeghan specimens and other material from

Middle to Late Miocene localities in Europe.

The size of the teeth of Deinotherium from

Varzeghan as figured in the plots is larger than

the material from Pontlevoy (MN5), Sansan

(MN 6) and Montredon (MN10) in France,

Deinotheriensande and Eppelsheim (MN 9) in

Germany, Pannonian basin in central Europe

(MN7-10) and Late Miocene localities in

Greece, Bulgaria and Hungary. Although

slightly smaller, the Deinotherium molars from

Varzeghan are comparable in size to the teeth

of Deinotherium giganteum from Obuhovka

(MN 12) of southern Russia (Bajgusheva and

Titov, 2006), but are smaller than teeth of D.

gigantissimum.

6 Biochronology and paleoenvironment

Although the new fossil localities at

Varzeghan (and Ivand) are separated from the

Maragheh Fm. by the Anatolian transform

(Fig.1), but because of their geographical

proximity (ca. 110-150 kilometres), faunal

comparison and correlation of these fossil sites

are of interest. Like other typical Turolian

localities of the old world, hipparions are the

most abundant material in the Varzeghan

district.

Among the known hipparionine horses

from Maragheh medium sized hipparion

species such as Hipparion moldavicum and

medium to large sized species such as H.

prostylum are the most common species. Both

these species occur in the Middle Maragheh

biostratigraphical interval, which is most likely

8 to 7 million years old (Bernor, 1985; 1986).

Species of similar size and related to H.

prostylum, such as H. dietrichi, have a wide

distribution in the eastern Mediterranean

localities and are recorded from many

localities in the Balkans and Anatolia.

Occurrence of this latter species is mostly in

MN12 (8-7 Ma), but it is also reported from

MN 11 localities. H. moldavicum is also very

common in the northern Black sea region and

is recorded from MN11-12 while being absent

from the Balkans (Koufos and Vlachou, 2005).

The only occurrence of H. moldavicum in

Anatolia is from Akkaşdaği at 7.1 Ma (Koufos

and Vlachou, 2005).

The Ivand locality is another new Late

Miocene fossil site in northwest Iran. The

Varzeghan district is closer to this locality (ca.

50 kilometers) than to Maragheh. Hipparionine

horses are also abundant in this area, but only a

large sized species very similar to H.

brachypus is recorded from Ivand and the rest

of the material has not been studied yet. The

occurrence of H. brachypus at Pikermi and

Samos (Greece), Akkaşdaği (Turkey), and

Hadjidimovo (Bulgaria) is in the middle

Turolian or MN12 (Koufos and Vlachou,

2005). Consequently a Middle Turolian age

(ca. 8-7 Ma) can be inferred for the levels

containing Hipparion sp. large at Ivand. This is

also consistent with the age span of the Group

1 hipparionine horses of Bernor (1985).

Although Deinotherium giganteum has a

broad stratigraphical range from the Middle

Miocene to the Pliocene (18-2 Ma), given the

deinotheriid trend toward increasing size

during the Miocene (Huttunen, 2002) and the

very large size of the deinotheriid material

from Varzeghan, a late Miocene age of this

locality is evident.


10

Based on the occurrence of

Deinotherium at Varzeghan, the locality can be

correlated with the Upper Maragheh

biostratigraphical interval, since the only

known Deinotherium from the Maragheh

Formation is recorded from the upper intervals

of this formation (K1 locality of Erdbrink et

al., 1976; identical to MMTT locality 31 of

Bernor, 1986). Absolute ages obtained from

Zircon fission track and K/Ar dating of

pumicites from some higher levels of the

Maragheh Formation implies an age of 7.4 Ma

for the uppermost part of this formation

(Bernor, 1986), which can be also assigned to

the Deinotherium-yielding upper levels of the

Maragheh Formation. D. giganteum is also

represented by several postcranial elements

from the Ivand locality. Based on the presence

of H. brachypus at this locality a Middle

Turolian age has been proposed for the

locality.

In conclusion, comparisons of the fossil

material from Varzeghan with other localities

in northwest Iran and the chronological

occurrences of similar taxa, suggest a middle

Turolian age (MN 12 equivalent) for the fossil

sites near Varzeghan.

The morphology of the deinotheriid

cheek teeth, which are lophodont and

brachyodont, indicates a folivorous diet and

implies a forested environment. However, the

very large size of the animal suggests a

landscape more open than that of a forest

(Poulakakis et al. 2005). Therefore, the

presence of this species in the new Late

Miocene fossil localities in northwest Iran

indicates either woodland or an open forest as

the predominant environment.

7 Conclusions

The area near Varzeghan in northwest Iran has

produced several new fossil localities. As a

consequence, our knowledge of Late Miocene

mammals in Iran is no longer restricted to the

fauna of the Maragheh Formation.

Hipparionine horses and deinotheres are

among the fossil material so far collected from

this area. The study of mandibular fragments

of hipparionine horses has resulted in

identification of two hipparion species based

on size and dental morphology. These fossil

materials belong to small to medium sized

hipparion species such as Hipparion

moldavicum and to medium to large sized

species such as H. prostylum or H. dietrichi.

According to the age and distribution of

these latter taxa in the eastern Mediterranean

and northern Black Sea regions and correlation

of the new localities with Maragheh and Ivand,

a Middle Turolian age (MN 12 equivalent) is

proposed for the sites near Varzeghan. The

presence of Deinotherium giganteum is very

important for correlation of these new

localities to biostratigraphical intervals of the

Maragheh Formation and Ivand and has proved

the existence of semi-open environments in the

region. More systematic collection of fossils at

Varzeghan and study of additional material

such as the maxillary cheek teeth of the

hipparionine horses will enhance our

understanding of these new localities and their

relationships with contemporaneous faunas in

northwest Iran and the adjacent Caucasus

region.

Acknowledgements

Iran’s Environment Protection Organization

(formerly Department of Environment)

facilitated this research and the field work. We

appreciate the pioneering work of Dr Z.

Purabrishemi, Tabriz University, in the

Varzeghan district. We are also grateful to

Prof. J. Barry, Peabody Museum, for

improving the text of this paper and his

constructive comments. MMA thanks the

support from Academy of Finland and RHOI

project.


11

Fig. 4 Scatter plots of M1, M2 and M3 length against width in Deinotherium giganteum from Abkhare

village, Varzeghan District, and deinotheres from twelve European localities (data from Huttunen,

2002; Poulakakis et al., 2005; Bajgusheva and Titov, 2006)

× Deinotherium giganteum, Varzeghan; - D. giganteum, Sansan (MN6); + D. giganteum, Eppelsheim

(MN9); D. giganteum, Hungary (L. Miocene); D. giganteum, Obuhovka (MN11); ▲ D.

giganteum, Pontlevoy (MN5); Δ Prodeinotherium, Austria (Pannonian); D. giganteum, Siteia

(MN11); D. giganteum, Bulgaria (L. Miocene); ● D. giganteum, Deinotheriensande (MN9); ○ D.

giganteum, Montredon (MN10); ■ D. giganteum Austria (MN7-10); □ D. gigantissimum, Romania (L.

Miocene)


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