1. MESOWEAR ANALYSIS OF RUMINANTS FROM MELITZIA CAVE, SOUTHERN PELOPONNESE GREECE
Marianthi Tzortzi1*, George Iliopoulos1 ,Andreas Darlas2
1Department of Geology, University of Patras, GR-26504, Rio, Patras, Greece
2 Ephoreia of Paleoanthropology and Speleology, Greek Ministry of Culture and Sports, Ardittou 34b, 1636 Athens, Greece
*mtzortzi@upatras.gr
Melitzia Cave is located on the west coast of Mani Peninsula, Greece (fig. 1),
where several sites, containing deposits with cultural remains from the
Middle Palaeolithic to the Neolithic period have been found to date. The
excavation revealed numerus lithic artefacts, together with the strong
presence of modified skeletal remains, testifying the intense hominin
occupation of the cave. The site revealed frequent occupation of the cave by
hominin groups between 24.000 and 11.000 cal BP (Darlas and Psathi, 2016).
The excavation pit (fig.2) consists
of six squares of 1x1 m were
excavated, reaching a depth of
190cm and one of them reached to
the depth of 300 cm.
Moving from top to the bottom, 7
stratigraphical units were recovered,
each unit is inclined towards the
East with a difference of 25 cm, a
slight inclination (<10cm) towards
the South is also observed (fig.3).
Fig 1: Simplified geological map presenting the geological and tectonic setting of the area surrounding the
Melitzia
Fig 2: Excavation pit
I
II
III
IV
V
VIb
VIa
VII
Fig 2: Stratigraphic section of Th22
square
2. Occlusal
relief
Cusp
shape
y/x ≥ 0.25
valley between cusps ≤ 90°
0.25 > y/x ≥ 0.125
valley between cusps >90°
0.125 > y/x ≥ 0.05
0.05 > y/x > 0
y/x ≤ 0
Sharp with lens 12x
Sharp with naked eye at
20cm, round with lens 12x
Clearly round with naked
eye, length ≤ ½ cusp length
Clearly round with naked
eye, length > ½ cusp length
Platform present
hh
h
hl
l
fn
s
rs
r
rr
b
Using the Extented Dental Mesowear
analysis (Winkler and Kaiser, 2011)(table
1, 2), all the maxillar molars that belongs to
ruminants were studied.
Table 1: Scoring scheme for the extented mesowear method. Occlusal relief
categories ‘high-high’ (hh), ‘high’ (h), ‘high-low’ (hl), ‘low’ (l) and ‘flat-
negative’ (fn) and cusp shape categories were scored as ‘sharp’ (s), ‘round-
sharp’ (rs), ‘round’ (r), ‘round-round’ (rr) or ‘blunt’ (modified from Taylor
et al. 2016)
Overall, 35 teeth of 3 different ruminant taxa (Dama dama, Cervus elaphus, Capra
ibex) were categorized by evaluating the relative relief and sharpness of the upper
molar's cusp apices through digital photography with the aid of the software tpsDig2
2.17. From the available dental material, only the adult isolated teeth and tooth rows
bearing molars were sampled, whereas those in very early wear stages or extremely
worn or even taphonomically altered were discarded from the analysis. The inclusion
of M1, M3 fulfil the need of a larger dataset.
Occlusal
relief
hh h hl hh h hl hh h hl h hl l l l l hl l
Cusp
shape
s s s rs rs rs r r r rr rr s rs r rr b b
Mesowear
score
1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17
1cm
Table 2: Combination of occlusal relied and cusp shape in conjunction with Mesowear score by Winkler and Kaiser 2011
Material and Methods
3. Results
0 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17
Mesowear Score
C. ibex
D. dama
C. elaphus
C. elaphus
D. dama
C. ibex
C. elaphus
D. dama
C. ibex
C. elaphus
D. dama
C. ibex
C. elaphus
D. dama
C. ibex
C. elaphus
D. dama
C. ibex
I
IV
V
VIa
VIb
VII
o Mesowear data for each taxon and among
different units have revealed a dietary range from
browsing to a mixed feeding strategy.
o The vast majority of the analyzed sample belongs
to Cervus elaphus, mainly deriving from remains
of unit VI, dated to approximately MIS2. Overall
mesowear scores were ranging from 2 to 9 for the
upper molars in Cervus elaphus.
o The predominant mesowear pattern among fossil
samples is high cusp relief combined with rounded
sharp cusp apices (table 3).
o Most of the samples derived from unit Via and
unfortunately the samples from the units VII, VIb,
IV and I are one or two, so the results are
questionable. Although, a tentative dietary
assumption can be made using three or even one
molar (Kaiser et al., 2003 Rivals et al., 2017).
Species n hh% h% hl% l% ll%
Cervus elaphus 21 0 85.7 9.5 4.8 0
Dama dama 5 0 100 0 0 0
Capra ibex 9 0 77.8 22.2 0 0
Species n s% rs% r% rr% s%
Cervus elaphus 21 14.3 52.3 28.6 4.8 14.3
Dama dama 5 40 20 40 0 40
Capra ibex 9 11 44.5 33.5 11 11
Table 3: Summary of the mesowear data for the molars examined in this study
4. Discussion
• Our findings seem to suggest general mixed feeding traits
closer to browsing traits, as it was expected according to the
literature for the most abounded taxa, Cervus elaphus.
• The lack of a grassy cover is attested as none of the present
taxa seems to be specialized to grazing.
• According to Ackerman (2020) Cervus elaphus prefer a
browsing to fixed diet.
• According to Lister (1984) The Late Pleistocene Cervus
elaphus shows a wide range of mesowear, corresponding to
significant dietary plasticity.
• According to Rivals and Lister (2016) Cervus elaphus shows a
preference for mixed feeding and leaf browsing traits.
• In terms of average wear traits among the different units, it is
more accurate to conclude that there is no significant variation
throughout the stratigraphic sequence and therefore, throughout
time. Although, it is observed a progressive replacement of the
Dama dama by the Cervus elaphus.
• According to Darlas and Psathi (2013) the same replacement is
occurred at the Mani peninsula and is an evidence of the
change toward drier and colder climatic conditions.
• Kolendrianou et al. (2020) examine mesowear signal of the
ruminant population from Kalamakia Cave, a nearby area with
similar age to lowermost units of Melitzia Cave and
characterize the site as a partly wooded environment.
Overall, the dietary traits of the ungulates of the Melitzia Cave indicate an open partly
wooded environment, such as Mediterranean shrublands. Although, the progressive
replacement of a clearly browser taxon (Dama dama) by a more flexible one (Cervus
elaphus) points out the restriction of the Mediterranean forest at the transition of an
interglacial period of the lowermost units to a glacial period of the middle and upper
units.
5. Conclusion
References
• Ackerman, N., L. 2020. The history of mesowear: a review. PeerJ 8:e8519
• Darlas, A., Psathi, E., 2016. The middle and upper paleolithic on the western coast of the Mani peninsula (southern Greece). In: Harvati, K., Roksandic, M. (Eds.), Palaeoanthropology of the Balkans and
Anatolia, Human Evolution and its Context. Springer Science and Business Media, Dordrecht: 95–118
• Kaiser TM, Fortelius M. 2003. Differential mesowear in occluding upper and lower molars: opening mesowear analysis for lower molars and premolars in hypsodont horses. Journal of Morphology
258(1):67–83
• Kolendianou, M., Ligkovanlis, S., Maniakas, I., Tzortzi, M., Iliopoulos, G. 2020. The Palaeolithic cave of Kalamakia (Mani Peninsula), Greece: new insights on the palaeoenvironment using
microvertebrates and mesowear analysis of ruminant teeth. Heliyon 6 e03958
• Lister, A.M. 1984. Evolutionary and ecological origins of British deer. Proceedings of the Royal Society of Edinburg Vol. 82 B(4): 205-229
• Rivals, F., Lister, A.M. 2016. Dietary flexibility and niche partitioning of large herbivores through the Pleistocene of Britain. Quaternary Science Reviews 146: 116-133.
• Rivals F, Uzunidis A, Sanz M, Daura J. 2017. Faunal dietary response to the Heinrich Event 4 in southwestern Europe. Palaeogeography, Palaeoclimatology, Palaeoecology 473: 123–130
• Taylor LA, Müller DWH, Schwitzer C, Kaiser TM, Castell JC, Clauss M, Schulz-Kornas E. 2016. Comparative analyses of tooth wear in free-ranging and captive wild equids. Equine Veterinary Journal
48(2): 240–245
• Winkler DE, Kaiser TM. 2011. A case study of seasonal, sexual and ontogenetic divergence in the feeding behavior of the moose (Alces alces, Linné, 1758). Verhandlungen des Naturwissenschaftlichen
Vereins Hamburg 46: 331–348
The current study marks the first mesowear analysis contacted on three different ungulate taxa found
in the mammal assemblange of Melitzia Cave. The results indicate that there is no significant variation
through time and the environment could be characterized as open partly wooded. However, due to the
limited amount of eligible specimens for mesowear analysis the results mentioned herein cannot give
a more precise picture of the site’s palaeoenvironment and therefore more sophisticated method
should be applied in the future.