The article is devoted to the primary description of the third (?) terminal phalanx of the left (?) Homo brushes found in cultural deposits of Denisova Cave in 2011. In terms of its longitudinal dimensions, the width of the distal tuberosity, and the degree of relative flatness, it is close to the Neanderthal phalanx, but differs in a much larger latitudinal hypertrophy of the diaphysis. The latter can be interpreted as a more archaic feature. Comparison with the Upper Paleolithic sapiens Sungir 1, including using computed tomography, showed that with similar bone length and cross-sectional area in the middle of the diaphysis, the phalanx from Denisova Cave has a fundamentally different structure: massive walls and a narrow medullary canal. Microfocus radiography revealed the presence of a powerful sclerotization zone in its upper part, a massive trabecular network, and traces of active internal rearrangements of bone tissue, which is not analogous in the normal anatomy of modern humans. The owner of the terminal phalanx from Denisova Cave can be attributed to the carriers of the so-called archaic morphology with some unique features.

Keywords: Denisova cave, end phalanx of the hand, archaic morphology, computed tomography, microfocus X-ray.

Introduction

Archaeological research of Denisova Cave, painstakingly carried out over the past decades by the expedition of the Institute of Archeology and Ethnography of the Siberian Branch of the Russian Academy of Sciences, attracts the exclusive attention of scientists from all over the world. Here we can trace the evolutionary changes in the material complex of the inhabitants of Gorny Altai during the entire period from the Early Middle to the end of the Upper Paleolithic-280-10 Ka BP (Derevianko, 2009, 2011; Derevianko and Shunkov, 2009).

A colossal breakthrough in the knowledge of the taxonomic status of the oldest population of Southern Siberia was achieved thanks to paleogenetic studies. The terminal phalanx of the little finger of the hand belonging to the girl from layer 11 of Denisova Cave became a source for the isolation of mitochondrial paleoDNA (Krause et al., 2010). In this study, it was found that the Altai hominin shows almost 2 times more differences from modern humans than Neanderthals. According to the results of phylogenetic analysis, this lineage separated from the common trunk of Neanderthals and modern humans much earlier - about ca. 1 million years ago (or, to be precise, 1 313 500 - 779 300 l. n.).

The male who owned the tooth also found in layer 11 of Denisova Cave had a mitochondrial genome very similar to the one studied in the girl [Reich et al., 2010]. Decoding the nuclear DNA of this person allowed us to correct the conclusions: this group of hominins had a common ancestor with Neanderthals, diverging about 600 thousand years AGO, after which its history developed independently. Thanks to this publication, Denisovans received a stable name, although not framed by the Linnean nomenclature, since it would be premature to talk about the rank of this taxon now.,

as, in fact, about the rank of Neanderthals. The most striking result was the detection of Denisovian genes in the modern inhabitants of Melanesia. Consequently, the sapient ancestors of the latter at some point came into genetic contact with the Denisovans, and this is unlikely to have happened in Southern Siberia.

The hypothesis of a very wide range of Denisovans in Asia was confirmed after their genetic heritage was identified in 33 other modern populations of Asia and Oceania (Reich et al., 2011). The descendants of the Denisovans include Australian Aborigines, Polynesians, residents of the island of Fiji, eastern Indonesians, and one of the Negrito groups (Mamanwa in the Philippines). These findings were taken as evidence of gene flow from Denisovans to common ancestors of New Guinea Aborigines, Australians, and Mamanwa. At the same time, most modern East Asian peoples do not have a Denisovian heritage, which means that their ancestors could have settled this territory later. Researchers conclude that Denisovans had an exceptionally wide geographical distribution from Siberia to tropical Asia (Meyer et al., 2012). In another model study, a similar conclusion was made: a mixture of Denisovans (or rather related groups) and the ancestors of modern East Asians was confirmed (Skoglund and Jakobsson, 2011).

It is still difficult to assess all the possible consequences of hybridization of ancient sapiens with Asian owners of the relict genome. But, in particular, P. Parham and a large group of co-authors [Abi-Rached et al., 2011], studying the human leukocyte antigen system (HLA), described a rare variant of B-73. The Denisov girl had two similar variants on the corresponding section of the sixth chromosome. From this, it was concluded that outside of Africa, adaptive introgression of archaic alleles could have a significant impact on the immune system of modern humans.

Meanwhile, morphologically, Denisovans continue to remain a mystery. The tooth from which the DNA was extracted has some extremely archaic features [Reich et al., 2010]. If this is the third upper molar, then due to its huge size, it goes beyond the variations of all fossil taxa of the genus Homo, with the exception of N. habilis, N. rudolfensis, and is even comparable to the teeth of Australopithecus. If this is the second molar, then it is larger than in Neanderthals or early modern humans, but similar to the teeth of erectus and habilis. This suggests that other anatomical features of the Denisovian genotype carriers in Southern Siberia were far from modern.

It should be added that the Denisovans were not the only inhabitants of Southern Siberia in the period of 50-40 thousand years AGO. Thanks to archaeological (Derevyanko, 2007, 2009, 2011; Derevyanko, Markin, 1992; Derevyanko, Markin, and Shunkov, 2013), paleogenetic (Krause et al., 2007), and morphological studies of materials from the Okladnikov and Chagyrskaya caves (Mednikova, 20116; Buzhilova, 2011; Viola et al., 2012) the fact that Neanderthals settled Gorny Altai can be considered proven.

A natural question arises: if the results of genetic contacts between anatomically modern and Denisovian populations are revealed, is it possible to assume the possibility of similar hybridization of Denisovans with Neanderthals in the contact zone of Southern Siberia ca. In particular, this could explain the presence of separate, more archaic features in the Neanderthals from the Okladnikov Cave (Mednikova, 20116).

Data on the appearance of Denisova Cave inhabitants are still missing. In addition to the above-mentioned tooth, the proximal phalanx of the foot originates from layer 11, which morphologically occupies an equidistant position from Neanderthals and Middle Eastern sapiens of the Skhul group, while being distinguished by a colossal (archaic?)shape. the massiveness of the diaphysis [Mednikova, 2011a].

Material and methods

This publication describes a new skeletal material discovered in 2011 in the sediments of layer 12 of the eastern gallery of Denisova Cave (sector 6, layer 12, level 4, square G-3, sub-square A). This is a small tubular bone of the hand or foot with a strongly destroyed proximal part (Figure 1). The maximum fragment length (19.10 mm) can be used as a characteristic of articular length due to the fact that the dorsal part of the double proximal facet has been preserved.

Identification. The section of the diaphysis is podoval, not rounded, respectively, this is the phalanx of the hand, not the foot. The characteristic flatness of the distal end and the presence of tuberosity (tuberositas phalangis distalis) allow us to determine the studied bone as the terminal phalanx of the hand.

Determination of biological age. In the ontogenesis of modern humans, the growth of epiphysis of the distal phalanges of the hand occurs at 13.5 years in girls and at 16 years in boys [Schaefer, Black, Scheuer, 2009, p. 228]. The discovery from Denisova Cave demonstrates the absence of both recent synostosis and degenerative-dystrophic changes characteristic of the older age category. Features-

Fig. 1. Terminal phalanx of the hand from layer 12 of Denisova cave, a - dorsal surface; b - palmar; c - medial (?); d - lateral (?) surface; d - base; d - head.

The rapid development of tuberosity (see below) suggests a high level of biomechanical influence, which could contribute to early bone tissue rearrangements. Since there are no external involutive features, the Denisova Cave individual was quite young, within the category of adultus.

Beam detection. In the normal anatomy of modern humans, the nail phalanx of the thumb is well recognized due to its length and sufficient massiveness. However, since we have reason to assume a specific taxonomic status of the Pleistocene phalanx owner, as well as a different level of biomechanical stress from today, modern diagnostic criteria should be applied with caution. Therefore, conclusions about the bone fragment belonging to a particular finger and right or left hand will, if possible, be made after a comparative examination of the find against the background of the structure of similar bones of fossil hominids (material for comparison from the works: [Bonch-Osmolovsky, 1941; Musgrave, 1973; Trinkaus, 1983; Sladek et al., 2000; Shang, Trinkaus, 2010]). When elucidating the features of the internal structure, I relied on data from my own studies of the Neanderthal from Kiik-Koba and the Upper Paleolithic sapiens Sungir 1.

Methodology. Macromorphological measurements were performed with an electronic caliper with a resolution of 0.01 mm or 0.0005" (Digital Electronic Caliper, model GMC-190). The study of the internal structure was carried out after scanning on a Discovery HD 750 volume tomograph (manufactured in the USA) and with microfocus radiography (developed by the Department of Electronic Devices and Devices of the St. Petersburg State Electrotechnical University and the company EltehMed),

Morphological features of the phalanx

At the first approximation, the terminal phalanx from Denisova Cave is characterized by a rather massive diaphysis and pronounced hypertrophy of the distal tuberosity, which distinguishes it from its modern counterparts. The dorsal surface of the diaphyseal part is straight, smooth and even, with the exception of the lateral surfaces of the prin-ulnar region, where there is a cluster of large feeding holes. The palmar surface of the phalanx carries even more feeding holes-both in the tuberosity area and in the central part of the diaphysis. Anatomically, it is slightly concave.

Despite the extensive damage at the base of the phalanx, enough can be done.-

conclusions about its size and configuration. The following measurements (mm)were obtained:

Special attention was paid to the comparison with the reference group of Neanderthals from the Shanidar Cave in Iraq and with the "anatomically modern" man from the Tianyang Cave in China. The longitudinal dimensions of the terminal phalanx of the thumb in Shanidarians (N 3-6) vary within 23.6 - 24.7 mm (Trinkaus, 1983, p. 281). This suggests that the phalanx from Denisova Cave belonged not to the first, but to the second - fifth rays. According to the ratio of articular length and height in the middle of the diaphysis, the bone from Denisova cave is closest to the second (?) The left nail phalanx of the Shanidar individual is 6. At the same time, it shows an exceptional width in the middle of the diaphysis, significantly exceeding the variations within the Shanidar group (5.3 - 7.4 mm) or the value characteristic of the Tianyang person. This trend is also confirmed by the distribution of the cross-sectional index values in the middle of the diaphysis (Fig. 2). Only the terminal phalanges of the third and fourth fingers of the Shanidar 4 individual are sufficiently close to the Denisova cave bone in terms of the degree of flatness and relative width of the diaphysis. The Crimean Neanderthal from Kiik-Koba is noticeably inferior to the Denisovan in terms of the diaphyseal massiveness of the terminal phalanx of the third finger (latitude index of massiveness 37.3 vs. 44.4; cross-section index 68.0 vs. 52.6). The massiveness in the middle of the diaphysis of the phalanx from Denisova Cave is also indicated by the circumference value, which goes far beyond the variations in the reference Shanidar group (17.5-20.5 mm).

It is difficult to judge the development of the proximal part; however, the greatest height of the base falls within the range of variability of the Iraqi sample of Neanderthals (6.2-7.6 mm).

The distal edge is noticeably flattened, in terms of size, the bone in this part is similar to the terminal phalanx of the little finger of Shanidar 3 or the fourth (to a lesser extent fifth) finger of Shanidar 4.The width of the distal tuberosity excludes the possibility of this bone belonging to the little finger. It significantly differs from the current values of this feature of all four rays, being located at the locus of intermediate values of the tuberosity width of the third finger of Neanderthals (Fig. In particular, the tuberosity of the Denisova Cave phalanx is wider than that of Shanidar 3, 5, and Regurdu 1, but narrower than that of Shanidar 4, Amud 1, and Kiik-Koba 1. However, quite broad variants could also occur among the phalanxes of the second ray (Fig. 4).

2. Cross-sectional index in the middle of the diaphysis of the terminal phalanges of the hand in some Pleistocene hominids of Asia (the ray number is indicated in parentheses). 1-Denisova cave; 2-Shanidar 3 (3); 3-Shanidar 3 (5); 4-Shanidar 4 (3); 5-Shanidar 4 (4); b-Shanidar 5 (2); 7-Shanidar 5 (3); 8-Shanidar 6 (2); 9 - Tianyang (2).

3. Width of the distal tuberosity of the terminal phalanges of the third ray of the hand. 1-Denisova cave; 2-Shanidar 3; 3-Shanidar 4; 4-Shanidar 5; 5-Amud 1; 6-La Ferrasi 1; 7-Kiik-Koba 1; 8-Regurdu 1; 9-modern Europeans.

4. Width of the distal tuberosity of the terminal phalanges of the second ray of the hand. 1-Denisova cave; 2-Shanidar 5; 3-Shanidar 6; 4-Amud 1; 5-La Ferrasi 1; b-Kiik-Koba 1; 7-Krapina 206.12; 8-Tianyang; 9-modern Europeans.

5. Articular length and width in the middle of the diaphysis of the terminal phalanges of the hand in hominids of different taxonomies (the ray number is indicated in parentheses). 1-Denisova cave; 2-Shanidar 3 (3); 3-Shanidar 3 (5); 4-Shanidar 4 (3?); 5-Shanidar 4 (4?); b-Shanidar 5 (2); 7-Shanidar 5 (3?); 8-Shanidar 6 (2?); 9-Kiik-Koba (3); 10-Kiik-Koba (4); 11-Tianyang (2); 12-Omo 1 (5?); 13 - Russkie (3); 14-Russkie (4); 15-Dolni Vestonice 3 (2 3); 16-Dolni Vestonice 3 (2 or 3); 1 7-Dolni Vestonice 3 (4); 18-Dolni Vestonice 3 (5); 19-Dolni Vestonice 13 (3 or 4); 20 - Dolni Vestonice 13 (3 or 4); 21-Dolni Vestonice 15 (2 or A); 22-Dolni Vestonice 16 (2 or 3); 23-Dolni Vestonice 16 (2 or 3); 24 - Dolni Vestonice 16 (2 or 3); 25 - Dolni Vestonice 16.

b. Width and height in the middle of the diaphysis of the terminal phalanges of the hand in hominids of different taxonomies. See Figure 5 for additional information.

Table 1. Dimensions of the hand phalanx from Denisova cave against the background of the average parameters of the terminal phalanges of the third ray in modern Altai peoples, mm*

* Comparative data on: [Mednikova, 1988].

If the bone from Denisova cave is from the second finger, it is inferior in latitudinal development of tuberosity only to the corresponding phalanges of the Neanderthals Kiik-Koba 1 and Amud 1.

In terms of body structure, the bone from the Altai is the most massive, although its length is shorter than in some of the largest variants typical of Neanderthal males (Fig.

According to the ratio of height and width in the middle of the diaphysis, the Denisova cave phalanx occupies the farthest position from the corresponding bones of representatives of modern anatomy, approaching Neanderthal (Figure 6). Comparison with the parameters of the terminal phalanges of the modern population of the Altai region shows that it is not only longer, but also much wider in the distal and central parts (Table. 1).

Discussion

Returning to the questions of direct identification of the bone from Denisova Cave, it can be stated that according to various measuring features, it belongs to the second or fourth fingers. The little finger, which even in Neanderthals is characterized by a narrower distal tuberosity, should probably be excluded.

G. A. Bonch-Osmolovsky [1941, p.109], quoting the early work of V. Pfitzner, emphasized that the most important distinguishing feature is the "body thickness" of the phalanx, which is greatest for the third ray and decreases in the row: the fourth, second and fifth rays. Unfortunately, the discovery from Denisova Cave is a single one, it cannot be compared with other terminal phalanges of the hand of the same individual. Nevertheless, the exceptional massiveness of the diaphysis is quite consistent with the version that it belongs to the third ray. It is noteworthy that according to the cross-section index, the phalanges of the third and fourth fingers of the Neanderthal man Shanidar 4 are closest to the Altai bone.

Based on the size, the version that the phalanx from Denisova Cave belongs to the index finger cannot be completely rejected. But G. A. Bonch-Osmolovsky [Ibid.] pointed out that the second phalanx is always more or less pointed at the end. In this case, such a "sharpness" is not observed, therefore, we are talking about the third or fourth finger.

When determining the side, following V. Pfitzner, G. A. Bonch-Osmolovsky [Ibid.] recommended paying special attention to the asymmetry of the nail blades, most often developed weaker on the outer side (radial in the second and third rays and ulnar in the fourth and fifth). In addition, the radial edge of the phalanx of the third finger, when viewed from the distal end, deviates to the back side. It seems that the Denisova cave find, from this point of view, finds the greatest agreement in the structure of the left nail phalanx of the third ray in the Crimean Neanderthal.

Having determined the anatomical identity of the bone, we will try to identify morphological trends that can clarify its taxonomic status. Unfortunately, the paleontological record of the distal phalanx structure in human evolutionary progenitors is not very detailed. The second and third ray phalanges of Homo habilis from Olduvai were studied (Susman and Creel, 1979). They differ from modern ones by their greater thickness in the middle of the diaphysis, and there is an increase in both the width and height of the body.

End phalanx of the little finger (?) According to researchers (Pearson, Royer, and Fleagle, 2008), Omo 1 in the oldest H. sapiens combines a broad base, an accentuated attachment area of T. flexor digitorum profundus, and a broad diaphysis with a "flattened" head. However, as can be seen from the comparison made (see Figures 5, 6), in terms of the ratio of articular length, width, and height in the middle of the diaphysis, it is located in the field of variability of the values of Upper Paleolithic sapiens, sharply differing from the find from the Denisova Cave and from the corresponding Neanderthal bones. The morphology of the latter's hand has been discussed repeatedly [Aiello and Dean, 2006]. In particular, the observations of G. A. Bonch-Osmolovsky retain their significance in the Russian scientific literature [1941, p. 4]. In the Neanderthal from Kiik-Koba, all the terminal phalanges are close to the modern maximum in length, and the values of the body width, base, and especially the head are higher than the largest modern ones. The absolute height of the body, base, and head of the Kiikkobin is close to or exceeds the maximum values of modern humans. The terminal phalanges of the Neanderthal hand are flattened. Cro-Magnons, according to Bonch-Osmolovsky, showed the opposite trend - with an increased absolute height, their phalanges give the impression of being relatively thicker than modern ones, not to mention Neanderthal ones.

A comparison was made between the features of not only the external but also the internal structure of the phalanx from Denisova Cave and a similar bone from the Upper Paleolithic sapiens Sungir 1 (Table 2). While the overall dimensions (length of the phalanx, maximum height of the base) are similar, the Altai find is characterized by a wide distal tuberosity, absolutely and relatively low and wide diaphysis. Even more striking differences can be seen in the internal structure of the phalanges. With a similar cross-sectional area (TA)

Table 2. Comparative characteristics of the structure of the terminal phalanges of the third ray of the hand in Homo from Denisova Cave and the Sungir man 1

these fossil bones differ sharply in the degree of corticalization (%CA). Heavy walls of the Altai Homo tubular bone correspond to parameters typical of archaic morphology carriers, among which Neanderthals from the Okladnikov Cave were among the most massive, starting from childhood [Mednikova, 20116].

The use of microfocus radiography, which allowed us to obtain high-quality enlarged images of the terminal phalanges of fossils and modern forms, revealed new facets of the peculiar internal structure of the Denisova Cave bone (Fig. 7). For example, when comparing the end of the bone with the end of the Denisova Cave (Fig.-

7, 8). When compared with the phalanges of the present - day 20-year - olds, it shows not only a coarser coarse - cell structure, but also a rather extensive sclerotization in the upper part of the diaphysis, as well as repeated erosion of the distal tuberosity (Fig. 55-year-old representatives of N. sapiens bone from Denisova cave also finds no analogies. However, in modern populations, some diseases are accompanied by the so-called dentoid epiphysis, in which, along with osteosclerosis, different levels of phor are recognized-

7.Microfocus X-ray images of the terminal phalanges of fossils and modern humans. Anterior-posterior projection, a-Denisova cave; b-Kiik-Koba 1 (left hand, 3rd ray); c-Sungir 1 (right hand, 3rd ray); d-modern 20-year-old man (right hand, 3rd ray); e-modern 55-year-old male (right hand, 3rd beam).

8. Images of the terminal phalanges obtained by microfocus X-ray. Side projection, a-Denisova cave. In the distal part of the phalanx, there is an area of sclerotization, in the proximal part - massive irregular trabeculae; b-Sungir 1 (right arm, 3rd ray). The medullary canal is clearly visible, and the structure of the trabeculae in the proximal part is more ordered. There are up to six Harris lines that reflect growth retardation in childhood.

mri of trabeculae. Differential diagnosis includes hypothyroidism, osteopetrosis, familial finger arthropathy (avascular necrosis or Thyemann syndrome), etc. [Castriota-Scanderberg. Dallapiccola, 2005, p. 428]. Local erosion of the distal tuberosity can mean an inflammatory process and necrotization of the nail region, for example, in psoriatic arthritis, which is also associated with bone compaction to a "tooth-like" state [Jacobson et al., 2008, p. 384]. To this list, it is worth adding necrosis due to frostbite, a diagnosis made by D. G. Rokhlin to the Kiik-Koba Neanderthal, which was recently confirmed by a new study [Buzhilova et al., 2008].

It is known that an anatomically modern person with age has a decrease in the number of bone plates and bone rarefaction (osteoporosis), as well as excessive bone formation in the form of growths (osteophytes). Cases of osteolysis, a pathological process accompanied by bone resorption, have been clinically described in old age with extensive osteoporosis. From this point of view, the picture revealed by microfocus X-rays fits into the framework of advanced, although not senile, changes without signs of osteoporosis.

So, in a person from Denisova Cave, the condition of bone tissue corresponding to an older age category is recorded than it is visually determined, and there is reason to assume an insurmountable pathological process. It seems likely that the detected" wear and tear " of the distal phalanx reflects not the aging of the individual, but the working hypertrophy under enormous biomechanical influence. Prior to histological examination, the question of the biological age of this person should preferably be left open.

As a result, if the 2011 find from layer 12 of Denisova Cave represents the third (?) end phalanx of the left hand of Noto, according to its parameters (elongation, large width of the body and distal tuberosity, relative flatness) it fully corresponds to the known features of Neanderthal morphology. Moreover, the degree of relative flatness and latitudinal development of the diaphysis of this phalanx exceeds the level typical of both Middle Eastern (Shanidarian) and European Neanderthals. The hypertrophied massiveness of the Altai bone diaphysis can also be regarded as a more archaic feature. In this connection, we would like to recall the proximal foot phalanx found in layer 11 of Denisova Cave in 2010 (Mednikova, 20116). It is characterized by an extraordinary massiveness, including an exceptional width of the diaphysis. Again, the study of the bones of the postcranial skeleton of Neanderthals from the Okladnikov cave revealed a systemic morphological trend, which manifests itself in a specific expansion of the articular surfaces and latitudinal patellar hypertrophy (Mednikova, 2011a).

In sum, these scattered and so far few facts can mean both the presence of a corresponding mutation in an isolated population of Neanderthals in Altai, and their genetic contact with a more relict population, from which the morphogenetic program of latitudinal development could be borrowed. The mysterious Denisovans remain contenders for the role of such a more archaic population in Southern Siberia.

Conclusion

The tubular small bone from layer 12 in Denisova Cave represents the terminal phalanx of the left (?) brushes are most likely of the third or fourth ray. It belonged to an adult, most likely a man. According to the set of features (elongation, large body width and distal tuberosity, relative flatness), the phalanx is close to the corresponding elements of the skeleton of the Anterior Asian and European Neanderthals.

At the same time, summing up the preliminary results. it is necessary to distinguish the relative and absolute massiveness of the terminal phalanx diaphysis from the Denisova cave, which is manifested in hypertrophy of the width and, as a consequence, the perimeter. Among modern forms, such features cannot be traced or may be extremely rare. Along with the increased development of distal tuberosity, the massiveness of the diaphysis can be evidence of working hypertrophy of bone tissue under the influence of very intense physical exertion. However, since the Denisova Cave individual stands out in this respect even against the background of the Neanderthal population, it cannot be excluded that such a development of small tubular bone reflects the genetic peculiarity of the representative of the Altai population of paleoanthropes or even a special taxonomic status. The use of computed tomography and microfocus radiography revealed extreme massiveness of the diaphysis walls, abnormal bone compaction (sclerotization), along with the formation of massive trabeculae. Erosion of the distal part of the head does not exclude necrotic processes of various etiologies.

Acknowledgements

I take this opportunity to express my deep appreciation for the opportunity to explore the unique material from Denisova Cave to A. P. Derevyanko and M. V. Shunkov. I am grateful to T. S. Baluyeva for the opportunity to examine the skeleton of Sungir 1, V. E. Sinitsyn for cooperation in performing computed tomography, N. N. Potrakhov and V. B. Bessonov for invaluable assistance in microfocus radiography. The skeleton from Kiik-Koba was X-rayed in 2008 together with V. I. Khartanovich, N. N. Potrakhov, A. P. Buzhilova and M. V. Dobrovolskaya.

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The article was submitted to the Editorial Board on 01.06.12.

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