FEEDING ECOLOGY OF THE EARLY PLEISTOCENE GERAKAROU HERBIVORE MAMMAL COMMUNITY (MYGDONIA BASIN, GREECE).

1. *Corresponding author: evi0812@yahoo.gr
FEEDING ECOLOGY AND PALEOENVIRONMENT OF THE EARLY PLEISTOCENE GERAKAROU
HERBIVORE MAMMAL COMMUNITY (MYGDONIA BASIN, GREECE) AS INFERED BY DMTA ANALYSIS
Introduction
E. Alifieri1,* , E. Berlioz2, D.S. Kostopoulos1 and G. Merceron3
1School of Geology, Aristotle University of Thessaloniki, Thessaloniki, Greece
2TRACES (UMR5608), CNRS, Université Toulouse 2 Jean Jaurès, Toulouse, France
3PALEVOPRIM, CNRS and University of Poitiers, Poitiers, France
The Early Pleistocene in Europe was characterized by glacial-interglacial cycles, which shifted the global trend to colder temperatures,
linked with habitat openings during the glacial events (Kahlke et al., 2011). Understanding the impact of Early Pleistocene’s climatic
oscillations on the vegetation of local environments could further help with explaining the dispersal of Homo, especially in southeastern
Europe.
Important insights into the environmental conditions of that time and area can be provided by studying the fairly rich Early Pleistocene fossil
site of Gerakarou-1 (2–1.8 Ma; Konidaris et al., 2021), situated in northern Greece (Mygdonia Basin). This can be done by using the
locality’s herbivore assemblage, since the herbivore species’ diet is reflective of the vegetal resources in their habitat. Using in particular,
Dental Microwear Texture Analysis (DMTA) on the fossil teeth, a proxy that corresponds to the diet of the animal during its last few days or
weeks of life (Winkler et al., 2020).
The aim of this study was to draw conclusions on the dietary preferences of the extinct artiodactyls and the equid from Gerakarou-1 by
using extant species with known dietary habits as a reference. Both Scale-Sensitive Fractal Analysis (SSFA) parameters and
complementary Surface Texture Analysis (STA) parameters were used. It allowed us inferring the paleoenvironment of this early pleisto
locality.

2. Gerakarou-1
Fossil material
Method
Figure 1: Geological map of the
Mygdonia Basin (modified after
Konidaris et al., 2015; data from
Koufos et al., 1995).
• First discovered in 1978 and excavated by Emeritus Prof. G. D. Koufos.
• Stratigraphy: upper part of the Gerakarou Formation (Koufos et al., 1995).
• Age: ~2–1.8 Ma (Konidaris et al., 2021).
• Fauna: typical “Villafranchian” faunal elements, and more recent taxa. Artiodactyls make >50% of the faunal assemblage.
• Previous paleoenvironmental reconstruction: fluvio-terrestrial environment (Konidaris et al., 2015; Fig.1).
Comparative Material*
BOVIDAE
Bovidae indet. (N=1)
Leptobos (N=1)
Antilope (N=3)
Gazella (N=31)
EQUIDAE
Equus (N= 14)
CERVIDAE
Croizetoceros (N=33)
Eucladoceros (N=5)
SUIDAE
Sus (N= 1)
Dental Microwear Texture Analysis (DMTA)
Scanned on shearing facets
of the lower or upper molars
- 5 SSFA parameters (Scott et al., 2006):
We considered 5 variables from Scale
Sensitive Fractal Analysis (SSFA)
especially anisotropy (epLsar) and
complexity (Asfc).
- 5 STA parameters (Calandra et al., 2012):
And 5 variables from Surface Texture
Analysis (STA), especially mean dale area
(Sda) and valley material portion (Smr2).
194 wild shot specimens of 7 extant
species European bovids and cervids
covering the dietary spectrum from
browsers to grazers.
*(Berlioz et al. 2021; Hermier et al., 2020; Merceron et
al., 2014; 2021)
Asfc
epLsar
Smr2
Sda

3. Figures 2 and 3 : Bivariate plots of Asfc vs epLsar and of Smr2 vs Sda for the extant (a) and fossil (b) taxa. Each point represents the mean, except for A. koufosi, L. cf. etruscus and
Bovidae indet., for which each point represents the values of one facet. Uncertainty crosses represent the 95% confidence interval of the standard error.
SSFA Parameters:
→ High Asfc =
brittle/hard food items
(ex. A. alces; Fig.2a).
→ High epLsar =
abrasive (ex. B.
taurus; Fig.2a) or
tough foods (ex. C.
capreolus; Fig.2a).
ISO Parameters:
→ Smr2= opposite trend
to Asfc.
→ High Sda= increase of
seed consumption only
(not for all hard items,
ex. C. elaphus; Fig 3a)
0 0 5 0 5 6 0 5
As c
00 5
0
00 0
0
00 5
0
0060
0
00 5
0
00 0
0
ep
sar
A
Anisotropy
(epLsar)
Complexity (Asfc)
Fossil
Figure 2b
82 8 8 85 86 8
S r2
00
50
200
250
00
S
a
F
Close
dale
area
(Sda)
Valley material portion (Smr2)
Fossil
Figure 3b
Close
dale
area
(Sda)
Valley material portion (Smr2)
Extant
Figure 3a
Anisotropy
(epLsar)
Complexity (Asfc)
Extant
Figure 2a
E. altidens
E. tegulensis
C. ramosus
gerakarensis
L. cf. etruscus
A. koufosi
G. bouvrainae
Fossils
Bovidae indet.
Legend
Alces alces
(N=44)
Capreolus
capreolus (N=18)
Rupricapra
rupicapra (N=21)
Cervus elaphus
(N=21)
Bison taurus
(N=24)
Bos bonasus
(N=44)
Ovis ammon
(N=22)
Extant
Grazer
Mixed Feeder
Browser
Results

4. Gerakarou-1 inferred paleoenvironment:
• The diets of the herbivorous ungulates in
Gerakarou-1 show a dominance of mixed feeders.
• Browsers are slightly more frequent than grazers.
→ Mosaic habitat: open forest / savanna woodland
environment with temperate towards warm
conditions,
This is on agreement with previous studies (Kahlke et
al., 2011).
Discussion
Grazers Mixed feeders Browsers
Figure 4: The interrelations of the dietary categorizations for
the fossil species in Gerakarou-1 (Icons by Zimices, CC BY 3.0
https://creativecommons.org/licenses/by-sa/3.0/).
Inferred diets:
The unique specimen of S. strozzii (n=1) was a mixed feeder
(intermediate to low epLsar), engaging perhaps in rootling behaviors (low H9).
Bovidae indet. (n=1) was a mixed feeder, that also incorporated seeds in
its diet.
L. cf. etruscus (n=1) was a frugivore-browser, eating foliage, fruits and
bark.
A. koufosi was a mixed feeder, also consuming a considerable amount of
seeds, and/or bark.
G. bouvrainae was a mixed feeder, that had almost no hard components
in its diet.
E. tegulensis was a browser including a high amount of hard items such
as seeds and bark. It had the most variable diet among the fossil.
C. ramosus gerakarensis was the most grazing mixed feeder, if not a
grazer.
E. altidens was probably more of a mixed feeder than a grazer.
Fig. 4

5. Summary
References
• The DMTA showed most of the extinct ungulates in Gerakarou-1 were mainly mixed feeders.
• The addition of Surface Texture Analysis (STA) parameters like mean dale area (Sda) and valley material portion (Smr2), apart from
Scale Sensitive Fractal Analysis (SSFA) ones, helps better distinguish the dietary groups.
• Gerakarou-1 had a mosaic environment of open forests-savanna woodlands, with temperate towards warm conditions.
• Berlioz et al., 2021. Palaeogeogr. Palaeoclimatol. Palaeoecol. 110754.
https://doi.org/10.1016/J.PALAEO.2021.110754
• Calandra et al., 2012. J. Hum. Evol. 63, 85–98.
https://doi.org/10.1016/j.jhevol.2012.05.001
• Hermier et al., 2020. Geobios 61, 11–29.
https://doi.org/10.1016/j.geobios.2020.06.006
• Kahlke et al., 2011. Quat. Sci. Rev. 30, 1368–1395.
https://doi.org/10.1016/j.quascirev.2010.07.020
• Konidaris et al., 2021. Quaternary 4, 1.
https://doi.org/10.3390/quat4010001
• Konidaris et al., 2015. Comptes Rendus - Palevol 14, 353–362.
https://doi.org/10.1016/j.crpv.2015.05.004
• Koufos et al., 1995. Geobios 28, 243–249.
https://doi.org/10.1016/S0016-6995(95)80171-5
• Merceron et al., 2021. Palaeogeogr. Palaeoclimatol. Palaeoecol. 562,
110077. https://doi.org/10.1016/j.palaeo.2020.110077
• Merceron et al., 2014. For. Ecol. Manage. 328, 262–269.
https://doi.org/10.1016/j.foreco.2014.05.041
• Ramdarshan et al., 2017. Palaeontol. Electron.
https://doi.org/10.26879/762
• Scott et al., 2006. J. Hum. Evol. 51, 339–349.
https://doi.org/10.1016/j.jhevol.2006.04.006
• Winkler et al., 2020. Palaeogeogr. Palaeoclimatol. Palaeoecol. 557.
https://doi.org/10.1016/j.palaeo.2020.109930