Native Greek horses: from man- and fish-eaters BC to DMRT3 gaiters in modern times

Theodore G. Antikas, DVM, PhD

Assoc. Prof., Aristotle Univ. of Thessaloniki (ret.); Visiting Onassis Prof., U of Washington Dept. of History

Abstract: ‘Mythical’ or prehistoric data re man eating horses in the 3rd or 2nd millennium BC reported by Greek
writers, historians and tragedians in classical times are elucidated by a new approach reflecting the taming,
domestication, zoogeography and evolution of Equus caballus in Greece, from Thrace to the Peloponnese.
Recent finds of fish eating horses in Central Macedonia of the 7th-4th centuries BC reveal that Herodotus was
reporting facts in his Histories thus rejecting the epithet ‘father of lies’ given to him by ancient and/or modern
writers. The ongoing collaborative research project of the Aristotle and Texas A&M Universities concerning the
presence of the DMRT3 gait mutation in autochthonous Greek horses suggests that it is fixed and homozygous
or heterozygous in 87% of 155 tested horses. The next phase of this study entails aDNA tests on horse skeletal
remains dating from the 16th to the 4th century BC from archaeological finds, so as to confirm the presence of
the mutation in ancient horses. It is noteworthy that gaiting athletic and war horses were depicted on vases,
coins, frescoes, reliefs and statues dating from the Mycenaean to the Classical, Hellenistic and Roman periods.

I. Man eating horses: Myth or Myth-history?

It is our belief that factual events in stories written by ancient writers or painted on vases by artists were
thought to represent unreal or mythical data. Less sceptical minds however, begin to realize that many mythical
events of the past were simply historical facts to which oral tradition, poets and tragedians had added mythical
elements to render them “heroic” or “divine”. A typical example is Homer’s Troy, Mycenae, Pylos, Ithaca and
Argos described in the Iliad and the Odyssey. Homer’s heroes, horses and city-states had been considered
unreal, until the pick axe of archaeologists unearthed Troy, Mycenae and Pylos. As if this weren’t enough,
hundreds of Linear B’ tablets describing the number of chariots stored in palaces, and tombs containing the
cremains of heroes such as Agamemnon or Nestor, are now the delight of museum visitors.

Examples of fact misinterpreted as myth are also found in the tragedies of Aeschylus1 and Euripides2. They both
reported the existence of man eating horses in what we call prehistory. In Aeschylus’ tragedy Glaucus was the
first son of Sisyphus, lived in the Boeotian city of Potnies, owned mares, and to render them more aggressive he
fed them human flesh. Alas when he took them to Iolkos in Thessaly to take part at the funeral games (probably
chariot races) organized by Acastus to honor his father Pelias, the mares turned crazy and devoured him. The
reason? Human flesh, the ‘food’ they were accustomed to, wasn’t available in Iolkos. Instead the mares were
given local hay. Another Boeotian myth claimed that this was not the reason the mares went crazy; instead, it
was Aphrodite who turned angry with Glaucus and punished him because he would not allow his mares to
reproduce. Note that Glaucus was worshipped in Corinth as hero, the locals called him Taraxippos (horse
terrorizer) and claimed he scared the horses at the Isthmian and the Olympic equestrian events. Jockeys and
charioteers used to sacrifice and offer libations to Taraxippos before racing to appease the hero-daemon3,4.

Heracles, a worshipped Pan-Hellenic hero thanks to his twelve athla (labors), was involved in a very interesting
myth reported by Euripides, Aelian5 and Diodorus6. The eighth labor of Heracles was to steal the man eating
mares of king Diomedes of Thrace and bring them to Argos. A black-figured kylix c. 510 BC at the Hermitage
depicts his bravery much earlier than Euripides (Fig. 1). The hero is holding a stallion and at the same time
threatens the animal with his club. The horse is trying to escape, and out of his mouth are the remains of a man
(Abderos or Diomedes). Well, it seems that horses were meat eaters probabaly fond of human flesh in the 2nd
or 3rd millenium BC. In one version of the myth, Heracles brought along many heroes to help him. They took
the mares but were chased away by Diomedes and his men. Heracles was not aware that the horses, Podagros
(the fast), Lampon (the shining), Xanthos (the blond) and Deimos (the terrible), were tethered to a bronze
manger because they were crazy and uncontrollable due to their unnatural diet consisting of human flesh.

1
Aeschylus, Glaucus Potnieus, passim
2
Euripides, Alcestis 480 ff.
3
Pausanias, Touring Greece VI.20.19, IX.8.2
4
Plinius, Natural History 25.94
5
Aelian, On Animals 15.25
6
Diodorus, Library of History 4.15.13
Heracles left his friend Abderos in charge of the horses while he fought Diomedes but on his return to the
stables he realized Abderos had been eaten by them. In revenge, Heracles fed Diomedes to the mares and then
founded the city of Abdera next to the boy's tomb. This city--where Democritos was born--exists until today in
western Thrace. In another version, Heracles stayed awake so that he wouldn't have his throat cut by Diomedes
in his sleep, then cut the chains binding the horses and scared them onto high ground. When king Diomedes
returned, Heracles killed him and fed the body to the mares to calm them. Both versions conclude that eating
human flesh calmed the horses. Once subdued, Heracles brought them back to King Eurystheus who dedicated
the horses to goddess Hera, who allowed them to roam freely around Argos as they were permanently calm.
Eurystheus ordered the horses taken to Olympus to be sacrificed to Zeus. Diodorus Siculus7 writes that when
the crazy horses were brought to Eurystheus, he dedicated them to Hera, they roamed freely to Thessaly, and
their breed continued down to the reign of Alexander the Great producing the famous Macedonian cavalry.

It is tempting to think that if modern horse, cattle or chicken breeders had taken some time to study Greek
mythology, they might have prevented zoonoses such as BSE (mad cow), H1N1 and other ailments. Such a
postulate may sound exaggerated but it is logical: the crazy mares of Diomedes symbolize nothing more than
the fact that ‘savage’ equids (not domesticated) had reached the Greek mainland from central Asia through
Thrace, an area joining Asia to Europe. Following the ‘heroic’ labor of Herakles the Thracian mares were tamed
(domesticated) and taken by sea or land (moved) to Argos long before the Trojan War. Finally, the horses were
dedicated to a female deity and were left to roam freely on the Greek mainland. In Troy we note hundreds of
war horses8,9: from Mycenae and Sparta (Agamemnon’s Aethe, Menelaus’ Podargos, Eumelus’ and Diomedes’
chariot mares); from Thessaly (Achilles’ Balios, Xanthos, Pedasus); from Crete (Meriones’ chariot horses); from
Thrace (King Rhesus’ chariot mares); and from Troy in Asia Minor (Hector’s mare Podarge). Eons later their
descendants reach Macedonia to breed cavalry horses. Ergo: the myths on man eating horses are symbolic and
explanatory at the same time. The primary lesson taught by them is that Man’s hubris of feeding meat to
herbivorous animals is a grave insult to mother Nature and will inevitably lead to tragedy and disaster. In a
very subtle manner the myth-history of Glaucus’ and Diomedes’ man eating horses explains the zoogeography,
the movement, the domestication and last but not least, the expansion of Equus caballus on the Greek mainland
during the late third or early second millennium BC. Moreover, there is a hard lesson to learn from the Greek
tragedies: people have reverted to eating horse meat for fear of Bovine Spongiform Encephalopathy. I call
innocent horses exported or stolen to supply the kitchens of consumers ’from stable to table.’

II. Fish eating horses: Was Herodotus lying?

In 2004 the pick axe of archaeology unearthed five horses and two dogs dating to the 7th-4th centuries BC
buried in a human cemetery found at Sindos, near Thessaloniki. A zooarchaeological study on these important
finds which reflect the Macedonian and Mycenaean10 habit of burying heroes with their horses and dogs has
been published11. The preservation of the five horse skeletons was so impressive (Fig. 2) that prompted the
research team to send samples for stable isotope analyses to two laboratories which came up with more or less
identical and surprising results (Fig. 3a). Namely, the 13C, 15N values and C/N ratios indicate that at least 20%
of the horses’ feed consisted of protein of fish origin12. As to the fish species Herodotus called papraces and
tilones, the labs have gone as far as to identify the species found until today in the Axios and Aliakmon rivers
close to Sindos (Fig. 3b). More analyses are necessary to compare the equine data to those of herbivorous
(cattle, capra/ovis) and omnivorous animals (canines, felines) from the same time period and the same region,
i.e., the Macedonian plains, but this is part of a future research project due to lack of funds.

Herodotus has been criticized for including “exaggerated stories” in his Histories, however recent studies by
archaeologists, archaeozoologists, geneticists, paleobotanists, and physicists have found elements of truth in his
exaggerations. As surprising Herodotus’ statement on fish eating horses may seem, a less skeptical approach
should lead modern scholars into considering his text from a new scientific angle.

7 Diodorus Siculus, Historic Library 4.15.3

8
Homer, Iliad, 23; see also his long list of horses owned by Greek and Trojan heroes
9
Antikas TG, Hippoi and Hippologia, To Tetarto Editions, Katerini 2012
10
Homer, Iliad, 23.171-74
11
Antikas TG, Fish Eating Horses in Macedonia, 5th c. BC: Was Herodotus Right? Veterinarija ir Zootechn. 44(66):31-37, 2008
12 I. Maniatis, National Ctr. For Scientific Research “Democritos”, Archaeometry Lab Report, Athens 2004
Firstly, one has to wonder what reason led four ancient writers to mention man or fish eating horses. None of
them was known to be a liar. Secondly, why should the perissodaktylon (odd-fingered) Equus need four sizable
canine teeth, if it were purely herbivorous, must be answered. Last but not least, we should consider whether
Herodotus, an invitee at the Macedonian court at Aegae, just 40 km from Sindos, should refer to “fish eating
horses and other pack animals” unless he witnessed or heard from locals that this was a custom. In conclusion,
Herodotus does not seem to have recorded myths in speaking of fish eating horses in the plains of Central
Macedonia13. Just as Homer before him was not referring to a mythical Troy but to a real citadel, Herodotus was
seemingly telling a story in his capacity as the father of history—not as a father of lies.

III. Origins of the Greek horse breeds and the DMRT3 gait mutation

Linear B’ scripts from Knossos dating to the second millennium BC and decoded by Chadwick and Ventris have
brought to light two ideograms: a-to-ro-qo (άνθρωπος) and i-qo (ίππος, hence equus). Horse skeletal remains
have been unearthed in Archanes-Crete, Dendra-Argos, Nemea-Corinth and Marathon-Attica. Linear B’ tablets
stating the number of chariot wheels stored in the palaces of Knossos and Pylos date to the 15th-16th c. BC, long
before the Trojan War. Zooarchaeologists may debate the origins of ancient horses, however there is data in
written sources that may help in suspecting some origins: (a) King Nestor of Pylos, after his war with the Eleans
brings 300 xanthes (chestnut) mares-in-foal back home14; (b) Agamemnon owns the fast chariot mare Aethe
from Sikyon in the Argolid15; and offers twelve victorious chariot horses of Argos to Achilles to apease him16;
(c) Eumelos owns mares bred in Pieria17; (d) Hector and Rhesus own chariot horses bred in Thrace and Asia
Minor; (e) Meriones owns Cretan horses. As the Trojan War was fought in the mid-13th century BC, i.e., one
century after the battle of Qadesh between Ramses and Muawatalli, I don’t see how we can ignore the origins of
Greek, Egyptian or Hittite horse breeds. In the archaic and classical eras (680 BC to 241 AD) many vase scenes
depict gaiting war horses (Figs. 4-5); and race or chariot horses at the festivals of Olympia, Pythia, Isthmia and
Nemea (Figs. 6-7)18. Interestingly, all these horses are gaiters thus forcing the postulate that the DMRT3 gait
mutation was present in their genome some 2,700 years ago. Also, that this important gait was preferred by
riders and drivers for speed in racing and charioteering, as well as in cavalries consisting of war horses. In
classical and hellenistic times, Macedonian kings minted coins depictig gaiters: Alexander I the Philhellene, 498-
454 BC, who rode to Plataies in 472 (Fig. 8); Perdikas II, 448-413 (Fig. 9); Archelaos, 413-399, first Macedonian
to win a wreath at Olympia’s tethrippon race in 408 (Fig. 10); Pausanias, 399 (Fig. 11); Aeropos, 399-396 (Fig.
12); and Philip II, 359-336, who won three wreaths at Olympia’s keles, tehrippon and synoris races (Fig. 13). It
is also noteworthy that the Thracian Getae and the Skythians of King Ateas (Fig. 14) who were defeated by
Philip in 339 BC had been riding gaiters since the times Chersonesos was governed by Miltiades in the sixth
century BC. In fact, Theopompus and Justin inform us that after defeating the Skythian king Ateas, Philip
brought back to Greece 20,000 “well fed” Skythian mares and 20,000 women and children to populate
Macedonia19. It is postulated that most Macedonian kings may have used gaiters in their cavalry. If one
considers that Philip II rode some 25,000 km in his campaigns and Alexander III more than 45,000 km riding
bareback from Pella to Egypt and Pakistan, it would be no surprise to this author that everyone, king or soldier,
would prefer to ride gaiters so as to save their gluteus muscles from turning into raw meat.

At the same time period, tribal Thracian Kings in the northwest who fought the neighboring Macedonians
sporadically but with little success as they were eventually defeated by Philip II and later by his son Alexander
the Great, also rode gaiters. Among them were the Bisaltai (Fig. 15) and the Thracian kings Sparadokos (Fig.
16) and Seuthes (Fig. 17) in the mid-fourth century BC.

13 Herodotus, Histories 5.16: Their horses and other pack animals they feed on fish which are so abundant… that when they
open the trapdoor and let down an empty bucket on a rope, they have only a minute to wait before they pull it up again, full.
The fish are of two kinds… papraces and tilones.
14 Homer, Iliad 11.680 ff.
15 Homer, Iliad 23.294
16 Homer, Iliad 9.123-24, 9.265-66
17 Homer, Iliad 2.765
18 Antikas TG, Ολυμπικά Ιππικά και Γυμνικά Άθλα. To Tetarto editions, Katerini 2012
19 Theopompus frag. 162, Justin 9.2, 9.3
The preference to use gaiters in cavalries was followed by Alexander’s successors. Coins minted by Eukratidas
in the second (Fig. 18), and Azes (Fig. 19) in the first century BC, depict gaiters. As to Alexander himself, after
329 BC he replaced his exhausted cavalry with Persian, Parthian, Bactrian and Indian men and horses. His
example was naturally followed by several of his Epigonoi (successors) in the Middle East, Asia and Egypt.

In the post-Christian era, King Gondophares (Gr: Ινδοφέρρης) minted coins depicting gaiters (Fig. 20). It was he
who took over the Kabul, the Punjab and the Sindh regions from king Azes, the Indo-Scythian successor of the
two governors Telephos and Hippostratos Alexander had left behind in the third century AD. As to Byzantine
kings as Justinian (Fig. 21), saints as Theodoros Tyron and Stratelates (Fig. 22) or laymen (Fig. 23), we can
only theorize that their gaiters were crosses of Greek to Eurasian breeds. It is interesting that the use of gaiting
horses was later followed both in the east and the west: many English, French, Swiss, Italian and Μamluk
manuscripts of the 14th, Chinese mabuscripts of the 15th, and Epirote empbroideries of the 17th century AD
depict gaiters. It is hard to find sufficient written data in the Dark Ages in Europe or furing the rise of the
Ottoman, Arab, Mamluk and Mongol chieftains in the east. Pictorial evidence however, may convince scientists
that the gait mutation was fixed on horses both in the East and the West. Such gaiters show up in ecclesiastical
texts, paintings, textile, carpets, icons and other artifacts. As to which equine breeds gave birth to these gaiters
it remains an enigma. Will aDNA testing tell us? I wonder as there is no sufficient DNA testing done thus far.

IV. The research project of DNA and aDNA testing native Greek horses

A two-year research project (1998-2000) run by the Department of Reproductive Physiology of the College of
Agriculure at the Aristotle University of Thessaloniki to define the phenotype of indigenous Greek horse breeds
established that five breeds, from Mt. Pindus in Epirus to Mt. Ida in Crete, were gaiters. Their population had
declined significantly in the late 20th century due to exports for slaughter, a sad horse reality: in 1950, Greece
had ca. 280,000 horses, 170,000 mules and 450.000 donkeys (total, 900,000). In 1980, 105,000, 115,000 and
230,000, respectively (450,000); and in 2000, 34,509, 41,319 and 88,837 (~165,000), respectively. In a hard
effort to save Greece’s native horse breeds from extinction the project had registered on a data base 4,000
horses from Epirus to Crete (Fig. 26), defined six indigenous breeds, and published its finds in 200020.

Thankfully, the horse genome was mapped successfully a few years later and so did the outstanding find of the
DMRT3 mutation on chromosome-23 of the horse. These two major discoveries led to an inter-university,
interdisciplinary project led by the Hellenic Association of Traditional Equitation (HASTE) in collaboration with
the Aristotle University and the Texas A&M University’s Dept. of Veterinary Intergrative Bioscience headed by
Prof. E. Gus Cothran. From 2011 to the present, blood and hair samples of 177 native horses have been tested
as to their origins (Tables I-II) and 155 as to the presence of the DMRT3 gait mutation (Table III).

V. Conclusions

The DNA tests and statistics contained in the tables below have come as no surprise: in short, they indicate that
(a) the origins of Greece’s native horses include three Asiatic breeds, namely AT-Akhal Teke, KU-Kurdish and
TU-Turkmen; (b) the majority of horses from five native Greek breeds are either homozygous or heterozygous
for the DMRT3 gait mutation. The next phase of this pioneer project should be running aDNA tests in ancient
horse skeletons found in archaeological excavations from Dendra and Nemea in the Peloponnese (16th-15th c.
BC) to Sindos, Polykastro and Therme in Central Macedonia (7th-4th) and analyzed by zooarchaologists21. The
ambitious purpose of this future research project is to discern whether the DMRT3 mutation has been fixed on
the genome of Greek horses during the last four or five millenia. It is my hope that Poseidon Hippios and Athena
Hippia will be favorable to this ambitious project presently suffering from lack of necessary funds.

20T. A. Alifakiotis, transl. by T. G. Antikas, The Indigenous Horses in Greece. Ministry of Agriculture (ed), Athens 2000
21T. G. Antikas, Fish Eating Horses in Central Macedonia, 5th c. BCE: Was Herodotus Right? Veter. Zootechn. 44(66):31-37,
2008; see also The Honour to be Buried with Horses: From Mycenaean Nemea to Macedonian Vergina. In: Les Equides dans le
Monde Mediterraneen Antique. Arnelle Gardeisen (ed.), CNRS 143-51, France
Acknowledgements: grateful thanks to the 117 owners from Crete, Epirus, Macedonia, Thessaly, Thrace and
the Peloponnese who were kind to help us get blood and hair samples from their native horses. Thanks are also
due to Dr. E. Gus Cothran for his invaluable contribution without which this project would have beeen a utopia.

Table I. Match frequency to the closest native horse breed after DNA testing¹

Greek Horses 3 matches 2 matches 1 match 0 match

Region Tested ♂ ♀ ♂ ♀ ♂ ♀ ♂ ♀
Crete 73 5 3 8 12 16 14 7 6
Macedonia 54 2 1 6 15 10 10 6 6
Peloponnese 18 0 0 1 4 7 6 0 0
Nafpaktos 11 0 1 3 1 1 4 1 0
E. Thrace 10 0 0 0 1 1 6 1 1
Thessaly 4 0 0 1 0 3 0 0 0
Epirus 3 0 1 1 0 0 0 1? 0
Cephalonia 3 0 0 1 1 0 0 0 1
Attica 1 0 0 0 0 0 1 0 0
Total-Percent % 177 13-7.345% 55-31.073% 79-44.633% 30-16.949%
¹ the Asiatic breeds AT, KU, TU are considered quasi native due to close origins with Greek breeds since the 4th century BC

Table II. Indigenous breed frequency after DNA testing²

Greek Horses EM-EZ PD-TH AT-KU-TU Foreign

Region Tested Origins Origins Origins Origins
Crete 73 46 25 22 13
Macedonia 54 31 19 10 11
Peloponnese 18 10 3 4 1
Nafpaktos 11 0 8 5 1
E. Thrace 10 4 5 0 2
Epirus 3 2 2 1 0
Thessaly 3 4 1 1 0
Cephalonia 3 2 8 0 1
Attica 1 0 0 1 0
Total-Percent 177 99-55.932% 71-40.112% 44-24.858% 30-16.949%
² EM-EZ=Elis Mountain-Elis/Zakynthos; PD-TH=Pindos-Thessalian; AT-KU-TU=Akhal Teke-Kurdish-Turkmen

Table III. Frequency of DMRT3 presence after DNA testing

Greek Horses Homozygous or GG-Greek¹ Foreign Undetermined

Region Tested Heterozygous Gaiters Gaiters (test failed)²
Crete 73 67 56 11 6
Macedonia 49 49 37 12 0
Peloponnese 16 14 2 0 2
E. Thrace 10 1 1 0 0
Epirus 3 2 2 0 1
Cephalonia 3 1 1 0 1
Attica 1 1 1 0 0
Total-Percent % 155 135-87.097% 100-64.516% 23-14.838% 12-7.741%
¹ the term GG is given to 2nd and 3rd generation horses born in Greece regardless of their particular native breed

² inconclusive tests although clinical tests to check the actual kinetics of these twelve horses showed true gaiting
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