Abstract
The origins of horseback riding remain elusive. Scientific studies show that horses were kept for their milk ~3500 to 3000 BCE, widely accepted as indicating domestication. However, this does not confirm them to be ridden. Equipment used by early riders is rarely preserved, and the reliability of equine dental and mandibular pathologies remains contested. However, horsemanship has two interacting components: the horse as mount and the human as rider. Alterations associated with riding in human skeletons therefore possibly provide the best source of information. Here, we report five Yamnaya individuals well-dated to 3021 to 2501 calibrated BCE from kurgans in Romania, Bulgaria, and Hungary, displaying changes in bone morphology and distinct pathologies associated with horseback riding. These are the oldest humans identified as riders so far.
Human skeletal remains indicate the practice of horseback riding as early as the 30th century BCE.
INTRODUCTION
Multidisciplinary sources of evidence for earliest horsemanship
Using horses for transport was a decisive step in human cultural development. Trade and cultural exchange as well as conflicts and migrations leapt with the increase in speed and range provided by horsemanship. Archeological, archeozoological, and paleogenetic research into the beginnings of horse domestication and the initial expansion of domesticated horses (Equus caballus) has recently seen much progress (1, 2), as has our understanding of the appearance of horse-drawn fast chariots with spoked wheels ~2000 BCE (3).
However, information for earliest horseback riding so far is sparse (see section S1 for a detailed review). Possible bit wear in premolar teeth of horses from Botai (Kazakhstan) dating to <3500 BCE were extensively debated during the past three decades (4–6). Information from the Botai site such as horse demography, horse dung finds, potential paddock fences, or horse milk traces in pot shards (7, 8), as well as horse milk peptides in the calculus of Yamnaya individuals from Krivyanskiy 9 (Russia; ~3000 BCE) (9), suggests that domestication became widely established during the second half of the fourth millennium BCE. However, these do not provide direct evidence for riding.
Depictions from the Mesopotamian Ur III period shortly before 2000 BCE may be the earliest figurative evidence for riding, probably on a horse (rider seated forward, falling mane, and bushy tail hair) but possibly on an ass (Equus asinus) or ass-onager (Equus hemionus) hybrid (10). During the Old Babylonian period of the early second millennium BCE, indubitable images and literary mentions in cuneiform texts (11) prove that horses were ridden. However, it is apparent that organized cavalry was introduced not before the very end of the second millennium BCE (4).
While this provides a rough time frame and geographical setting, our understanding of how horsemanship developed between mid-fourth and early second millennium BCE in the Pontic-Caspian steppe and the Middle East is still vague. This time span also sees the first horse dispersals to the west and south (1, 2, 10, 12), the origins of modern horse breeds (1), the widespread introduction of cattle-pulled wheeled carts and wagons (13), and the Yamnaya (~3200 to 2500 BCE) expansions eastward to the Altai and Mongolia in the form of the Afanasievo culture (14) and westward into the southeast of Europe, coming to a hold at the Tisza river in eastern Hungary (15). Latest research into this event (Fig. 1) indicates its rapid accomplishment within one or two centuries just before and after 3000 BCE. Considering the vast geographical distances of 4500 km between the Tisza river and the Altai mountains, the absence of roads, and the small overall population sizes, it is difficult to envision how this expansion could have taken place without improved means of transport.
Fig. 1. Map of the Yamnaya and Afanasievo overall distribution.
Sites with individuals with skeletal markers for horsemanship are marked (black circles, Yamnaya; yellow circles, graves dated to other periods) (the background map is made with Natural Earth; free vector and raster map data at naturalearthdata.com; QGIS software). 1a, Strejnicu mound I grave 3 (I/3) grave in situ (photo credit: A. Frînculeasa, Prahova County Museum of History and Archaeology).
Yamnaya people had horses, as we know from their few settlements, where horse bones ranged widely from 1 to 2% up to 80% of animal bones, as well as from occasional horse bones found in kurgan fillings (fig. S1) [(16), pp. 150–157, (17)]. What we lack is archeozoological or artifactual evidence for their use, whether as livestock, as beast of burden or mount and draft animal, or as prestige good. However, the quantity and preservation of Yamnaya horse remains are insufficient for biomechanics studies regarding possible use. In addition, specialized riding tack is not essential for riding or can be made from perishable materials only, so its absence in findings is no proof against riding.
RESULTS
Osteological evidence for earliest horsemanship: Strejnicu mound I grave 3 and other Yamnaya individuals
Horsemanship has two interacting components: the horse as mount and the human as rider (18). Horseback riding is a demanding physical activity, and adaptive changes of the musculoskeletal apparatus as response to frequent specific biomechanical stressors are well documented (19–21). Human skeletons from archeological sources are available in higher numbers than horse skeletons and often are better preserved because of proper burials. They therefore provide a more approachable source of information about horse riding (see section S2 and fig. S2 for details). There is continuing research in this direction [(22), pp. 28–38], but diagnosing an individual as a rider by skeletal traits alone still encounters limitations. The influence of genetics, age, sex, height, weight, limb proportions, diet, or pathological conditions on the relevant skeletal traits is not fully understood; there are no experiments with regard to the thresholds of strain necessary to provoke adaptive responses. Some activities other than riding, such as barrel making or basket weaving, could result in similar biomechanical stress and, thus, similar reactions of bone tissue. Not all possible indicators are equally reliable, and there is no established evaluation system, so interpretations can vary [(22), p. 40]. No doubt, a comprehensive basic study on reliable control groups (e.g., Early Neolithic Linearbandkeramik (LBK) versus Early Medieval Eurasian nomads) with proven rider and nonrider background is lacking. However, a number of studies of historical human skeletons with known equestrian activity demonstrate the informative value of a certain set of osteological traits specifically associated with horse riding, on which we base our observations (23–25).
Here, we report five Yamnaya individuals from the sites of Strejnicu in Romania, Malomirovo and Vetrino in Bulgaria, and Dévaványa and Balmazújváros in Hungary. We also briefly discuss two 1750 to 1540 and 1611 to 1446 calibrated BCE Middle Bronze Age individuals from two Medgidia mounds in Romania and the cases of a 3331 to 2927 calBCE “pre-Yamnaya” individual of Blejoi in Romania and a 4442 to 4243 calBCE Copper Age individual of Csongrád-Kettőshalom in Hungary (individual nos. 064, 116, 118, 213, 215, 153, 161, 032, and 209 in Tables 1 and 2). These individuals display ≥4 of 6 (diagnostic threshold of >50%) skeletal traits indicative of the so-called “horsemanship syndrome” (26) with a high level of diagnostic certainty. We also list 15 more individuals, of which 9 are Yamnaya, with three positive diagnostic trait categories and, thus, lower probability (Table 1). These were among 217 mostly “steppe” individuals from 39 sites dated between fifth and second millennium BCE studied 2019 to 2022, of which ~150 are archeologically assigned to the Early Bronze Age Yamnaya culture (see Materials and Methods and Fig. 2 for details; see also section S3). Note that recognizing evidence of horseback riding was not the intention of the undertaken study on the mentioned skeletal materials; finding the traits was incidental and rather unexpected.
Table 1. List of individuals with possible “horsemanship syndrome” displaying at least three of the six diagnostic traits.
Overview of trait appearances (+, present; −, absent; ?, not preserved; numbers in brackets indicate the relative weight of diagnostic specificity).
Ind. no. | 032 | 034 | 064 | 081 | 082 | 092 | 103 | 116 | 118 | 130 | 135 | 148 |
---|---|---|---|---|---|---|---|---|---|---|---|---|
Femoral/pelvic entheses (3) | + | + | + | + | + | − | + | + | + | + | + | + |
Ovalization of acetabulum (3) | ? | ? | + | ? | ? | ? | ? | ? | + | ? | ? | − |
Femoroacetabular lesion (2) | + | + | + | ? | + | + | + | ? | + | + | + | + |
Platymeric femur (2) | + | + | + | + | + | ? | ? | + | + | + | − | − |
Specific vertebral degeneration (1) | + | − | + | + | − | + | − | + | + | ? | + | + |
Specific trauma (1) | − | − | + | − | − | + | + | + | − | − | − | − |
Number of traits | 4 | 3 | 6 | 3 | 3 | 3 | 3 | 4 | 5 | 3 | 3 | 3 |
Ind. no. | 153 | 161 | 164 | 166 | 170 | 174 | 177 | 186 | 198 | 209 | 213 | 215 |
Femoral/pelvic entheses (3) | + | + | + | + | + | + | + | + | ? | + | + | + |
Ovalization of acetabulum (3) | + | + | ? | ? | ? | ? | ? | ? | − | + | + | + |
Femoroacetabular lesion (2) | + | − | + | ? | + | + | + | + | + | + | + | − |
Platymeric femur (2) | − | + | − | + | + | + | + | + | ? | − | + | + |
Specific vertebral degeneration (1) | + | + | + | ? | ? | ? | ? | ? | + | + | + | + |
Specific trauma (1) | + | − | − | + | ? | − | − | ? | + | + | − | − |
Number of traits | 5 | 4 | 3 | 3 | 3 | 3 | 3 | 3 | 3 | 5 | 5 | 4 |
Table 2. List of individuals with possible horsemanship syndrome displaying at least three of the six diagnostic traits.
Individual data and relative diagnostic weight in the applied scoring system. Highlighted are the individuals with ≥4 of 6 traits and ≥7 of 12 points. RO, Romania; BG, Bulgaria; HU, Hungary; CZ, Czechia; m, male; f, female; B.P., Before Present.
Ind. no. | Country | Tag | Complete | Sex | Age | Date | Score |
---|---|---|---|---|---|---|---|
032 | RO | Blejoi (Prahova District, Romania) 2016, mound III, grave 3 | 95% | m | 25–35 years | Pre-Yamnaya (DeA-8814) 4437 ± 34 before the present (B.P.), 3331–2927 calBCE | 7 |
034 | RO | Blejoi (Prahova District, Romania), mound III, grave 5 | 100% | m | 25–35 years | Pre-Yamnaya (DeA-8815) 4452 ± 33 B.P., 3338–2939 calBCE | 5 |
064 | RO | Strejnicu (Prahova District, Romania) 2011, mound I, grave 3 | 95% | m | 30–40 years | Yamnaya (Hd-30719) 4106 ± 38 B.P., 2869–2501 calBCE (BRAMS-3586) 4190 ± 28 B.P., 2891–2669 calBCE | 12 |
081 | BG | Boyanovo (Yambol District, Bulgaria), “Bajlar Kajrak,” mound I, grave 13 | 85% | m | 22–26 years | Yamnaya | 5 |
082 | BG | Boyanovo (Yambol District, Bulgaria), “Bajlar Kajrak,” mound III, grave 2 | 90% | m | 35–45 years | Post-Yamnaya | 5 |
092 | BG | Boyanovo (Yambol District, Bulgaria), “Bajlar Kajrak,” mound I, grave 17 | 90% | m | 25–30 years | Pre-Yamnaya (early Yamnaya?) | 3 |
103 | CZ | Vliněves (Mělník District, Czech Republic), grave 4214A; inv. no.: P7A 41603 | 95% | m | 25–35 years | Corded Ware (CRL-9194) 4133 ± 87 B.P., 2896–2488 calBCE (MAMS-44711) 4174 ± 25 B.P., 2881–2669 calBCE | 5 |
116 | BG | Malomirovo (Yambol District, Bulgaria) 2021, grave 17 | 90% | m | 65–75 years | Yamnaya (Poz-141946) 4315 ± 35 B.P., 3018–2884 calBCE | 7 |
118 | BG | Vetrino (Varna District, Bulgaria) 2020, Necropole 1, mound XXXIV, grave 3 | 95% | m | 25–35 years | Yamnaya (SUERC-95535) 4138 ± 22 B.P., 2873–2623 calBCE | 10 |
130 | BG | Vetrino (Varna District, Bulgaria), Necropole 3, mound I, grave 9 | 80% | (f) | 35–45 years | Yamnaya (SUERC-97452) 4172 ± 27 B.P., 2883–2635 calBCE | 6 |
135 | BG | Chudomir (Razgrad District, Bulgaria), mound I, grave 9 | 85% | m | 35–50 years | Yamnaya | 4 |
148 | RO | Medgidia (Constanţa District, Romania), mound V/VI, grave 1a | 100% | m | 35–40 years | Post-Yamnaya | 3 |
153 | RO | Medgidia (Constanţa District, Romania) 2010, mound VI, grave 6 | 100% | m | 40–50 years | Middle Bronze Age (DeA-9728) 3254 ± 28 B.P., 1611–1446 calBCE | 9 |
161 | RO | Medgidia (Constanţa District, Romania) 2010, mound V, grave 4 | 85% | m | 45–60 years | Middle Bronze Age (DeA-9667) 3361 ± 32 B.P., 1750–1540 calBCE | 7 |
164 | RO | Medgidia (Constanţa District, Romania), mound V, grave 7 | 95% | (m) | 15–18 years | Yamnaya | 4 |
166 | RO | Medgidia (Constanţa District, Romania), mound III, grave 1–2, individual 1 | 70% | m | 35–45 years | Pre-Yamnaya | 6 |
170 | RO | Medgidia (Constanţa District, Romania), mound III, grave 11 | 65% | (m) | 25–35 years | Yamnaya (BRAMS-3579) 4129 ± 28 B.P., 2870–2581 calBCE | 7 |
174 | RO | Medgidia (Constanţa District, Romania), mound III, grave 16 | 75% | (f) | 17–20 years | Yamnaya (BRAMS-3582) 4106 ± 28 B.P., 2865–2505 calBCE | 6 |
177 | RO | Aliman (Constanţa District, Romania), grave 1/s | 75% | m | 40–50 years | Yamnaya (BRAMS-3575) 4096 ± 28 B.P., 2859–2500 calBCE | 6 |
186 | HU | Debrecen (Hajdú-Bihar District, Hungary), Basahalom 1906/1326/32 | 30% | (m) | 20–40 years | Yamnaya (DeA-34604) 4322 ± 32 B.P., 3018–2888 calBCE | 7 |
198 | HU | Kunhegyes (Jász-Nagykun-Szolnok District, Hungary), Nagyállás-halom grave 18 | 75% | m | 35–50 years | Yamnaya (Poz-39456) 4195 ± 35 B.P., 2895–2636 calBCE | 3 |
209 | HU | Csongrád (Csongrád-Csanád District, Hungary) 1963, Bárdos-farmstead, -Kettőshalom, grave 1 | 100% | m | 25–35 years | Copper Age (Poz-41865) 5470 ± 40 B.P., 4442–4243 calBCE | 9 |
213 | HU | Dévaványa (Békés District, Hungary) 1969, Barcé-halom, grave 1 | 90% | m | 40–50 years | Yamnaya (DeA-8221) 4279 ± 22 B.P., 2916–2881 calBCE | 10 |
215 | HU | Balmazújváros (Hajdú-Bihar District, Hungary) 1964, Árkusmajor, –Kettőshalom, grave 1 | 40% | m | 35–45 years | Yamnaya (Poz-39461) 4320 ± 35 B.P., 3021–2886 calBCE | 7 |
Fig. 2. Distribution of the 217 individuals from 39 sites that were bioanthropologically evaluated for this study.
The background map displays all excavated Yamnaya kurgans—yellow dot—in Romania (RO), Bulgaria (BG), Hungary (HU), and Serbia (Fig. 9 of 15, amended); light blue squares denote potential Yamnaya kurgans outside the main distribution. Sites, white triangles: RO 1, Ariceştii-Rahtivani; 2, Blejoi; 3, Coada Izvorului; 4, Nedelea; 5, Păuleşti; 6, Ploieşti; 7, Strejnicu; 8, Târgşoru Vechi; 9, Boldeşti-Grădiştea; 10, Aliman; 11, Medgidia. BG 12, Kamentsi; 13, Chudomir; 14, Vetrino; 15, Mogila; 16, Boyanovo; 17, Malomirovo. HU 18, Kétegyháza; 19, Dévaványa; 20, Sárrétudvari; 21, Püspökladány Kinczesdomb; 22, Debrecen; 23, Balmazújváros; 24, Mezőcsát; 25, Bojt; 26, Tiszavasvári; 27, Nagyhegyes-Elep; 28, Hajdunánás-Tedej; 29, Hajduböszörmény; 30, Földeák; 31, Berettyóújfalu; 32, Csongrád. Czechia (early Corded Ware) 33, Plotiště; 34, Neratovice; 35, Obříství; 36, Vliněves; 37, Trmice; 38, Stadice; 39, Konobrže.
The starting point of our study is the well-preserved skeleton (Fig. 1A) from Strejnicu (Prahova District, Romania), mound I, grave 3 (hence, I/3), dated to ~2879 to 2633 calBCE (see section S4 and figs. S3 to S5) and displaying a typical Yamnaya culture burial (15, 27). The individual, morphologically male, died at an age of 30 to 40 years. With ~165 cm of height, he was rather short compared to other males of the same population (regional mean of ~172 cm) but shared their robust phenotype. Osteological examination showed one of the best records of distinct adaptive, degenerative, and traumatic traits of any of the examined skeletons, thus providing valuable insights about physical activity of the deceased. Although there is no consensus on an optimal set of diagnostic traits [(22), p. 40], we recorded the following, which are used widely as indicators of more than occasional horseback riding activity:
1) Entheseal stress reactions on pelvis and femur (Fig. 3, A to C and E): The individual from Strejnicu I/3 exhibits pronounced entheseal marks (25, 28, 29), especially on the attachment sites of adductor muscles on pelvis and femur, namely, Musculus glutaeus medius and minimus, M. adductor brevis, M. adductor longus, M. adductor magnus, M. pectineus and M. iliacus. They are not completely uniform, but they generally show raised margins, dense and rugose surfaces, and only slight enthesophytes but no cortical defects or porosities. When sitting astride on a moving mount, especially without saddle and stirrups, a rider must hold on to the mount’s back and balance each step by continuous and sometimes forceful contractions of his lower body and thigh muscles. In everyday locomotive activity, these muscles usually experience a continuous but rarely intense workload.
Fig. 3. The Strejnicu I/3 individual: Adaptive changes to bone morphology.
(A) The sclerotic plaque caused by femoroacetabular contact. (B) The elevated entheses of the M. adductor magnus and the thickened lateral to superior acetabular rim. (C) The entheses of M. iliacus. (D) The entheses of M. glutaeus minimus and medius. (E) The entheses of M. pectineus, M. adductor magnus, M. adductor brevis, and M. vastus lateralis (photo credit: M. Trautmann, University of Helsinki). (F) Femoral shaft cross sections in the computed tomography (CT) (see section S5).
2) Acetabular ovalization (Fig. 3A): The pelvis of Strejnicu I/3 is only partially preserved, so measurements of the acetabulum were not available. Still, the anterosuperior margo acetabuli appears extended, thickened, and very dense, possibly a response to pressure or impact stress inflicted by frequent close contact of this structure with the collum femoris in a sitting position with updrawn legs [(22), pp. 85–87, (30, 31)]. In theory, anterosuperior ovalization of the acetabulum could also develop from normal standing and walking activity in individuals with a very high body weight.
3) Femoroacetabular alterations (Fig. 3D): Both femurs of the Strejnicu I/3 individual show distinct impression dents with a dense and raised bony margin on the anterosuperior part of the collum femoris (32, 33). This so-called antero-iliac plaque probably shares its etiology with Poirier’s facet but may be a less-reliable symptom of horse riding [(22), pp. 67 and 143). It indicates a frequent and long-lasting sitting position with spread and drawn-up legs. Pincer-like protrusions of the acetabular rim or an asymmetric shape of the collum femoris as connected with the two types of femoroacetabular impingement condition were not observed. The surface is elevated and not cribrous, which rules out Allen’s fossa, and not clearly connected to the articular surface as typical for Poirier’s facet.
4) Bone-shaft cross-sectional shape (Fig. 3F; see also section S5): Both femoral diaphyses display anteroposterior flattening (platymeric index 81.4) and a thicker medial and lateral cortical mass in the subtrochanteric diaphysis (34–36). This can be understood as an adaptation to mediolateral bending and traction stress on the proximal femur shaft, as in horse riding. Shape adaption due to mechanic demands of this kind mainly develops during adolescence. This makes it probable that the individual from Strejnicu I/3 rode regularly from a young age (37). The upper femur-shaft cross-sectional shape is potentially also influenced by relative pelvis breadth and femur angle, because female individuals usually show a tendency toward a higher platymeric index than males.
5) Stress-induced vertebral degeneration (Fig. 4, A and B): The Strejnicu I/3 individual shows signs of a medium-level spondylosis of the lower thoracal and lumbar region, with sclerosis of the anterior rim and small osteophytes. Conditions such as Diffuse Idiopathic Skeletal Hyperostosis (DISH) that may influence the assessment were not observed. The vertebrae are affected symmetrically. Their end plates are slightly concave as a symptom of vertical pressure stress on the intervertebral discs and show small herniation imprints (Schmorl’s nodes). Degenerative changes of the spine are common in premodern individuals, but the general degree of spondylopathies appears comparatively low in the Yamnaya population samples so far examined. In addition, asymmetrical alterations that indicate load with a side preference caused by tool use are rarely observed. The changes found here are more in concert with the repetitive vertical impact stress in an upright position with pronounced lordosis, as typically suffered from riding [(22), pp.115 and 159, (38, 39)].
Fig. 4. The Strejnicu I/3 individual: Degenerative and traumatic traits.
(A and B) The thickened and sclerotic frontal margin of two vertebrae and the concave deformation of the end plates. (C) The misaligned processus spinosus of the first sacral vertebra (photo credit: M. Trautmann, University of Helsinki). (D) A CT image of the same feature (lateral view), displaying the replacement of spongious tissue by compact bone (see section S5)
6) Trauma by accident (Fig. 4, C and D): The os sacrum of the Strejnicu individual shows a well-healed but slightly misaligned processus spinosus of the uppermost sacral vertebra. No structural weakening by inflammative, infectious, or neoplastic cause is visible, so a traumatic etiology is probable. A fatigue fracture can be ruled out, because the bone part is positioned too high to be affected by any type of sitting position; a forceful fall on the backside is the most likely trauma scenario. Falls from horseback are the most common cause of injury in conjunction with equestrianism, often resulting in fractured bones of the limbs or the trunk (40, 41). Other common injuries from handling horses are from bites (usually the hands are affected), from kicks (most often resulting in pelvic and thoracal injuries), or from stepping on the foot.
DISCUSSION
A scoring system for assessing horsemanship syndrome and earliest horse riding
Each single trait as described above may not be a specific “occupational marker” for horse riding. The simultaneous occurrence of all six traits in the case of the Strejnicu Yamnaya man, however, gives the interpretation of habitual horsemanship a good degree of plausibility. Apart from our threshold of more than half of relevant diagnostic traits (≥4 of 6 traits) related to biomechanical stress from frequent horseback riding, we applied an additional scoring point system to better assess the degree of symptomatic probability between individuals. This is due to the consideration that a syndrome does not necessarily need to display all correlated diagnostic symptoms, but diagnostic reliability increases with the number of positive symptoms, and some symptoms have a higher and more specific diagnostic value than others. This scoring system (Table 2; see also section S3; the table also provides a summary of the other burials) takes the occurrence of certain traits, their severity of expression, preservation, and relative importance [(22), p. 159] into account. By doing so, we set a second threshold of ≥7 points out of the possible maximum value of 12 (again, more than half).
Besides Strejnicu, particularly the near-complete skeletons of a man of Vetrino (N1, XXXIV/3; fig. S10), 25 to 35 years old at death and a near-coeval dated to 2873 to 2623 calBCE, scores nearly as high, followed by an earlier (2916 to 2881 calBCE) 40- to 50-year-old man of Dévaványa (Barcé-halom, grave 1; fig. S13). Both display pelvic and femoral entheses, acetabular ovalization, femoroacetabular lesions, platymeric femora, and specific vertebral degeneration but lack fall-related traumata. Two more Yamnaya individuals, still displaying four traits, also meet our scoring threshold of ≥7 of 12 points. Of these, the Malomirovo grave 17 (fig. S9) individual, male and 65 to 75 years old, strongly displays the distinctive characteristics and would possibly score higher, if more relevant skeletal regions would be preserved. The Malomirovo and the Balmazújváros (-Kettőshalom grave 1; fig. S11) individuals represent, with radiocarbon dates of 3018 to 2884 and 3021 to 2886 calBCE, respectively, an early Yamnaya horizon.
We also list four more graves meeting the threshold, which are however non-Yamnaya. While the two mature individuals of Medgidia V/4 and VI/6 (figs. S7 and S8) are securely dated to the mid-second millennium BCE, the 25- to 35-year-old man buried in the late fourth millennium BCE in Blejoi III/3 (fig. S6) is culturally displaying a mixture of local elements with those of either pre-Yamnaya or (chronologically possible) incoming first Yamnaya pastoralists (15, 27). Special attention is deserved by the case of the individual of Csongrád-Kettőshalom in Hungary (fig. S12). Displaying five traits, this 25 to 35 years old scores as high as our five Yamnaya individuals and thus meets our requirements to qualify as a rider with a sufficiently high probability. However, his Copper Age date in the second half of the fifth millennium BCE and his geographical isolation call for caution because we lack comparably assessed skeletons of this period and his special cultural context [(42), pp. 249–257; see also section S1 for his Copper Age/Eneolithic context).
A further nine Yamnaya individuals (Table 2), displaying three traits, however, do not meet the requirements set by our scoring system and, thus, fall short for us to plausibly call them riders. The same applies in our set for three more pre-Yamnaya individuals of the late fourth millennium BCE and a Corded Ware man from the Czech Republic (Vliněves, grave 4214A). Besides the men of Medgidia V/4 and VI/6, for which criteria are met, two extra steppe kurgan burials of the mid-second millennium BCE from Bulgaria and Romania also fall short. While we imagine that horsemanship was possibly widespread in the steppes by this time (43), our sample demonstrates that the practice may not always show up reliably in all traits, which is to be expected because of the aforementioned varying influences. This prevents us from securely quantifying horse riding in steppe societies of the third and second millennium BCE. However, having five plausible Yamnaya cases from three different countries in southeast Europe, spanning the near entire duration of Yamnaya culture in these regions, may well speak for a wider practice.
Biomechanical stress markers on human skeletons provide a viable way to further investigate the history of horseback riding and may even provide clues about riding styles and equipment. Depictions of Bronze Age riders (Fig. 5) usually show a position called “chair seat.” This style is mainly used when riding without padded saddle or stirrups to avoid discomfort to horse and rider. It is physically demanding, with the legs exerting constant pressure to cling to the mount’s back and needs continual balancing, but would not preclude activities such as combat or the handling of herd animals, as numerous historical examples demonstrate. The osteological features described here fit well with this riding style and may have been typical for the earliest period of horsemanship. With the later introduction of shaped and padded supporting saddles and stirrups (28), other riding styles such as the so called “split seat,” “dressage seat,” and “hunt seat” evolved (see section S2 and fig. S2).
Fig. 5. Pictorial evidence of horsemanship in the Bronze Age (c. 2100 to 1200 BCE).
(A to C) Mesopotamia. (D to F) Egypt. (G to I) Aegean-Cyprus. (A) Drawing of a seal impression depicting a horse rider, Ur III period (81). (B) Baked clay plaque mold depicting a rider, Old Babylonian period [The Trustees of the British Museum; shared under a Creative Commons Attribution–NonCommercial-ShareAlike 4.0 International (CC BY-NC-SA 4.0) license; except where otherwise noted, content within this article is licensed under a CC BY 4.0 license] (82). (C) Drawing of a seal imprint of Ili-pada, Middle Assyrian Empire (courtesy of F. Wiggermann, Leiden) (83). (D) Astarte on horseback: an Egyptian graffito, Nineteenth Dynasty (photo credit: S. Steiß, Berlin) (84). (E) Egyptian plaque of glazed steatite showing a horse rider trampling a fallen enemy, Nineteen Dynasty (The Metropolitan Museum of Art) (85). (F) Limestone relief with a messenger on horseback from the Horemheb tomb, Saqqara, Late Eighteenth Dynasty (Museo Civico Archeologico di Bologna) (86). (G) Clay figurine of the so-called “cavalryman” from Mycenae, early LH IIIB (courtesy of J. Kelder, Leiden) (87). (H) Horseman on an LHIIIB krater in the Allard Pierson Museum, Amsterdam (courtesy of J. Kelder, Leiden) (87). (I) Drawing of a sherd showing a horse rider from Minet el-Beida, tomb VI, LH IIIB2 (courtesy of J. Kelder, Leiden) (87).
Together, our findings provide a strong argument that horseback riding was already a common activity for some Yamnaya individuals as early as ~3000 calBCE. This supports other tentative third millennium BCE evidence of an early onset of equines as mounts (44). However, because of the lack of specialized gear and a comparably short breeding and training history, early horses were probably hard to handle. As Librado et al. (1) demonstrate, Yamnaya horses were markedly closer to the equid lineage known as DOM2, including all modern domestic horses, than were wild steppe horses from the sixth millennium BCE. When DOM2 was bred from late Yamnaya horses in the steppes during the second half of the third millennium BCE, genes for reduced anxiety/fear response were selected and retained in all later DOM2 horse breeds. Even DOM2 horses can be highly strung and excitable animals, so a still greater anxiety response in early Yamnaya horses probably made them even more likely to “bolt” from violent or loud actions. The military benefit of equestrianism may therefore have been limited; but nevertheless, rapid transport to and away from the site of raids would have been an advantage, even if combat was carried out on foot. Riding certainly was useful for patrolling wide tracts of land and controlling larger herds of livestock (45). It consequently would have contributed substantially to the overall success of pastoral Yamnaya society.
MATERIALS AND METHODS
A large contingent of skeletons from the modern European countries of Romania, Bulgaria, Hungary, and Czechia were bioanthropologically evaluated between 2019 and 2022 in the context of an ongoing multidisciplinary research project centered around Yamnaya burials and kurgans (Fig. 2). Most skeletons are from the Prahova and Constanţa Districts of Romania and the Yambol region of Bulgaria and were investigated during fieldwork in Ploiești (Romania) in 2019, in Yambol (Bulgaria) and Prague (Czechia) in 2020, in Budapest and Szeged (Hungary) as well as in Yambol and Bucharest (Romania) in 2021, and in Sofia (Bulgaria) and Yambol in 2022. Two hundred seventeen individuals from 39 sites were evaluated for the present study. Most (~150) have been assigned to the Yamnaya culture by radiocarbon dating and archeological context of burial customs (15, 16, 27, 42), but some individuals are ambiguous or are—as buried in the same burial mound, called kurgan—dated earlier or later, thus belonging to another culture group.
Established osteological macroscopic examination methods were applied to provide standard data for individual and demographic analysis (46–51). A selection of cranial and postcranial measurements following Howells and Martin for phenotype comparison and possibly kinship analyses (52, 53) were taken, as well as for biomechanical workload analyses. The grade and type of joint surface degeneration and entheseal characteristics (28, 54–57) were specifically recorded. Dental status including tooth loss, caries, calculus, periodontal diseases, enamel hypoplasia, dental wear grade and characteristics were documented to allow for dietary reconstruction (58–62). Furthermore, all observable paleopathological changes, traumata, and activity markers (63–67) were noted as health status indicators, as well as inheritable morphological variants (68, 69) for studies on kinship and population heterogeneity.
All data were recorded on an adapted recording sheet based on guidelines of the Global History of Health Project and formerly established documentation routines used by the lead author (50, 70). While burials do not usually provide much direct insights into daily life and activities, the remains of the deceased are a rich indirect source of information. Skeletons are archives of past lives with regard to appearance, kinship, health, diet, and activity (71–75). Taphonomic changes such as discolorations, postmortem defects, animal bite marks, or deformations were documented, and bone and tooth samples for paleogenetic and isotope studies were acquired. High-resolution photos, x-ray, and/or Computed Tomography (CT) scans were taken for a selection of skeletons if necessary.
The burial practice of depositing the body in a timber-covered underground chamber protected by a mound of earth (kurgan) collected from the surrounding surface helped to shield the inhumations against mechanical damage from animals or agriculture but produced increased chemical stress from organic acids originating from decomposition, decaying wood, and rainwater filtering through the cover of loose humus (70). Thus, the condition of bone material in general was mediocre. To assess it individually, the preserved percentage of skeletal remains (completeness) and the overall structural integrity (preservation) described by the terms hard, firm, fragile, and brittle was estimated. Values varied depending on factors such as local soil characteristics, specific burial customs, or depth below surface. The overall mean completeness was 57%, with most skeletons showing a firm to fragile condition. For all skeletal individuals, detailed excavation reports and archeological analyses are available (see sections S3 and S4 for the individuals in focus).
Preliminary analysis shows a nonrepresentative demographic selection: 132 individuals were determined as male or probably male, 65 as female or probably female, and 20 were not determined (Masculinity Index 2031). Sixty-one individuals of 217 (28.1%) died at subadult age. The detailed analysis of biomarkers regarding population characteristics and lifestyle indicators awaits the inclusion of further samples and relevant non-Yamnaya population data. However, some trends are already visible: Yamnaya-related individuals show a generally much more robust morphology of cranium and postcranium and a higher mean body height. Dental wear and workload-related joint degeneration correlated to age appear comparatively lower, while muscle attachment marks are more pronounced. Signs of interpersonal violence are very rare. So far, differences in everyday activity patterns and diet are probable and would support the assumption of a transition from a more agrarian pre-Yamnaya Neolithic population to a pastoralist Yamnaya population (76–80).
Acknowledgments
Radiocarbon dates were made available by C. Schuster, C. Ştefan, and A. Morintz, Vasile Pârvan Institute of Archaeology, Romania. The background map of Fig. 1 was provided by B. Olariu, Bucharest. The Corded Ware Bohemian samples were assessed as part of the Praemium Academiae Award Project of Michal Ernée. We thank Michal Ernée, Miroslav Dobeš, Peter Limburský (Institute of Archaeology, Czech Academy of Sciences, Prague) and Peter Velemínský (National Museum, Prague) for making the samples available for our research.
Funding: We acknowledge the funding by the European Research Council (ERC) under the European Union’s Horizon 2020 research and innovation program under grant agreement no. 788616-YMPACT. The CT scanning of osteological remains was supported by a grant of the Romanian Ministry of Education and Research, CNCS-UEFISCDI, project number PN-III-P1-1.2-PCCDI-2017-0686, within PCCDI, contract no. 52PCCDI. P.W. and S.A. wishes to mention a grant from the National Science Centre (Kraków, Poland) “From steppes to the Balkans. Yamnaya culture in Thrace” (no. NCN OPUS 2017/25/B/HS3/2516). T.H.’s work was supported by grant FK128013 of the Hungarian Research, Development, and Innovation Office, the Bolyai Scholarship of the Hungarian Academy of Sciences and ÚNKP-22-5 New National Excellence Program of the Ministry for Culture and Innovation from the source of the National Research, Development, and Innovation Fund (no: ÚNKP-22-5-ELTE-1124). Open Access funded by Helsinki University Library.
Author contributions: M.T. and V.H. conceived the study. M.T. developed the methodology and analyzed data. M.Pe. and M.F. performed laboratory work (CT scan). A.F., B.P.-B., S.A., N.A., P.W., M.Po., J.D., T.H., Z.B., R.B., A.I., A.M., D.M., A.-D.P., D.S., G.V., and V.H. assembled archeological material and prepared the site descriptions. M.T., A.F., B.P.-B., D.A., and V.H. wrote the manuscript and compiled supplementary sections with the input of all other coauthors. M.T., A.F., B.P.-B., and V.H. prepared the illustrations with the input of all other coauthors. D.A. and T.H. conducted critical review, commentary, and revision. V.H., P.W., and T.H. acquired funding.
Competing interests: The authors declare that they have no competing interests.
Data and materials availability: All data needed to evaluate the conclusions in the paper are present in the paper and/or the Supplementary Materials.
Supplementary Materials
This PDF file includes:
Supplementary Text
Figs. S1 to S13
References
REFERENCES AND NOTES
- 1.P. Librado, N. Khan, A. Fages, M. A. Kusliy, T. Suchan, L. Tonasso-Calvière, S. Schiavinato, D. Alioglu, A. Fromentier, A. Perdereau, J.-M. Aury, C. Gaunitz, L. Chauvey, A. Seguin-Orlando, C. Der Sarkissian, J. Southon, B. Shapiro, A. A. Tishkin, A. A. Kovalev, S. Alquraishi, A. H. Alfarhan, K. A. S. Al-Rasheid, T. Seregély, L. Klassen, R. Iversen, O. Bignon-Lau, P. Bodu, M. Olive, J.-C. Castel, M. Boudadi-Maligne, N. Alvarez, M. Germonpré, M. Moskal-del Hoyo, J. Wilczyński, S. Pospuła, A. Lasota-Kuś, K. Tunia, M. Nowak, E. Rannamäe, U. Saarma, G. Boeskorov, L. Lōugas, R. Kyselý, L. Peške, A. Bălăşescu, V. Dumitraşcu, R. Dobrescu, D. Gerber, V. Kiss, A. Szécsényi-Nagy, B. G. Mende, Z. Gallina, K. Somogyi, G. Kulcsár, E. Gál, R. Bendrey, M. E. Allentoft, G. Sîrbu, V. Dergachev, H. Shephard, N. Tomadini, S. Grouard, A. Kasparov, A. E. Basilyan, M. A. Anisimov, P. A. Nikolskiy, E. Y. Pavlova, V. Pitulko, G. Brem, B. Wallner, C. Schwall, M. Keller, K. Kitagawa, A. N. Bessudnov, A. Bessudnov, W. Taylor, J. Magail, J.-O. Gantulga, J. Bayarsaikhan, D. Erdenebaatar, K. Tabaldiev, E. Mijiddorj, B. Boldgiv, T. Tsagaan, M. Pruvost, S. Olsen, C. A. Makarewicz, S. Valenzuela Lamas, S. Albizuri Canadell, A. Nieto Espinet, M. Pilar Iborra, J. Lira Garrido, E. Rodríguez González, S. Celestino, C. Olària, J. L. Arsuaga, N. Kotova, A. Pryor, P. Crabtree, R. Zhumatayev, A. Toleubaev, N. L. Morgunova, T. Kuznetsova, D. Lordkipanize, M. Marzullo, O. Prato, G. Bagnasco Gianni, U. Tecchiati, B. Clavel, S. Lepetz, H. Davoudi, M. Mashkour, N. Y. Berezina, P. W. Stockhammer, J. Krause, W. Haak, A. Morales-Muñiz, N. Benecke, M. Hofreiter, A. Ludwig, A. S. Graphodatsky, J. Peters, K. Y. Kiryushin, T.-O. Iderkhangai, N. A. Bokovenko, S. K. Vasiliev, N. N. Seregin, K. V. Chugunov, N. A. Plasteeva, G. F. Baryshnikov, E. Petrova, M. Sablin, E. Ananyevskaya, A. Logvin, I. Shevnina, V. Logvin, S. Kalieva, V. Loman, I. Kukushkin, I. Merz, V. Merz, S. Sakenov, V. Varfolomeyev, E. Usmanova, V. Zaibert, B. Arbuckle, A. B. Belinskiy, A. Kalmykov, S. Reinhold, S. Hansen, A. I. Yudin, A. A. Vybornov, A. Epimakhov, N. S. Berezina, N. Roslyakova, P. A. Kosintsev, P. F. Kuznetsov, D. Anthony, G. J. Kroonen, K. Kristiansen, P. Wincker, A. Outram, L. Orlando, The origins and spread of domestic horses from the Western Eurasian steppes. Nature 598, 634–640 (2021). [DOI] [PMC free article] [PubMed] [Google Scholar]
- 2.S. Guimaraes, B. S. Arbuckle, J. Peters, S. E. Adcock, H. Buitenhuis, H. Chazin, N. Manaseryan, H.-P. Uerpmann, T. Grange, E.-M. Geigl, Ancient DNA shows domestic horses were introduced in the southern Caucasus and Anatolia during the Bronze Age. Sci. Adv. 6, eabb0030 (2020). [DOI] [PMC free article] [PubMed] [Google Scholar]
- 3.S. Lindner, Chariots in the Eurasian Steppe: A Bayesian approach to the emergence of horse-drawn transport in the early second millennium BC. Antiquity 94, 361–380 (2020). [Google Scholar]
- 4.D. Anthony, D. Brown, The secondary products revolution, horse-riding, and mounted warfare. J. World Prehist. 24, 131 (2011). [Google Scholar]
- 5.W. T.-T. Taylor, C. I. Barrón-Ortiz, Rethinking the evidence for early horse domestication at Botai. Sci. Rep. 11, 7440 (2021). [DOI] [PMC free article] [PubMed] [Google Scholar]
- 6.A. K. Outram, R. Bendrey, R. P. Evershed, L. Orlando, V. F. Zaibert, “Rebuttal of Taylor and Barrón-Ortiz 2021: Rethinking the evidence for early horse domestication at Botai” (2021). 10.5281/zenodo.5142604 [accessed 1 November 2022]. [DOI] [PMC free article] [PubMed]
- 7.A. K. Outram, N. A. Stear, R. Bendrey, S. Olsen, A. Kasparov, V. Zaibert, N. Thorpe, R. P. Evershed, The earliest horse harnessing and milking. Science 323, 1332–1335 (2009). [DOI] [PubMed] [Google Scholar]
- 8.A. Outram, A. Polyakov, A. Gromov, V. Moiseyev, A. W. Weber, V. I. Bazaliiskii, O. I. Goriunova, “Archaeological supplement B to Damgaard et al. 2018: Discussion of the archaeology of Central Asian and East Asian Neolithic to Bronze Age hunter-gatherers and early pastoralists, including consideration of horse domestication” (2018). 10.5281/zenodo.1240521 [accessed 19 June 2022]. [DOI]
- 9.S. Wilkin, A. Ventresca Miller, R. Fernandes, R. Spengler, W. T.-T. Taylor, D. R. Brown, D. Reich, D. J. Kennett, B. J. Culleton, L. Kunz, C. Fortes, A. Kitova, P. Kuznetsov, A. Epimakhov, V. F. Zaibert, A. K. Outram, E. Kitov, A. Khokhlov, D. Anthony, N. Boivin, Dairying enabled Early Bronze Age Yamnaya steppe expansions. Nature 598, 629–633 (2021). [DOI] [PMC free article] [PubMed] [Google Scholar]
- 10.A. Bennett, J. Weber, W. Bendhafer, S. Champlot, J. Peters, G. M. Schwartz, T. Grange, E.-M. Geigl, The genetic identity of the earliest human-made hybrid animals, the kungas of Syro-Mesopotamia. Sci. Adv. 8, eabm0218 (2022). [DOI] [PMC free article] [PubMed] [Google Scholar]
- 11.D. Bodi, The mustering of tribes for battle in 1 Samuel 11 and in Arm II 48 and the donkey as the Hebrew royal symbol in light of Amorite customs. Revue Int. d’Histoire Militaire Ancienne 5, 7–31 (2017). [Google Scholar]
- 12.R. Kyselý, L. Peške, Horse size and domestication: Early equid bones from the Czech Republic in the European context. Anthropozoologica 51, 1–39 (2016). [Google Scholar]
- 13.S. Burmeister, Early wagons in Eurasia: Disentangling an enigmatic innovation, in Appropriating Innovations Entangled Knowledge in Eurasia, 5000–1500 BCE, P. W. Stockhammer, J. Maran, Eds. (Oxbow, 2017), pp. 69–77. [Google Scholar]
- 14.W. Honeychurch, L. Rogers, C. Amartuvshin, E. Diimaajav, N.-O. Erdene-Ochir, M. E. Hall, M. Hrivnyak, The earliest herders of East Asia: Examining Afanasievo entry to Central Mongolia. Arch. Res. in Asia 26, 100264 (2021). [Google Scholar]
- 15.B. Preda-Bălănică, A. Frînculeasa, V. Heyd, The Yamnaya impact North of the Lower Danube: A tale of newcomers and locals. Bull. Soc. Préhist. française 117, 85–101 (2020). [Google Scholar]
- 16.E. Kaiser, Das dritte Jahrtausend im osteuropäischen Steppenraum (Berlin Studies of the Ancient World, Edition Topoi, 2019), vol. 37. [Google Scholar]
- 17.S. Bőkőnyi, The earliest waves of domestic horses in East Europe. J. Indo Eur. Stud. 6, 17–76 (1978). [Google Scholar]
- 18.H. M. Clayton, S.-J. Hobbs, The role of biomechanical analysis of horse and rider in equitation science. Appl. Anim. Behav. Sci. 190, 123–132 (2017). [Google Scholar]
- 19.G. Pálfi, O. Dutour, Activity-induced skeletal markers in historical anthropological material. Int. J. Anthropol. 11, 41–55 (1996). [Google Scholar]
- 20.R. Jurmain, F. Alves Cardoso, C. Henderson, S. Villotte, Bioarchaeology’s holy grail: The reconstruction of activity, in A Companion to Paleopathology, A. L. Grauer, Ed. (Wiley/Blackwell, 2012), pp. 531–542. [Google Scholar]
- 21.C. S. Larsen, Bioarchaeology: Interpreting Behavior from the Human Skeleton (Cambridge Univ. Press, ed. 2, 2015). [Google Scholar]
- 22.W. Berthon, “Bioarchaeological analysis of the mounted archers from the Hungarian Conquest period (10th century): Horse riding and activity-related skeletal changes,” thesis, University of Szeged, Hungary and École Pratique des Hautes Études, PSL University, Paris, France (2019). [Google Scholar]
- 23.E. Bagagli, F. Cantini, F. Mallegni, F. Bartoli, “Horseman Syndrome” in the Tuscan Early Middle Age: The Sk888 case. J. Biol. Res. 85, 203–204 (2012). [Google Scholar]
- 24.J. Zaia, Saddle Sore: Skeletal Occupational Markers of Habitual Horseback Riding (Senior Honors Theses 269, Brockport NY, 2019); http://hdl.handle.net/20.500.12648/6765
- 25.K. Djukic, N. Miladinovic-Radmilovic, M. Draskovic, M. Djuric, Morphological appearance of muscle attachment sites on lower limbs: Horse riders versus agricultural population. Int. J. Osteoarchaeol. 28, 656–668 (2018). [Google Scholar]
- 26.G. Pálfi, Traces des activités sur les squelettes des anciens Hongrois. Bull. et Mémoires Soc. d’Anthropologie de Paris 4, 209–231 (1992). [Google Scholar]
- 27.A. Frînculeasa, B. Preda, V. Heyd, Pit-graves, Yamnaya and Kurgans along the Lower Danube: Disentangling 4th and 3rd millennium BC burial customs, equipment and chronology. Praehist. Zeitschrift 90, 45–113 (2015). [Google Scholar]
- 28.S. Villotte, C. J. Knüsel, Understanding entheseal changes: Definition and life course changes. Int. J. Osteoarchaeol. 23, 135–146 (2013). [Google Scholar]
- 29.S. Stefanović, M. Porčić, Between-group differences in the patterning of musculo-skeletal stress markers: Avoiding confounding factors by focusing on qualitative aspects of physical activity. Int. J. Osteoarchaeol. 23, 94–105 (2013). [Google Scholar]
- 30.J. D. Erickson, D. V. Lee, J. E. A. Bertram, Fourier analysis of acetabular shape in Native American Arikara populations before and after acquisition of horses. Am. J. Physical Anthropol. 113, 473–480 (2000). [DOI] [PubMed] [Google Scholar]
- 31.W. Berthon, B. Tihanyi, L. Kis, L. Révész, H. Coqueugniot, O. Dutour, G. Pálfi, Horse riding and the shape of the acetabulum: Insights from the bioarchaeological analysis of early Hungarian mounted archers (10th century). Int. J. Osteoarchaeol. 29, 117–126 (2019). [Google Scholar]
- 32.S. Villotte, C. J. Knüsel, Some remarks about femoroactebular impingement and osseous non-metric variations of the proximal femur. Bull. et Mémoires Soc. d’Anthropologie de Paris 21, 95–98 (2009). [Google Scholar]
- 33.N. Radi, V. Mariotti, A. Riga, S. Zampetti, C. Villa, M. G. Belcastro, Variation of the anterior aspect of the femoral head-neck junction in modern human identified skeletal collection. Am. J. Phys. Anthropol. 152, 261–272 (2013). [DOI] [PubMed] [Google Scholar]
- 34.C. J. Knüsel, Bone adaptation and its relationship to physical activity in the past, in Human Osteology in Archaeology and Forensic Science, M. Cox, S. Mays, Eds. (Greenwich Medical Media Ltd. and Cambridge Univ. Press, 2000), pp. 381–402. [Google Scholar]
- 35.D. J. Wescott, “Structural variation in the humerus and femur in the American Great Plains and adjacent regions: Differences in subsistence strategy and physical terrain,” thesis, The University of Tennessee, Knoxville, TN (2001). [Google Scholar]
- 36.B. K. McIlvaine, L. A. Schepartz, Femoral subtrochanteric shape variation in Albania: Implications for use in forensic applications. Homo 66, 79–89 (2015). [DOI] [PubMed] [Google Scholar]
- 37.C. Shaw, J. Stock, Intensity, repetitiveness, and directionality of habitual adolescent mobility patterns influence the tibial diaphysis morphology of athletes. Am. J. Phys. Anthropol. 140, 149–159 (2009). [DOI] [PubMed] [Google Scholar]
- 38.C. N. Kraft, P. H. Pennekamp, U. Becker, M. Young, O. Diedrich, C. Lüring, M. von Falkenhausen, Magnetic resonance imaging findings of the lumbar spine in elite horseback riders: Correlations with back pain, body mass index, trunk/leg-length coefficient, and riding discipline. Am. J. Sports Med. 37, 2205–2213 (2009). [DOI] [PubMed] [Google Scholar]
- 39.K. D. Williams, N. J. Meinzer, C. S. Larsen, History of degenerative `joint disease in people across Europe, in The Backbone of Europe Health, Diet, Work and Violence over Two Millennia, R. H. Steckel, C. S. Larsen, C. A. Roberts, J. Baten, Eds. (Cambridge Univ. Press, 2018), pp. 253–299. [Google Scholar]
- 40.R. T. Loder, The demographics of equestrian-related injuries in the United States: Injury patterns, orthopedic specific injuries, and avenues for injury prevention. J. Trauma 65, 447–460 (2008). [DOI] [PubMed] [Google Scholar]
- 41.W. Berthon, B. Tihanyi, O. A. Váradi, H. Coqueugniot, O. Dutour, G. Pálfi, Riding for a fall: Bone fractures among mounted archers from the Hungarian Conquest period (10th century CE). Int. J. Osteoarchaeol. 31, 926–940 (2021). [Google Scholar]
- 42.D. Anthony, The Horse, the Wheel, and Language: How Bronze-Age Riders from the Eurasian Steppes Shaped the Modern World (Princeton Univ. Press, 2007). [Google Scholar]
- 43.K. Kanne, Riding, ruling, and resistance: Equestrianism and political authority in the Hungarian Bronze Age. Curr. Anthropol. 63, 289–329 (2022). [Google Scholar]
- 44.T. Molleson, J. Blondiaux, Riders’ bones from Kish, Iraq. Cambridge Archaeol. J. 4, 312–316 (1994). [Google Scholar]
- 45.A. Khazanov, Nomads and the Outside World (Cambridge Univ. Press, 1984). [Google Scholar]
- 46.D. H. Ubelaker, Human Skeletal Remains: Excavation, Analysis, Interpretation (Taraxacum, 1978). [Google Scholar]
- 47.D. R. Brothwell, Digging Up Bones, The Excavation, Treatment and Study of Human Skeletal Remains (Cornell Univ. Press, ed. 3, 1981). [Google Scholar]
- 48.J. E. Buikstra, D. H. Ubelaker, Standards for Data Collection from Human Skeletal Remains. Proceedings of a Seminar at the Field Museum of Natural History (Arkansas Archaeological Survey Research Series 44, 1994). [Google Scholar]
- 49.A. Chamberlain, Demography in Archaeology (Cambridge Manuals in Archaeology, Cambridge Univ. Press, 2006). [Google Scholar]
- 50.R. H. Steckel, C. S. Larsen, P. W. Sciulli, P. L. Walker, The Global History of Health Project. Data Collection Codebook (2006); https://www.uv.es/paleolab/Codebook-08-25-051%5B1%5D.pdf.
- 51.G. Grupe, M. Harbeck, G. McGlynn, Prähistorische Anthropologie (Springer-Verlag Berlin Heidelberg, 2015). [Google Scholar]
- 52.W. W. Howells, “Who’s who in skulls. Ethnic identification of crania from measurements” (Papers of the Peabody Museum of Archaeology and Ethnology 82, Peabody Museum, 1995). [Google Scholar]
- 53.W. W. Howells, “Cranial variation in man. A study by multivariate analysis of patterns of differences among recent human populations” (Papers of the Peabody Museum of Archeology and Ethnology 67, Peabody Museum, 1973). [Google Scholar]
- 54.E. Cunha, C. Umbelino, What can bones tell about labour and occupation: The analysis of skeletal markers of occupational stress in the Identified Skeletal Collection of the Anthropological Museum of the University of Coimbra (preliminary results). Antropol. Port. 13, 49–68 (1995). [Google Scholar]
- 55.C. S. Larsen, P. L. Walker, R. H. Steckel, P. Sciulli, H. D. Klaus, J. Blondiaux, G. Grupe, R. Jankauskas, G. Maat, G. McGlynn, A. Papathanasiou, C. Roberts, M. Teschler-Nicola, U. Wittwer-Backofen, A. Agnew, S. Assis, Z. Bereczki, B. Bertrand, T. K. Betsinger, S. Boulter, C. Bourbou, A. Boylston, M. Brickley, L. Bürli, C. Cooper, A. Coppa, J. Coughlan, A. Drozd, E. During, J. Eng, F. Engel, S. Fox, M. Furtado, G. Gerhards, K. Haebler, K. Harkins, P. Holck, M. Holst, G. Hotz, H. Justus, K. Kaminska, A. Kjellström, C. J. Knüsel, T. Kozlowski, A. Lagia, C. Lopes, S. Manolis, A. Marcsik, C. Marques, C. Moenke, I. Moutafi, C. Niel, S. A. Novak, F. Novotny, J. Peck, I. Potiekhina, B. Rega, R. Richman, F. Rijpma, J. Rose, J. Ruiz, P. Sannen, A. Soficaru, M. Spannagl, R. Storm, M. E. Subirà, D. Swales, V. Tritsaroli, E. Tyler, S. Ulrich-Bochsler, S. Vatteoni, N. Villena-Mota, R. Wiggins, L. L. Williams, History of Degenerative Joint Disease in Europe: Inferences about Lifestyle and Activity (AAPA Symposium Reconstructing Health and Disease in Europe: The Early Middle Ages through the Industrial Period, 2009). [Google Scholar]
- 56.S. Villotte, S. Assis, F. Alves Cardoso, C. Y. Henderson, V. Mariotti, M. Milella, D. Pany-Kucera, N. Speith, C. A. Wilczak, R. Jurmain, In search of consensus: Terminology for entheseal changes (EC). Int. J. Paleopathol. 13, 49–55 (2016). [DOI] [PubMed] [Google Scholar]
- 57.M. Milella, F. A. Cardoso, S. Assis, G. P. Lopreno, N. Speith, Exploring the relationship between entheseal changes and physical activity: A multivariate study. Am. J. Phys. Anthropol. 156, 215–223 (2015). [DOI] [PubMed] [Google Scholar]
- 58.A. Azzopardi, D. W. Bartlett, T. F. Watson, B. G. Smith, A literature review of the techniques to measure tooth wear and erosion. Eur. J. Prosthodont. Restor. Dent. 8, 93–97 (2000). [PubMed] [Google Scholar]
- 59.K. W. Alt, F. W. Rösing, M. Teschler-Nicola, Dental Anthropology, Fundamentals, Limits and Prospects | Softcover reprint of the original 1st ed. 1998 (Springer, 2011). [Google Scholar]
- 60.C. Ganss, J. Klimek, N. Borkowski, Characteristics of tooth wear in relation to different nutritional patterns including contemporary and medieval subjects. Eur. J. Oral Sci. 110, 54–60 (2002). [DOI] [PubMed] [Google Scholar]
- 61.H. Schutkowski, Thoughts for food: Evidence and meaning of past dietary habits, in Between Biology and Culture (Cambridge Studies in Biological and Evolutionary Anthropology 56, 2008), pp. 141–164. [Google Scholar]
- 62.B. H. Smith, Patterns of molar wear in hunter-gatherers and agriculturalists. Am. J. Phys. Anthropol. 63, 39–56 (1984). [DOI] [PubMed] [Google Scholar]
- 63.A. C. Aufderheide, C. Rodríguez-Martín, The Cambridge Encyclopedia of Human Paleopathology (Cambridge Univ. Press, 1998). [Google Scholar]
- 64.A. L. Grauer, Macroscopic analysis and data collection in palaeopathology, in Advances in Human Palaeopathology, R. Pinhasi, S. Mays, Eds. (Wiley, 2008), pp. 57–76. [Google Scholar]
- 65.M. N. Cohen, G. J. Armelagos, Paleopathology at the Origins of Agriculture: Proceedings of the Conference on Paleopathology and Socioeconomic Change at the Origins of Agriculture at Plattsburgh, held April 25–May 1, 1982 (Academic Press, 1984). [Google Scholar]
- 66.D. J. Ortner, Identification of Pathological Conditions in Human Skeletal Remains (Elsevier, 2003). [Google Scholar]
- 67.C. A. Roberts, Adaptation of populations to changing environments: Bioarchaeological perspectives on health for the past, present and future. Bull. Mém. Soc. Anthropol. Paris 22, 38–46 (2010). [Google Scholar]
- 68.K. Alt, Odontologische Verwandtschaftsanalyse: Individuelle Charakteristika der Zähne in ihrer Bedeutung für Anthropologie, Archäologie und Rechtsmedizin (Fischer Verlag GmbH & Co. KG, 1998). [Google Scholar]
- 69.R. W. Mann, D. R. Hunt, S. Lozanoff, Photographic Regional Atlas of Non-Metric Traits and Anatomical Variants in the Human Skeleton (Charles C Thomas, 2016). [Google Scholar]
- 70.M. Trautmann, Die schnurkeramischen Bestattungen von Lauda-Königshofen. Steinzeitliche Hirtennomaden im Taubertal? Fundberichte aus Baden-Württemberg 32, 265–476 (2012). [Google Scholar]
- 71.J. E. Buikstra, L. A. Beck, Bioarchaeology: The Contextual Analysis of Human Remains (Academic Press, 2006). [Google Scholar]
- 72.D. J. Wescott, Effects of mobility on femur midshaft external shape and robusticity. Am. J. Phys. Anthropol. 130, 201–213 (2006). [DOI] [PubMed] [Google Scholar]
- 73.C. B. Ruff, Biomechanical analysis of archaeological human skeletons, in Biological Anthropology of the Human Skeleton, M. A. Katzenberg, S. R. Saunders, Eds. (Wiley-Liss, 2000), pp. 71–102. [Google Scholar]
- 74.C. S. Larsen, Biological changes in human populations with agriculture. Ann. Rev. Anthropol. 24, 185–213 (1995). [Google Scholar]
- 75.M. Wagner, X. Wu, P. Tarasov, A. Aisha, C. Bronk Ramsey, M. Schultz, T. Schmidt-Schultz, J. Gresky, Radiocarbon-dated archaeological record of early first millennium B.C. mounted pastoralists in the Kunlun Mountains, China. Proc. Natl. Acad. Sci. U.S.A. 108, 15733–15738 (2011). [DOI] [PMC free article] [PubMed] [Google Scholar]
- 76.M. Levine, C. Renfrew, K. Boyle, Prehistoric Steppe Adaptation and the Horse (McDonald Institute for Archaeological Research, 2003). [Google Scholar]
- 77.C. Renfrew, Pastoralism and interaction: Some introductory questions, in Ancient Interactions: East and West in Eurasia, K. Boyle, C. Renfrew, M. Levine, Eds. (McDonald Institute for Archaeological Research, 2002), pp. 1–12. [Google Scholar]
- 78.E. Rosenstock, M. Groß, A. Hujić, A. Scheibner, Back to good shape: Biological standard of living in the Copper and Bronze Ages and the possible role of food, in Human Development in Landscapes. Vol. 6. The Third Food Revolution? Setting the Bronze Age table: Common Trends in Economic and Subsistence Strategies in Bronze Age Europe. Proceedings of the International Workshop “SocioArchaeol Anthropol Sci Environmental Dynamics over the Last 12,000 Years: The Creation of Landscapes III (15th–18th April 2013)” in Kiel, J. Kneisel, M. Dal Corso, W. Kirleis, H. Scholz, N. Taylor, V. Tiedtke, Eds. (Habelt, 2015), pp. 121–152. [Google Scholar]
- 79.R. Steckel, C. S. Larsen, C. A. Roberts, J. Baten, The Backbone of Europe: Health, Diet, Work and Violence over Two Millennia (Cambridge Univ. Press, 2018). [Google Scholar]
- 80.M. Niskanen, C. B. Ruff, B. Holt, V. Sládek, M. Berner, Temporal and geographic variation in body size and shape of europeans from the Late Pleistocene to recent times, in Skeletal Variation and Adaptation in Europeans: Upper Paleolithic to the Twentieth Century, C. B. Ruff, Ed. (Wiley Blackwell, 2018), pp. 49–89. [Google Scholar]
- 81.D. I. Owen, The first equestrian: An Ur III glyptic scene. Acta Sumerologica 13, 259–273 (1991). [Google Scholar]
- 82.Collection of the British Museum, Inv. No. 22958; www.britishmuseum.org/collection/object/W_1897-0511-104 [accessed 5 April 2021].
- 83.F. A. M. Wiggermann, The seal of Ili-Padda, grand vizier of the Middle Assyrian Empire, in The Iconography of Cylinder Seals, P. Taylor, Ed. (University of London Press, 2006), pp. 92–99. [Google Scholar]
- 84.Collection of the Ägyptisches Museum und Papyrussammlung Berlin, ÄM 21826 (Photo credit: S. Steiß, Berlin). [Google Scholar]
- 85.Collection of the Metropolitan Museum New York, Acc. No. 05.3.263; www.metmuseum.org/art/collection/search/548708 [accessed 30 November 2022].
- 86.Collection of the Museo Civico Archeologico di Bologna, inv MCABo 1899; www.museibologna.it/archeologicoen/percorsi/66288/id/74955/oggetto/74958/ [accessed 9 November 2022].
- 87.J. Kelder, Horseback riding and cavalry in Mycenaean Greece. Anc. West East 11, 11–18 (2012). [Google Scholar]
- 88.J. Dani, Milleker’s pride and joy, in Danubian Route of the Yamnaya Culture: The Barrows of Vojvodina, P. Jarosz, J. Koledin, P. Włodarczak, Eds. (The Yamnaya Impact of Prehistoric Europe 3, Archaeolingua, 2021), pp. 195–206. [Google Scholar]
- 89.D. Makowicz-Poliszot, Analysis of animal bones from “Ciganska humka” in ŠajkaŠ, in Danubian Route of the Yamnaya Culture: The Barrows of Vojvodina, P. Jarosz, J. Koledin, P. Włodarczak, Eds. (The Yamnaya Impact of Prehistoric Europe 3, Archaeolingua, 2021), pp. 91–102. [Google Scholar]
- 90.S. Bökönyi, Copper Age vertebrae fauna from Kétegyháza, in The People of the Pit-grave Kurgans in Eastern Hungary, I. Ecsedy, Eds. (Fontes Arch. Hungariae, 1979), pp. 101–116. [Google Scholar]
- 91.C. Gerling, E. Bánffy, J. Dani, K. Köhler, G. Kulcsár, A. W. G. Pike, V. Szeverényi, V. Heyd, Immigration and transhumance in the Early Bronze Age Carpathian Basin: The occupants of akurgan. Antiquity 86, 1097–1111 (2012). [Google Scholar]
- 92.S. Alexandrov, Fourth/third millennium BC barrow graves in North-East Bulgaria (120 years of investigations), in Yamnaya Interactions. Proceedings of the International Workshop held in Helsinki, 25–26 April 2019, V. Heyd, G. Kulcsár, B. Preda-Bălănică, Eds. (The Yamnaya Impact of Prehistoric Europe 2, Archaeolingua, 2021), pp. 271–314. [Google Scholar]
- 93.N. Benecke, Diachroner Vergleich der Pferdehaltung im Karpatenbecken und in der osteuropäischen Steppe während der vorchristlichen Metallzeiten, in Das Karpatenbecken und die osteuropäische Steppe, B. Hänsel, J. Machnik, Eds. (Südosteuropa-Schriften 20, Prähistorische Archäologie Südosteuropa 12, Leidorf, 1998), pp. 91–98. [Google Scholar]
- 94.E. Gál, Animal bone remains from the Late Copper Age cemetery at Pilismarót-Basaharc, in The Late Copper Age cemetery at Pilismarót-Basaharc, M. Bondár, Ed. (Archaeolingua, 2015), pp. 367–379. [Google Scholar]
- 95.E. Gál, Animals at the Dawn of Metallurgy in South-Western Hungary (Archaeolingua, 2017). [Google Scholar]
- 96.N. Benecke, On the beginning of horse husbandry in the Southern Balkan peninsula—The horses from Kanlıgeçit, Kırklareli (Turkish Thrace). Turkish Acad. Sci. J. Archaeol. 12, 13–23 (2009). [Google Scholar]
- 97.L. György, Late Copper Age Animal Burials in the Carpathian Basin, in Moments in Time. Papers Presented to Pál Raczky on His 60th Birthday, A. Anders, G. Kulcsár, Eds. (Prehistoric Society, Eötvös Loránd Univ., L’Harmattan, 2013), pp. 627–642. [Google Scholar]
- 98.N. Spassov, L. Hristova, N. Iliev, The domesticated horses from the submerged prehistoric village of Urdoviza (Kiten) on the Bulgarian Black Sea coast—Among the oldest known. Hist. Nat. Bulga. 25, 11–14 (2018). [Google Scholar]
- 99.K. Lyublyanovics, Animal bones from the Bell Beaker Settlement of Albertfalva, Budapest, in A Bell Beaker Settlement in Albertfalva, Hungary (2470–1950 BC), A. Endrödi, L. Reményi, Eds. (Budapest History Museum, 2016), pp. 204–216. [Google Scholar]
- 100.C. Gaunitz, A. Fages, K. Hanghøj, A. Albrechtsen, N. Khan, M. Schubert, A. Seguin-Orlando, I. J. Owens, S. Felkel, O. Bignon-Lau, P. de Barros Damgaard, A. Mittnik, A. F. Mohaseb, H. Davoudi, S. Alquraishi, A. H. Alfarhan, K. A. S. Al-Rasheid, E. Crubézy, N. Benecke, S. Olsen, D. Brown, D. Anthony, V. Ken Massy, A. Pitulko, G. Kasparov, M. Brem, G. Hofreiter, N. Mukhtarova, L. Baimukhanov, V. Lõugas, P. W. Onar, J. Stockhammer, B. Krause, S. Boldgiv, D. Undrakhbold, S. Erdenebaatar, M. Lepetz, A. Mashkour, B. Ludwig, V. Wallner, I. Merz, V. Merz, E. Zaibert, P. Willerslev, A. K. Librado, L. O. Outram, Ancient genomes revisit the ancestry of domestic and Przewalski’s horses. Science 360, 111–114 (2018). [DOI] [PubMed] [Google Scholar]
- 101.D. Y. Telegin, A. L. Nechitailo, I. D. Potekhina, Y. V. Panchenko, Srednestogovskaya i Novodanilovskaya Kul’tury Eneolita Azovo-Chernomorskogo Regiona (Lugansk, 2001). [Google Scholar]
- 102.S. Reinhold, J. Gresky, N. Berezina, A. R. Kantorovich, C. Knipper, V. E. Maslov, V. G. Petrenko, K. W. Alt, A. B. Belinsky, Contextualising innovation. Cattle owners and wagon drivers in the North Caucasus and beyond, in Appropriating Innovations: Entangled Knowledge in Eurasia, 5000–1500 BCE, P. Stockhammer, J. Maran, Eds. (Oxbow, 2017), pp. 78–97. [Google Scholar]
- 103.P. F. Kuznetsov, A. N. Usachuk, Obshchee i osobennoe v izgotovlenii psaliev iz Ureni—Detalei upryadzhi boevykh kolesnits. Strat. Plus 2, 335–341 (2019). [Google Scholar]
- 104.D. W. Anthony, A. A. Khokhlov, S. A. Agapov, D. S. Agapov, R. Schulting, I. Olalde, D. Reich, The Eneolithic cemetery at Khvalynsk on the Volga River. Praehistorische Zeitschrift 97, 22–67 (2022). [Google Scholar]
- 105.E. Rosenstock, A. Masson, B. Zich, Moraines, megaliths and moo: Putting the prehistoric tractor to work, in Megaliths – Societies – Landscapes: Early Monumentality and Social Differentiation in Neolithic Europe, Volume 3, Proceedings of the International Conference »Megaliths – Societies – Landscapes. Early Monumentalityand Social Differentiation in Neolithic Europe« (16th–20th June 2015) in Kiel, J. Müller, M. Hinz, M. Wunderlich, Eds. (Dr. Rudolf Habelt GmbH, 2019) pp. 1099–1110. [Google Scholar]
- 106.A. Sherratt, The secondary exploitation of animals in the Old World. World Archaeol. 15, 90–104 (1983). [Google Scholar]
- 107.N. L. Morgunova, Pri-Uralskaya Gruppa Pamyatnikov v Sisteme Volzhsko-Ural’skogo Varianta Yamnoi Kul’turno-Istoricheskoi Oblasti (OGPU, 2014). [Google Scholar]
- 108.N. I. Shishlina, Reconstruction of the Bronze Age of the Caspian Steppes: Life Styles and Life Ways of Pastoral Nomads (British Archaeological Reports International Series 1876, 2008). [Google Scholar]
- 109.R. J. Schulting, M. P. Richards, Stable isotope analysis of neolithic to Late Bronze Age populations in the Samara Valley, in A Bronze Age Landscape in the Russian Steppes: The Samara Valley Project, D. W. Anthony, D. Brown, O. Mochalov, A. Khokhlov, P. Kuznetsov, Eds. (Monumenta Archaeologica 37, Cotsen Institute of Archaeology Press, 2016), pp. 127–147. [Google Scholar]
- 110.N. I. Shishlina, E. S. Azarov, T. D. Dyatlova, N. V. Roslyakova, O. P. Bachura, J. van der Plicht, P. I. Kalinin, I. A. Idrisov, A. V. Borisov, Innovatsioniye sezonnye i sistema zhineobespecheniya podvizhn’kh skotovodov v pustynno-stepnoi zone Evrazi: pol’ sotsial’nykh grupp. Strat. Plus 2, 69–90 (2018). [Google Scholar]
- 111.C. Knipper, S. Reinhold, J. Gresky, N. Berezina, C. Gerling, S. L. Pichler, A. P. Buzhilova, A. R. Kantorovich, V. E. Maslov, V. G. Petrenko, S. V. Lyakhov, A. A. Kalmykov, A. B. Belinskiy, S. Hansen, K. W. Alt, Diet and subsistence in Bronze Age pastoral communities from the southern Russian steppes and the North Caucasus. PLOS ONE 15, e0239861 (2020). [DOI] [PMC free article] [PubMed] [Google Scholar]
- 112.A. Scott, S. Reinhold, T. Hermes, A. A. Kalmykov, A. Belinskiy, A. Buzhilova, N. Berezina, A. R. Kantorovich, V. E. Maslov, F. Guliyev, B. Lyonnet, P. Gasimov, B. Jalilov, J. Eminli, E. Iskandarov, E. Hammer, S. E. Nugent, R. Hagan, K. Majander, P. Onkamo, K. Nordqvist, N. Shishlina, E. Kaverzneva, A. I. Korolev, A. A. Khokhlov, R. V. Smolyaninov, S. V. Sharapova, R. Krause, M. Karapetian, E. Stolarczyk, J. Krause, S. Hansen, W. Haak, C. Warinner, Emergence and intensification of dairying in the Caucasus and Eurasian steppes. Nat. Ecol. Evol. 6, 813–822 (2022). [DOI] [PMC free article] [PubMed] [Google Scholar]
- 113.S. V. Ivanova, O konseptsii vostochnogo poroisxhozhdeniya Yamnao kul’turno-istoricheskoi obshchnosti. Voprosy Arkheologii Povol’zhya (Samara) 4, 203–208 (2006). [Google Scholar]
- 114.N. Y. Merpert, Drevneishie skotovody Volzhsko-Ural’skogo Mezhdurech’ya (Nauka, 1974). [Google Scholar]
- 115.V. P. Shilov, Modeli skotovodcheskikh khozyaistv stepnykh oblastei Evrazii v epokhu eneolita i rannego bronzovogo veka. Sovietskaya Arkheologiya 1, 5–15 (1975). [Google Scholar]
- 116.D. W. Anthony, The Samara Valley Project and the evolution of pastoral economies in the western Eurasian steppes, in A Bronze Age Landscape in the Russian Steppes: The Samara Valley Project, D. W. Anthony, D. Brown, P. Kuznetsov, O. Mochalov, A. Khokhlov, Eds. (Monumenta Archaeologica 37, Cotsen Institute of Archaeology Press, 2016), pp. 3–36. [Google Scholar]
- 117.A. Bogaard, M. Fochesato, S. Bowles, The farming-inequality nexus: New insights from ancient Western Eurasia. Antiquity 93, 1129–1143 (2019). [Google Scholar]
- 118.K. Kristiansen, Archaeology and the Genetic Revolution in European Prehistory (Elements in the Archaeology of Europe) (Cambridge Univ. Press, 2022). [Google Scholar]
- 119.H. M. Frost, Wolff’s Law and bone’s structural adaptations to mechanical usage: An overview for clinicians. Angle Orthod. 64, 175–188 (1994). [DOI] [PubMed] [Google Scholar]
- 120.M. Benjamin, T. Kumai, S. Milz, B. M. Boszczyk, A. A. Boszczyk, J. R. Ralphs, The skeletal attachment of tendons—Tendon ‘entheses’. Comp. Biochem. Physiol. A Mol. Integr. Physiol. 133, 931–945 (2002). [DOI] [PubMed] [Google Scholar]
- 121.K. Djukic, P. Milovanovic, M. Hahn, B. Busse, M. Amling, M. Djuric, Bone microarchitecture at muscle attachment sites: The relationship between macroscopic scores of entheses and their cortical and trabecular microstructural design. Am. J. Phys. Anthropol. 157, 81–93 (2015). [DOI] [PubMed] [Google Scholar]
- 122.L. Capasso, K. A. R. Kennedy, C. A. Wilczak, Atlas of Occupational Markers on Human Remains (Edigrafital, 1999). [Google Scholar]
- 123.F. Alves Cardoso, C. Y. Henderson, The categorisation of occupation in identified skeletal collections: A source of bias? Int. J. Osteoarchaeol. 23, 186–196 (2013). [Google Scholar]
- 124.K. A. R. Kennedy, Skeletal markers of occupational stress, in Reconstruction of Life from the Skeleton, M. Y. İşcan, K. A. R. Kennedy, Eds. (Alan R. Liss Inc., 1989), pp. 129–160. [Google Scholar]
- 125.C. Y. Henderson, F. Alves Cardoso, Special issue entheseal changes and occupation: Technical and theoretical advances and their applications. Int. J. Osteoarchaeol. 23, 127–134 (2013). [Google Scholar]
- 126.C. Y. Henderson, “Musculo-skeletal stress markers in bioarchaeology: Indicators of activity levels or human variation? A re-analysis and interpretation,” thesis, University of Durham, Durham (2009). [Google Scholar]
- 127.S. M. Aguayo, “Variations in skeletal markers and pathologies between Southern Plains equestrian and Puebloan Native American populations,” thesis, Texas Tech University, Lubbock, TX (2012). [Google Scholar]
- 128.M. R. Fuka, Activity markers and horse riding in Mongolia: Entheseal changes among Bronze and Iron Age human skeletal remains (Master thesis) (Purdue University, West Lafayette, 2018). [Google Scholar]
- 129.A. Khudaverdyan, H. Khachatryan, L. Eganyan, The human skeleton from the Late Iron Age burial of Shirakavan (Armenia): A case study. Bull. Int. Assoc. Paleodontol. 11, 51–61 (2017). [Google Scholar]
- 130.Š. Anđelinović, I. Anterić, E. Škorić, Ž. Bašić, Skeleton changes induced by horse riding on medieval skeletal remains from Croatia. Int. J. Hist. Sport 32, 708–721 (2015). [Google Scholar]
- 131.R. K. Wentz, N. T. de Grummond, Life on horseback: Palaeopathology of two Scythian skeletons from Alexandropol, Ukraine. Int. J. Osteoarchaeol. 19, 107–115 (2009). [Google Scholar]
- 132.K. Zejdlik, Z. Nyárádi, A. Gonciar, Evidence of horsemanship in two Szekler noblemen from the Baroque period. Int. J. Osteoarchaeol. 31, 66–76 (2020). [Google Scholar]
- 133.C. Baillif-Ducros, G. McGlynn, M. C. Truc, Cavaliers du passé : Activité et marqueurs ostéologiques. Proposition d’une révision du "Syndrome du cavalier“: de L’Europe à l’Asie. 1839èmes journées de la Société d’Anthropologie de Paris (2014).
- 134.C. Baillif-Ducros, Christèle, La pratique de la monte à cheval au haut Moyen Âge (fin Ve -VIIe siècle) dans le nord-est de la Gaule. État des connaissances archéologiques, recherche méthodologique sur le "syndrome du cavalier" et application d’un nouveau protocole d’étude aux populations mérovingiennes (Université de Caen Normandie, 2018). [Google Scholar]
- 135.B. Bühler, S. Kirchengast, A life on horseback? Prevalence and correlation of metric and non-metric traits of the “horse-riding syndrome” in an Avar population (7th-8th century AD) in Eastern Austria. Anthropol. Rev. 85, 67–82 (2022). [Google Scholar]
- 136.C. Baillif-Ducros, G. McGlynn, Stirrups and archaeological populations: Bio-anthropological considerations for determining their use based on the skeletons of two Steppe riders. Bull. Soc. Suisse d’Anthropologie 19, 43–44 (2013). [Google Scholar]
- 137.G. Tommasini, “Maneggi and Jumps”. The basic exercises of Renaissance horsemanship (2014); http://worksofchivalry.com/maneggi-and-jumps-the-basic-exercises-of-renaissance-horsemanship-part-1/ [accessed 19 April 2021].
- 138.G. Tommasini, “A la brida” and “a la gineta”. Different riding techniques in the late Middle Ages and the Renaissance (2014); http://worksofchivalry.com/a-la-brida-and-a-la-gineta-different-riding-techniques-in-the-late-middle-ages-%e2%80%a8and-the-renaissance/ [accessed 19 April 2021].
- 139.C. Baillif-Ducros, The Merovingian rider and his horse: Impact of equestrian equipment on the rider’s posture and skeleton. EAA 2021, Session #509 Horseman-horse couple through time and space (Inrap, 2021). [Google Scholar]
- 140.T. A. Tichnell, “Invisible horsewomen: Horse riding and social dynamics on the steppe,” thesis, Michigan State University, East Lansing, MI (2012). [Google Scholar]
- 141.L. Klejn, The steppe hypothesis of Indo-European origins remains to be proven. Acta Archaeol. 88, 193–204 (2017). [Google Scholar]
- 142.V. M. Narasimhan, N. Patterson, P. Moorjani, N. Rohland, R. Bernardos, S. Mallick, I. Lazaridis, N. Nakatsuka, I. Olalde, M. Lipson, A. M. Kim, L. M. Olivieri, A. Coppa, M. Vidale, J. Mallory, V. Moiseyev, E. Kitov, J. Monge, N. Adamski, N. Alex, N. Broomandkhoshbacht, F. Candilio, K. Callan, O. Cheronet, B. J. Culleton, M. Ferry, D. Fernandes, S. Freilich, B. Gamarra, D. Gaudio, M. Hajdinjak, É. Harney, T. K. Harper, D. Keating, A. M. Lawson, M. Mah, K. Mandl, M. Michel, M. Novak, J. Oppenheimer, N. Rai, K. Sirak, V. Slon, K. Stewardson, F. Zalzala, Z. Zhang, G. Akhatov, A. N. Bagashev, A. Bagnera, B. Baitanayev, J. Bendezu-Sarmiento, A. A. Bissembaev, G. L. Bonora, T. T. Chargynov, T. Chikisheva, P. K. Dashkovskiy, A. Derevianko, M. Dobeš, K. Douka, N. Dubova, M. N. Duisengali, D. Enshin, A. Epimakhov, A. V. Fribus, D. Fuller, A. Goryachev, A. Gromov, S. P. Grushin, B. Hanks, M. Judd, E. Kazizov, A. Khokhlov, A. P. Krygin, E. Kupriyanova, P. Kuznetsov, D. Luiselli, F. Maksudov, A. M. Mamedov, T. B. Mamirov, C. Meiklejohn, D. C. Merrett, R. Micheli, O. Mochalov, S. Mustafokulov, A. Nayak, D. Pettener, R. Potts, D. Razhev, M. Rykun, S. Sarno, T. M. Savenkova, K. Sikhymbaeva, S. M. Slepchenko, O. A. Soltobaev, N. Stepanova, S. Svyatko, K. Tabaldiev, M. Teschler-Nicola, A. A. Tishkin, V. V. Tkachev, S. Vasilyev, P. Velemínský, D. Voyakin, A. Yermolayeva, M. Zahir, V. S. Zubkov, A. Zubova, V. S. Shinde, C. Lalueza-Fox, M. Meyer, D. Anthony, N. Boivin, K. Thangaraj, D. J. Kennett, M. Frachetti, R. Pinhasi, D. Reich, The formation of human populations in South and Central Asia. Science 365, eaat7487 (2019). [DOI] [PMC free article] [PubMed] [Google Scholar]
- 143.A. V. Poliakov, S. Svyatko, N. F. Stepanova, A review of the radiocarbon dates for the Afanasyevo culture (Central Asia): Shifting towards the “shorter” chronology. Radiocarbon 61, 243–263 (2019). [Google Scholar]
- 144.T. R. Hermes, A. A. Tishkin, P. A. Kosintsev, N. F. Stepanova, B. Krause-Kyora, C. A. Makarewicz, Mitochondrial DNA of domesticated sheep confirms pastoralist component of Afanasievo subsistence economy in the Altai Mountains (3300-2900 cal BC). Archaeol. Res Asia 24, 100232 (2020). [Google Scholar]
- 145.E. B. Vadetskaia, A. V. Poliakov, N. F. Stepanova, Svod pamiatnikov afanas’evskoi kul’tury (Azbuka, 2014). [Google Scholar]
- 146.V. Heyd, Yamnaya groups and tumuli west of the Black Sea, in Ancestral Landscapes in Ancestral Landscape. Burial mounds in the Copper and Bronze Ages. Proceedings of the International Conference held in Udine, May 15th–18th 2008, E. Borgna, S. Müller Celka, Eds. (Travaux de la Maison de l’Orient et de la Méditerranée, Série recherches archéologiques 58, 2011), pp. 535–555. [Google Scholar]
- 147.I. Mathieson, S. Alpaslan-Roodenberg, C. Posth, A. Szécsényi-Nagy, N. Rohland, S. Mallick, I. Olalde, N. Broomandkhoshbacht, F. Candilio, O. Cheronet, D. Fernandes, M. Ferry, B. Gamarra, G. González Fortes, W. Haak, E. Harney, E. Jones, D. Keating, B. Krause-Kyora, I. Kucukkalipci, M. Michel, A. Mittnik, K. Nägele, M. Novak, J. Oppenheimer, N. Patterson, S. Pfrengle, K. Sirak, K. Stewardson, S. Vai, S. Alexandrov, K. W. Alt, R. Andreescu, D. Antonović, A. Ash, N. Atanassova, K. Bacvarov, M. Balázs Gusztáv, H. Bocherens, M. Bolus, A. Boroneant, Y. Boyadzhiev, A. Budnik, J. Burmaz, S. Chohadzhiev, N. J. Conard, R. Cottiaux, M. Čuka, C. Cupillard, D. G. Drucker, N. Elanski, M. Francken, B. Galabova, G. Gantesovski, B. Gély, T. Hajdu, V. Handzhyiska, K. Harvati, T. Higham, S. Iliev, I. Jankovič, I. Karavanič, D. J. Kennet, D. Kamso, A. Kozak, D. Labuda, M. Lari, C. Lazăr, M. Leppek, K. Leshtakov, D. Lo Vetro, D. Los, I. Lozanov, M. Malina, F. Martini, K. Mcsweeney, H. Meller, M. Mandušič, P. Mirea, V. Moiseyev, V. Petrova, T. D. Price, A. Simalcsik, L. Sineo, M. Šlaus, V. Slavchev, P. Stanev, A. Starovič, T. Szeniczey, S. Talamo, M. Teschler-Nicola, C. Thevenet, I. Valchev, F. Valentin, S. Vasilyev, F. Veljanovska, S. Venelinova, E. Veselovskaua, B. Viola, C. Virag, J. Zaninovič, S. Zäuner, P. W. Stockhammer, G. Catalano, R. Krauß, D. Caramelli, G. Zarina, B. Gaydarska, M. Lillie, A. G. Nikitin, I. Potekhina, A. Papathanasiou, D. Borič, C. Bonsall, J. Krause, R. Pinhasi, D. Reich, The genomic history of southeastern Europe. Nature 555, 197–203 (2018). [DOI] [PMC free article] [PubMed] [Google Scholar]
- 148.J. Dani, G. Kulcsár, Yamnaya interactions in the Carpathian Basin, in Yamnaya Interactions. Proceedings of the International Workshop held in Helsinki, 25–26 April 2019, V. Heyd, G. Kulcsár, B. Preda-Bălănică, Eds. (The Yamnaya Impact of Prehistoric Europe 2, Archaeolingua, 2021), pp. 329–359. [Google Scholar]
- 149.E. Kaiser, K. Winger, Pit graves in Bulgaria and the Yamnaya Culture. Präehistorische Zeitschrift 90, 1–27 (2015). [Google Scholar]
- 150.S. Alexandrov, Bronze Age barrow graves in Upper Thrace—Old and new questions, in Repräsentationen der Macht, Beiträge des Festkolloquiums zu Ehren des 65. Geburtstags von Blagoje Govedarica, S. Hansen, Ed. (Harrasowitz Verlag Wiesbaden, 2020), pp. 147–170. [Google Scholar]
- 151.A. Frînculeasa, The Children of the Steppe: Descendance as a key to Yamnaya success. Studii de Preistorie 16, 129–168 (2019). [Google Scholar]
- 152.J. Koledin, U. Bugaj, P. Jarosz, M. Novak, M. M. Przybyła, M. Podsiadło, A. Szczepanek, M. Spasić, P. Włodarczak, First archaeological investigations of barrows in the Bačka region and the question of the Eneolithic/Early Bronze Age barrows in Vojvodina. Präehistorische Zeitschrift 95, 350–375 (2020). [Google Scholar]
- 153.A. Frînculeasa, M. Frînculeasa, I. Dumitru, C. Buterez, The dynamics of prehistoric burial mounds of Ploieşti metropolitan area (Romania) as reflected by cartographic documents of the 18th-20th centuries. Area 49, 533–544 (2017). [Google Scholar]
- 154.A. Frînculeasa, Burial mounds in the Lower Danube region. From the international to the local and the other way round, in Yamnaya Interactions. Proceedings of the International Workshop held in Helsinki, 25–26 April 2019, V. Heyd, G. Kulcsár, B. Preda-Bălănică, Eds. (The Yamnaya Impact of Prehistoric Europe 2, Archaeolingua, 2021), pp. 173–205. [Google Scholar]
- 155.A. Frînculeasa, Earthen burial mounds and the Coţofeni culture south of the Carpathians. Archaeological research in Ariceştii-Rahtivani-Movila pe Răzoare. Ziridava. Studia Archaeol. 34, 35–90 (2020). [Google Scholar]
- 156.A. Frînculeasa, B. Preda, O. Negrea, A. Soficaru, V. Dumitraşcu, M. Frînculeasa, Complexe funerare de la începutul mileniului al II-lea descoperite recent în judeţul Prahova. Mater. şi Cercet. Arheol. 8, 139–163 (2012). [Google Scholar]
- 157.C. Bronk Ramsey, S. Lee, Recent and planned developments of the program OxCal. Radiocarbon 55, 720–730 (2013). [Google Scholar]
- 158.A. Frînculeasa, B. Preda, C. Dumitrescu, O. Negrea, A.-D. Soficaru, Blejoi, jud. Prahova, in Cronica Cercetărilor Arheologice din România, Campania 2016, A LI-a Sesiune Naţională de Rapoarte Arheologice, Muzeul Naţional de Istorie a României, Bucureşti 24–27 Mai 2017 (Institutul Naţional al Patrimoniului, 2017), pp. 166–168. [Google Scholar]
- 159.C. Schuster, A. Morintz, R. Băjenaru, A. Ioniţă, A. Măgureanu, C. Ştefan, A. Popescu, D. Sârbu, D. Măgureanu, R Kogălniceanu, E. Dumitraşcu, M. Vasile, M. Constantin, C. Constantin, Peştera, com. Peştera, jud. Constanţa, “Peştera, com. Peştera, jud. Constanţa, Punct: Km 168+500–167+700 (Tumulul nr. 5 şi 6), km 168+600–169+100 (Tumulul nr. 3) şi km 169+800–171+000 (Valul mic de pământ)” in Cronica Cercetărilor Arheologice din România. Campania 2010, A XLIII-a Sesiune Naţională de Rapoarte Arheologice, Sibiu, 26–29 mai 2011 (Muzeul Brukenthal, 2011), pp. 215–217. [Google Scholar]
- 160.P. Włodarczak, T. Valchev, От степите към Балканите ...Проучване на надгробни могили в област Ямбол/Ze tepów na Bałkany...Badania kurhanów w obwodzie Jamboł/From the steppes to the Balkans...Excavation on burial mounds in Yambol district (Regional Museum of History Yambol Institute of Archaeology and Ethnology of the Polish Academy of Sciences, 2022). [Google Scholar]
- 161.S. Alexandrov, V. Slavchev, E. Tonkova, Rescue excavations of Bronze Age barrows in Vetrino region, northeast Bulgaria. Mater. şi Cercet. Arheol. 17, 5–48 (2021). [Google Scholar]
- 162.Gy. Gazdapusztai, Zur Frage der Verbreitung der sogenannten “Ockergräberkultur” in Ungarn. Móra Ferenc Múz. Évk. 65, 31–38 (1963).
- 163.I. Ecsedy, The People of the Pit-Grave Kurgan in Eastern Hungary (Fontes Archaeologici Hungariae, Akad. Kiadó, 1979). [Google Scholar]
- 164.T. Horváth, J. Dani, Á. Petö, L. Pospiezny, É. Svingor, Multidisciplinary contributions to the study of pit grave culture Kurgans of the Great Hungarian Plain, in Transitions to the Bronze Bronze Age. Interregional Interaction and Socio-Cultural Change in the Third Millennium BC Carpathian Basin and Neighbouring Regions, V. Heyd, G. Kulcsár, V. Szeverényi, Eds. (Archaeolingua, 2013), pp. 153–179. [Google Scholar]
- 165.C. Gerling, Prehistoric Mobility and Diet in the West Eurasian Steppes 3500 to 300 BC. An isotopic Approach (Topoi Berlin Studies of the Ancient World 25, De Gruyter, 2015). [Google Scholar]
- 166.K. Nagy, Csongrád-Kettőshalom-Petőfi Tsz. Régészeti Füzetek Ser. I/17 (Budapest, 1964), pp. 14. [Google Scholar]
- 167.I. Ecsedy, A new item relating the connections with the East in the Hungarian Copper Age (a Marosdécse type grave in Csongrád). A Móra Ferenc Múzeum Évkönyve 1971-2, 9–17 (1974). [Google Scholar]
- 168.J. Dani, T. Horváth, Őskori kurgánok a magyar Alföldön. A Gödörsíros (Jamnaja) entitás magyarországi kutatása az elmúlt 30 év során. Áttekintés és revízió (Archaeolingua Kiadó, 2012). [Google Scholar]
- 169.A. Marcsik, Data of the Copper Age anthropological find of Bárdos-farmstead at Csongrád-Kettőshalom. A Móra Ferenc Múzeum Évkönyve 1971-2, 19–27 (1974). [Google Scholar]
- 170.I. Ecsedy, Eine neue Hügelbestattung der “Grubengrab-Kultur” (Kupferzeit-Frühbronzezeit) in Dévaványa (Vorbericht). Archäologische Forschungen 1969. Antaeus – Mitteilungen des Archäologischen Instituts der Ungarischen Akademie der Wissenschaften 2, 45–50 (1971). [Google Scholar]
- 171.MRT 6: I. Ecsedy, L. Kovács, B. Maráz, I. Torma, Magyarország Régészeti Topográfiája 6. Békés megye régészeti topográfiája. A szeghalmi járás IV/1 (Akadémiai Kiadó, 1982). [Google Scholar]
- 172.Á. Bede, A tiszántúli halmok régészeti geológiai és környezettörténeti szempontú vizsgálati lehetőségei (Doctoral Dissertation, Department of Geology and Palaeontology, University of Szeged, Hungary, 2014); http://doktori.bibl.u-szeged.hu/id/eprint/2447/17/Bede%20Adam-tezisfuzet_magyar.pdf [accessed 20 March 2022].
- 173.Á. Bede, Beszámoló a Békés megyei Nagy-Sárrét halmainak felméréséről. (Report on mound survey in the Nagy-Sárrét region (Békés county, Hungary). Crisicum 8, 17–43 (2014).
- 174.A. Marcsik, The anthropological material of the pit-grave kurgans in Hungary, in The People of the Pit-Grave Kurgans in Eastern Hungary, I. Ecsedy, Ed. (Fontes Archaeologici Hungariae, Akadémiai Kiadó, 1979), pp. 87–98. [Google Scholar]
Associated Data
This section collects any data citations, data availability statements, or supplementary materials included in this article.
Supplementary Materials
Supplementary Text
Figs. S1 to S13
References