UDC 572

M. B. Mednikova ¹, M. V. Dobrovolskaya ¹, A. P. Buzhilova ¹, M. N. Kandinov ²

¹ Institute of Archeology of the Russian Academy of Sciences

19 Dm Ulyanova str., Moscow, 117036, Russia

E-mail: medma_pa@mail.ru; mk_pa@mail.ru; albu_pa@mail.ru

² Vernadsky State Geological Museum of the Russian Academy of Sciences

11/2 Mokhovaya St., Moscow, 103009, Russia

The article is devoted to the introduction into scientific circulation of a fossil find-the Homo humerus from Khoroshevsky Island near Khvalynsk, which was considered lost for 80 years. The structural features of the humerus studied by classical osteometry, radiography, computed tomography, histology, and paleopathology are discussed. Based on these data, it is concluded that it belongs to an adult paleoanthrope, most likely male, who had a specific hormonal status that does not find complete analogies in the normal and pathological physiology of modern humans.

Keywords: evolutionary anthropology, archaic morphology of the postcranial Homo skeleton, paleofauna of the Volga region, massiveness of the structure, paleohistology, computed tomography.

In 2007, while working in the collections of the Vernadsky State Geological Museum of the Russian Academy of Sciences, we found a fragment of a human humerus that was previously considered depasportized. A more detailed examination revealed the label of this storage unit, and it became clear that the skeletal fragment can be attributed to the humerus found near the town of Khvalynsk**. The geological and paleontological contexts of the discovery of this artifact indicate that this is a fossil object that belongs in a wide range to the period from the Late Assyrian to the Aurignacian-Solutrean time inclusive (Kandinov et al., 2008). Therefore, at this stage of the study, even if we do not yet have the results of radiocarbon dating***, we consider it our duty to give as detailed a morphological description of the bone fragment as possible. For this purpose, in addition to the visual description, methods of radiography, computed tomography and histology were used as part of the paleoanthropological study****.

* The study was funded by the Bioarchaeological Institute (San Francisco).

** Label text: "A fragment of a human skull and humerus were found in 1927 on Khoroshevsky Island on the left bank of the Volga River near Khvalynsk, Saratov Province, by V. F. Orekhov and K. Y. Gross." Below, in a different handwriting, it says: "No skull."

*** A sample of bone tissue for direct radiocarbon dating was transferred to the laboratory of the University of Oxford (Radiocarbon Accelerator Unit).

**** Radiography of the sample was performed by M. V. Dobrovolskaya and M. B. Mednikova; computed tomography was performed at the Radiation Diagnostics Department of the N. I. Priorov Central Institute of Traumatology and Orthopedics; histological examination was performed by M. V. Dobrovolskaya.

Results of a comprehensive study of the humerus

The majority of researchers considered the fossil finds from Khoroshevsky Island to be relatively late, but at the same time rather archaic in structure. Therefore, the morphological characteristics of the shoulder fragment we found are particularly important. Taking into account what has been repeatedly stated by various researchers (e. Eikngtedt, M. A. Gremyatsky, V. P. Alekseev, etc.) opinions on the primitive and even neanderthaloid features of the Khvalyn skull cap, found only 2 m from the humerus, our main task was to determine to which variant of the structure, archaic or modern, this bone belongs.

Before proceeding to the description of the Khvalyn fragment, let us recall some characteristic features of the humerus structure in Neanderthals. There are three main features that place Neanderthals among the extreme variants of morphological variability of modern humans [Aiello and Dean, 2006]. As noted by L. Aiello and K. Dean, based on the work of E. According to Trinkaus (1983, p.228), the first such feature is the impressive development of ridges at the site of attachment of the deltoid and pectoralis major muscles, reflecting the strong development of the latter in both European and Asian forms. In Neanderthals Shanidar 3, 4, and 6, the deltoid tuberosity is formed by a well-defined, uneven, slightly protruding ridge, which is sufficiently pronounced on the right humerus of Shanidar 3 to form a furrow between this tuberosity and the anterior edge of this bone. Such a furrow along the deltoid tuberosity is characteristic of both some European Neanderthals and modern humans [Ibid., p. 229]. However, other authors have emphasized that the deltoid tuberosity in Neanderthals is usually narrow and weakly protruding (Endo and Kimura, 1970), and it is also located more proximally than in modern humans. The area of attachment of the pectoralis major muscle, on the contrary, was very wide [Trinkaus, 1983, p. 229].

The second sign is flattening of the humerus diaphysis in the mediolateral direction. Apparently, this feature of Neanderthal postcranial morphology was first identified by the most authoritative Russian anatomist. Sinelnikov, describing the bone of a child from Teshik-Tash: "... in the proximal part it (the body of the bone. - Auth.) has one feature, due to which the determination of its belonging to the right or left humerus would be difficult if there was no place left in the distal part of the transition to the internal epicondyle. This feature is that at the level of roughness, the bone is flattened in the mediolateral direction. This flatness is so great that it even suggests postmortem deformity, although there is no other evidence for this... In any case, the proximal part of the bone has neither a cylindrical shape nor the shape of an irregular polygon, as in the modern type, but rather an ellipsoid shape. This feature is not clearly displayed in figures, but it is clearly visible on the bone" (Sinelnikov and Gremyatsky, 1949, p. 130).

The third sign is a very massive diaphysis when compared with modern humans. Moreover, this massiveness is recorded not so much by traditional external measurements, but rather by considering excessively thick compacts (Rokhlin, 1949; Ben-Itzhak, Smith, and Bloom, 1988). On average, Neanderthals (both males and females) had a wider cortical layer of the humeral diaphysis compared to modern Sapiens, with the highest values of compaction thickness found on the right upper limb of Neanderthal males. In addition, Neanderthals show not only a noticeable internal massiveness of tubular bones when compared with modern humans, but also a much more pronounced sexual dimorphism for this trait. Based on these observations, it was concluded that Neanderthals had more muscle mass and more intense load on the upper limb than Homo sapiens. They also differed from modern humans in noticeably different ways in the use of the right hand between the sexes. It has been suggested that regular javelin throwing by Neanderthal males could have led to such obvious hypertrophy. Such sexual dimorphism did not affect other parts of the Neanderthal postcranial skeleton. The longitudinal and girth dimensions of most postcranial bones lie within the range of variations in modern humans (Trinkaus, 1983; Heim, 1983). However, as noted by L. Aiello and K. According to Aiello and Dean (2006), hypothetical behavioral differences suggest that indicators of internal massiveness of Neanderthal tubular bones are generally more informative for distinguishing them from carriers of modern morphology.

Bone safety

The humerus from Khoroshevsky Island is right (Fig. 1). Its epiphysis is destroyed, but most of the diaphysis is preserved. The largest fragment length is 247 mm. The surface of the diaphysis was eroded and partially "doused" with water. The color is dark gray, turning into black, "muddy". Obviously, originally cor-

1. Fragment of the humerus from Khoroshevsky Island. Front view.

the tical layer was developed more strongly and, accordingly, the determined values of the diameters were larger. Nevertheless, the area of deltoid tuberosity is fixed quite clearly, which makes it possible to recreate some reference points necessary for a comparative description. In particular, the middle of the diaphysis is determined at the end of the deltoid tuberosity.

Macromorphological description

When visually examined, the humeral diaphysis gives the impression of gracile. The body of the bone is smooth, not bent. Dimensions (mm):

Unfortunately, the maximum width at the attachment point of m. pectoralis major cannot be determined by measurement due to surface damage.

For a comparative assessment of the external massiveness of the Khvalynsky diaphysis, we used data on the morphology of the right humerus of the Upper Paleolithic male Sungir 1 and a representative of the Bronze Age (Volga region, XVII century BC), characterized by extremely hypertrophied development of the musculoskeletal relief of the upper limb.

The values of the index of the mid-diaphysis cross-section in the Khvalyn individual (75.6) are significantly smaller than in the Sungir and Bronze Age individuals, which indicates its greater flatness in the mediolateral direction (Fig. 2). The absolute dimensions of the Sungir individual are larger than in the Khvalyn individual: this applies to both circles and diameters (Fig. 3, 4). As we mentioned above, a strong flattening of the proximal part of the diaphysis in the mediolateral direction was previously observed in a nine-year-old Neanderthal from Teshik-Tash [Sinelnikov and Gremyatsky, 1949, p. 130], which means that this feature appeared relatively early in the ontogenesis of paleoanthropes.

At the same time, a noticeable contrast between the thickness in the middle of the diaphysis and at the level of its maximum development in the Khvalyn individual (the cross-section index in this place reaches 95.65) indicates an increase in the musculature of the upper limb. With the external miniaturization of the bone, the relative extent of the deltoid tuberosity in the vertical direction is noteworthy.

The lateral contour of the bone below the deltoid tuberosity is concave rather than convex. Hambucken (1983) considered this feature to be characteristic of Neanderthal morphology. Another typical Neanderthal feature is a furrow delineating the anterior edge of the deltoid tuberosity (for example, in Shanidar 3 (Trinkaus, 1983))., -

2. Values of the index of the cross-section of the middle of the humerus diaphysis (Martin index 6 : 5,%) of the Khvalyn individual (a), Cro-Magnon from Sungir (b), and representative of the Bronze Age (c).

3. Values of the largest and smallest diameters in the middle of the diaphysis of the Khvalyn (a) and Sungir (b) humerus bones (Martin signs 5 and 6, mm, respectively).

it cannot be fixed due to extensive destruction of the surface layer.

A broader comparison of the diaphyseal parameters of the humerus shows that "Neanderthal" dimensions are part of the boundaries of modern human variability. The cross-sectional shape of the middle of the diaphysis may be more indicative, but there may also be exceptions. The cross-sectional index of the Khvalynets shows a median, "undifferentiated" position against the background of the spread of values for fossil hominids (Table 1). Among the early forms, it is most closely related to the KNM-WT 15,000 erectus, some Neanderthals (Kebara 2, La Kina 5, Regurdu 1, Neanderthal) and Cro-Magnons (Grot Children 4, 5; Przedmzy 4, 5; Barma Grande 2; Aren Candide 10). Velichi-

4. Values of the mid-diaphysis circumference and the smallest diaphysis circumference of the Khvalyn (a) and Sungir (b) humerus bones (Martin signs 7a and 7, mm, respectively).

Table 1.

Indications of the mid-diaphysis cross-section in fossils and modern hominids (values for right bones, except for the specified exceptions)*

* Calculated according to various authors (Pearson, 1997; Trinkaus, 1983; Endo and Kimura, 1970; et al.).

5. Differentiation of hominid fossils according to the index of the cross-section in the middle of the diaphysis (index 6 : 5 according to Martin) and the minimum circumference of the humerus (index 7 according to Martin).

The numbers correspond to the ordinal numbers in Table 1.

More than 80 % of them characterize the African "Heidelberg man" (Kabwe E898) and the Middle Paleolithic "anatomically modern" Africans (Omo 1 and Border Cave). The Levantine Mousterian Skhul-Kafzeh groups have similar, but mostly lower indicators in comparison with Omo 1 (but higher than in Khvalynets).

We can say that Neanderthals generally show the greatest flatness of the middle of the diaphysis (we have already listed all the exceptions above). Sapiens of the Early and Middle stages of the Upper Paleolithic have the most rounded section of the diaphysis (Cro-magnon 4293 and 2; Przedmaty 10 and 14, Pato 231; Neuessing, Oberkassel 1; Aren Candide 2). It is noteworthy that during the last temperature minimum in Europe, the values of the considered trait fall to quite "Neanderthal" (Chancelade, Pato 230, Continenza, Aren Candide 3, 5, 12). At the same time, the population of hot regions (Ain Gev 1, Nazlet Kater, Wadi Kubbaniya) has the same high values of this index as the early carriers of modern morphology in the Middle Paleolithic of Africa.

Unfortunately, the circumference of the humerus diaphysis also does not provide much for attribution of the fragment under study, since this feature is very variable and depends on many factors. The smallest circumference of the Khvalyn bone is identical to that of Le Regurd 1; similar values are found in Skhul 4, Mladech, Shanidar 1, Przedmmi 4 and 14.

By the ratio of the cross-sectional index and the smallest circumference of the diaphysis, the fragment under study occupies an intermediate position between Neanderthal and Madeleine materials, on the one hand,and most anatomically modern sapiens, on the other (Fig.

X-ray morphological characteristics

X - ray images of the bones of representatives of the Upper Paleolithic population - Sungir 1-3 (Mednikova, 2000a-b) - served as the main comparative material. Images of the Khvalynsky fragment were taken in anteroposterior and lateral projections on the Arman device at a focal length of 80 cm, 50 kV mode, 6 mA / s, on a domestic RT X-ray film.

The X-ray image was used to study the ratio of the compacts and the channel of the bone-brain space at the level of the middle of the bone diaphysis (in this plane, during osteometric examination, the largest and smallest diameters are measured, the circumference of the middle of the diaphysis-signs 5, 6 and 7a according to Martin, respectively). X-ray bone width and channel width (mm) were measured on the radiograph. Subtracting the second value from the first value, we get the total width of the compact layer.

The compactification index (IC) of all tubular bones is calculated by the formula

IC = WCL : WD x 100%,

where WCL is the total width of the compact layer; WD is the X-ray width of the bone.

To expand the possibilities of comparative analysis, a method was used to determine the degree of compaction of the humerus, proposed by A. Stirland [Stirland, 1998]. For this purpose, measurements were carried out at the level of the end of the inter-tubercular sulcus, the external development of which is largely determined by the functional activity of the pectoralis major muscle m. pectoralis major, in the images taken in the anteroposterior projection. Next, the total thickness of the compact layer was calculated. The area of the cortical layer in this section was determined by the formula for a circle

CA = π / 4 (WD2-M2) = 0.785 (WD2^{) where} is the channel width.

Table 2.

Comparative X-ray structural characteristics of the Khvalyn humerus

Taking into account the uniqueness of the studied material, we tried to give the most differentiated assessment of bone compaction. Therefore, the development of the anterior, posterior, medial, and lateral walls was evaluated (Table 2).

In the anteroposterior projection on the X-ray image of the Khvalynskaya find in the middle of the diaphysis, the X - ray width of the bone is 19 mm, the lateral compaction is 6 mm, the medial one is 5 mm, and the canal is 8 mm; IC is 57.9 %. At the site of the maximum development of deltoid tuberosity, the X - ray width of the bone is 22 mm, the lateral compact is 6 mm, the medial one is 3 mm, and the medullary canal is 13 mm; IC is 40.9 %. In the area under the deltoid tuberosity, the X-ray width of the bone is 18 mm, the channel width is 6 mm; IC is 66.67%.

In the lateral projection, the X - ray width of the bone in the middle of the diaphysis (at the end of the deltoid tuberosity) is 24 mm, the anterior compact is 7 mm, the posterior compact is 5 mm, and the canal is 12 mm; IC is 50.0 %.

When compared with Sungir materials, the internal massiveness of the Khvalyn bone is clearly visible: it is fixed due to the relative thickening of not only the medial and lateral, but also the anterior and posterior walls of the diaphysis. The side walls are most thickened, as can be seen when comparing the compaction indices at the level of the middle of the diaphysis (Fig. 6).

The differences between the Khvalyn individual and the Sungir people are less noticeable when considering the X-ray images taken in the lateral projection (Fig.

6. Compaction indices (IC,%) of humerus bones from Khvalynsk (X) and Sungir (C), taken in the anteroposterior projection. Middle of the diaphysis.

a - right, b-left.

7. Compaction indices (IC,%) of humerus bones from Khvalynsk (X) and Sungir (C), taken in the lateral projection. Middle of the diaphysis.

a - right, b-left.

8. Area of the cortical layer of humerus bones from Khvalynsk (X) and Sungir (C) at the level of m. pectoralis major (CA, ^mm2).

the compaction index in the middle of the diaphysis of the Khvalynskaya find coincides with the value of the trait in the Sungir girl (NW). However, given the extreme gracefulness of the lateral walls of the diaphysis in the anteroposterior norm in the individual Sungir 3, we can talk about a completely different structure of the cross-section of the diaphysis.

In general, when compared with the Upper Paleolithic materials from Sungir, the Khvalyn humerus is more "archaic" (Figure 8). Despite its external miniaturization, it is much more massive inside. Moreover, taking into account the damage to both the inner and outer surfaces of the cortical layer of this bone, the differences were more pronounced.

Results of computed tomography examination

Computed tomography (CT) is a method of radiation diagnostics associated with obtaining layer-by-layer X-rays of the study area of interest, which makes it possible to study the internal structure of the bone in more detail. Axial sections with a thickness of 1 mm and their reconstructions with a thickness of up to 0.2 mm were obtained on the Aura Philips spiral CT scanner, which allows creating multiplanar three-dimensional images in real time. The distances between anatomical points were determined using Centricity DICOM Viewer (version 2.2, 2004).

Figure 9. Contours of the right humerus bones in the middle of the diaphysis.

1-Khvalynsk; 2-Neanderthal 1; 3-La Kina 5; 4-control sample of the Bronze Age from the Volga region.

The CT-scanned image was compared with the control sample - data from a tomographic examination of the right humerus of a 20-24 - year-old Bronze Age man from the Volga region, who, judging by the extremely high indicators of relief development at the attachment points of m. deltoideus, m. pectoralis major, experienced increased biomechanical pressure associated with the daily motor activity of a blacksmith-foundry worker [Mednikova, 2001, fig. 8 to ch. 8] (his professional activities are indicated by the accompanying burial equipment). This object was chosen in order to assess the nature of the formation of the cortical layer of diaphysis with a greater measure of reasonableness, as well as to evaluate the variants of bone rearrangements during increased physical exertion.

When comparing cross-sections in the middle of the diaphysis, attention is drawn to the different shape of the contours (Fig. 9). The Khvalyn bone is subcircular in cross-section, with a distinct parallel flattening in the mediolateral plane. The canal cavity has an irregular shape due to postmortem local damage, but is generally close to ellipsoid. The shape of the external section of the diaphysis of the control sample is rounded in the posterior part; a deltoid tuberosity is very clearly delineated, forming a characteristic external protrusion. On the Khvalyn bone, nothing like this is observed, but a uniform and powerful increase in internal massiveness is recorded. The space of the medullary canal in humans of the Early Metal epoch has a rounded cross-section, in addition, the bone marrow occupied much more space than that of the Khvalynets (earlier, when comparing the Khvalynsky find with the Sungir materials, according to radiography data, we noted a similar trend).

Thus, even at the first assessment, there is an impression of greater internal massiveness of the Khvalyn bone compared to the modern one (with the exception of the hypertrophied thickness of the lateral wall in the control sample). At the same time, the external structure associated with the place of attachment of the deltoid muscle is not accentuated, while in a Bronze Age person the corresponding tuberosity forms a completely defined structural unit, clearly fixed on the section in the middle of the diaphysis. In addition, the localization of deltoid muscle attachment differs: in the control sample, this area is shifted by about 60° compared to the Khvalyn bone, located in the lateral quadrant of the diaphysis. In the normal anatomy of modern humans, the deltoid tuberosity is located on the anterior lateral surface of the humerus body, slightly above the nutrient opening foramen nutricium, leading to the distally directed nutrient channel canalis nutricius. Back door

Table 3.

Wall thickness in the middle of the humeral diaphysis determined by radiography and computed tomography, mm

See Table 4.

Indicators of the internal structure of the humerus diaphysis

the surface lying behind the deltoid tuberosity is indicated by the sulcus nervi radialis radial nerve furrow, which has a spiral course and is directed from top to bottom and from inside to outside. Due to superficial damage, this furrow on the Khvalyn bone is not fixed, however, the identification of the anteroposterior projection of the diaphysis due to the orientation of the lower metaphysis is not in doubt. The deltoid tuberosity occupies an anteromedial position. Thus, the contours of the cross-section of the Khvalyn bone and the localization of the m. deltoideus attachment site on it coincide with those characteristic of Neanderthals (Fig. 9). According to our observations, the actual tuberosity is not formed.

It is noteworthy that on the left humerus of a nine-year-old Neanderthal child from the Teshik-Tash grotto, N. A. Sinelnikov recorded a similar position of the deltoid muscle: starting on the dorsal surface, then it was located more medially [Sinelnikov and Gremyatsky, 1949, p. 130].

As you know, the entire muscle takes the arm from the trunk to a horizontal level, the front part bends the upper limb, and the back part extends it. The differences in the place of attachment of this muscle are not only formal (although this is extremely significant), they seem to reflect the different nature of daily loads. In Khvalynets, we can assume a large length of deltoid muscle fibers and its greater strength.

Computed tomography helps to correct measurements on the X-ray image (tab. 3) and evaluate the specific parameters of the internal structure (Table 4). These data convincingly demonstrate a higher compaction and internal massiveness of the Khvalyn bone in comparison with the much larger and more prominent human bone of the Bronze Age.

If we assume that postmortem surface and internal destruction of the cortical layer significantly reduced its volume, the total thickness of the cortical layer in Khvalynets will be even greater than in Neanderthals Neanderthal 1 and La Kina 5, who are characterized by a special hypertrophy of the right humerus (they had serious pathological changes affecting the left arm, resulting in the right upper limb an additional workload occurred, which caused a sharp increase in the internal massiveness of the proximal segment) [Trinkaus, Churchill, Ruff, 1994, p. 23, fig. 6].

When calculating biomechanical parameters for the middle of the humerus diaphysis, we used formulas for an ellipse. In terms of the percentage of cortical layer area and cross-section (%CA), Khvalynskaya nakhodka falls into the category of the most massive and thick-walled Neanderthal variants (Tabun 1, Le Regourdu), and taking into account the destruction of the cortical layer, it is possible that its internal massiveness exceeded these variants (Fig. 10, Table 5). Again, the author draws attention to the fact that note the sharp difference between the Khvalyn individual and the Sungir individual, which shows low rates of relative skeletal corticalization not found in the male Neanderthal population. The value of %CA in a Sungir man is lower than in a Bronze Age man.

It is noteworthy, however, that due to the extreme development of the lateral wall at the site of attachment of the deltoid muscle, the Bronze Age man also falls into the category of "Neanderthal size", approaching, for example, the European Neanderthal La Ferrasi1. Of course, the control sample chosen by us reflects one of the extreme variants of modern postcranial morphology, but this example once again proves the need to consider the complex of features in the differentiation of fossil and modern forms. However, the difference in the shape of the cross-section of the diaphysis already mentioned above remains important for distinguishing between archaic and modern variants of the structure. In addition, the more thickened Bronze Age human bone was much stronger from the biomechanical point of view, as indicated by the high value of the polar moment of inertia (Table 5).

Figure 10. Relative compaction index in the middle of the right humerus diaphysis (cortical layer area / cross-sectional area).

See Table 5. Comparative characteristics of the humerus structure from Khvalynsk in the middle of the diaphysis

Notes: 1. Comparative material - right bones of Neanderthals (Trinkaus, Churchill, Ruff, 1994; Ramsay, Weaver, and Seidler, 2005); representatives of the Bronze Age (Volga Region) and Upper Paleolithic (Sungir) (our data).

2. CA - area of the cortical layer, mm ²; MA - area of the medullary layer, mm ²; %CA - relative area of the cortical layer,%; Ip - polar moment of inertia, mm ⁴. * Calculated from data from [Trinkaus, Churchill, Ruff, 1994].

** Calculated from X-ray examination data.

Considering that the features of the Bronze Age control sample chosen by us may be too specific, we considered cross-sections of the right humerus bones of representatives of the medieval population of Yaroslavl - two women and three men.

Reference to the late sample was justified due to the external miniaturization of the diaphysis of the proximal segment of the upper limb, which brings them closer to the Khvalynskaya find. In addition, these medieval samples are characterized by moderate deltoid development.

Figure 11. Results of factor analysis based on CA, MA, %CA, and Ip characteristics.

To the individual values given in Table 5, we added data on medieval Yaroslavl residents (N 1,2-women, N 3-5-men).

bumpiness and extremely hypertrophied-large and small tubercles-a feature typical of many representatives of the Neanderthal population in both Europe and Asia. Comparison with these samples showed that the external measurement parameters of the Khvalyn bone correspond to the category of female sizes. However, the Khvalynets sharply differ from the females due to the thickening of the diaphysis in the place of the greatest development of deltoid tuberosity (70 versus 64 mm in the middle of the diaphysis), and in this indicator it is identical to the Yaroslavl males. It is noteworthy that a medieval resident of Yaroslavl (N 3) with external gracility found an increased massiveness of the walls of the humerus diaphysis, comparable to that of Neanderthals (Fig. To judge the nature of this massiveness in the Khvalynsky and Yaroslavsky individuals, reference is made to the methods of histology and paleopathology.

Results of histological examination

We conducted a histological study of the features of the tissue structure of the Khvalyn humerus. To do this, a steel hacksaw sawed out a segment of the diaphysis in the area of maximum relief development of the m. deltoideus, as well as more medial and more dorsal areas of the diaphysis. Further, grinding and polishing was performed on a Labopol-5 Straers grinding machine with a Dia Pro abrasive grinding mixture with a diamond grain size of 1 mk and OP-U polishing silicone suspension. The resulting full section was studied in reflected light using an Olympus BX-41 light microscope. The image was captured by a Color View digital camera. Osteon measurements were performed using the Olympus Cell digital image processing software. The size of the described image is 144x107 mk.

The diameters of osteons, Haversov channels, and the area of resorption cavities were measured. First of all, it should be noted that the histological structure of the cross-section at the site of the maximum development of deltoid tuberosity differs significantly from that observed in the more dorsal and medial areas without pronounced relief. Let's describe each of these pictures (Fig. 12).

1. Area of maximum development of deltoid tuberosity relief. The peripheral part of the slice. The average diameter of the Haversov canal is about 47 microns, that of the osteon is about 274 microns; in the central part, respectively, it is approx. 60 and approx. 270 mk. Resorption cavities with a diameter of 200-610 microns are found in the central part of the compacts. In the perimedullary zone, the average diameter of the Haversov channel is about 80 microns, and the resorption cavities occupy about half the area of the entire image.

2. Medial section of the bone diaphysis at the level of maximum development of deltoid tuberosity. In the peripheral part of the section, the average diameter of the Haversov channel is approx. 67 microns, and the Haversov system is approx. 255 mk; in the central region - approx. 106 and 290 mk, respectively; in the perimedullary region - 120 and 310 mk.

Thus, we observe two different trends in the dynamics of the size of Haversov systems. One of them is the enlargement of the Haversov channels from the periphery to the center of the diaphysis. It is expressed both in the medial region and partially in the place of tuberosity. Another trend is evident only in the place of development of the functional relief. Along with smaller osteons, we observe the appearance of extensive resorption cavities. Osteons with narrower Haversov channels indicate a rapid development of bone formation during the formation of deltoid tuberosity. Extensive resorption cavities cannot be the result of a general bone defect, as they are found only in the part of the bone that is most susceptible to constant mechanical stress from muscle tendons. Hypothetically, this picture can be explained by a physiological reaction to increased physical activity.

As a comparative object, a preparation was made from the humerus of an individual (N 3)

12. Histology of the cross-section of the Khvalyn bone at the level of maximum development of deltoid tuberosity.

- a peripheral part; b-medial area; c-central part. 20x magnification.

medieval series from Yaroslavl. Recall that morphologically, the width of the cortical layer in this individual reaches its maximum size. Histological cross-section revealed that the parameters of the osteon structure are extremely peculiar. The average diameter of the Haversov canal in the peripheral region is close to 30 microns, and in the perimedullary region-to 50 microns, while the average diameters of osteons are 285 and 260 microns, respectively. According to histomorphometric measurements, such osteon sizes are within the limits of the normal variability of modern human parameters, while the diameters of Haversov channels are significantly lower than normal. The causes of thickening of the inner walls of the cortical layer should be considered taking into account pathological processes.

The resulting histological picture is quite consistent with what is observed in diaphyseal hyperostoses: a biopsy shows the absence of any signs of an inflammatory or other significant pathological process, and there is only some uniform osteosclerosis with narrow Haversov channels between the compacted bone substance [Reinberg, 1964, book 1]*.

So, the conducted histological study showed that the fact of internal massiveness itself cannot be considered as a taxonomically valuable feature when describing the elements of the postcranial skeleton. An increase in the thickness of the compacts of the tubular bone diaphysis can be caused both by a pathological process and by the specifics of normal ontogenesis. When examining the humerus from Khvalynsk, we observed the second variant. The formation of large resorption cavities is caused by a morphofunctional response to extreme mechanical loads.

Discussion of the results of the humerus study

Summing up the preliminary results of the macromorphological examination of the fossil bone fragment from Khoroshevsky Island near Khvalynsk, we can identify several features that were mentioned by researchers as evidence of archaic (Neanderthal?)evolution. morphology. So, we met:

1) flatness of the diaphysis in the mediolateral direction;

2) expansion of the compact bone layer in the area of attachment of the deltoid and pectoralis major muscles;

3) thickening of the anterior and posterior walls of the diaphysis throughout compared to Cro-Magnons of Sungir;

* In this series, there are several other cases of thickening of the diaphyseal part of the tubular bones by the type of diaphyseal hyperostosis. The disease is quite rare, inherited according to the autosomal dominant type [Volkov, Meerson, Nechvolodova, 1982].

4) ellipsoid shape of the brain canal section;

5) a different localization of the deltoid tuberosity, which itself is narrow and weakly protruding above the surface of the bone body, compared to the modern one;

6) concave rather than convex shape of the bone contour below the deltoid tuberosity;

7) the extraordinary massiveness of the diaphysis walls, which is at the upper limit of individual Neanderthal values.

Based on these characteristics, the owner of the described shoulder diaphysis should be attributed to carriers of archaic postcranial morphology. However, how specific could such a combination of features be? Are they autopomorphs, i.e. exceptional features of archaic, for example Neanderthal, morphology?

Our comparison with data on anatomically modern sapiens from the Upper Paleolithic, Bronze Age, and Middle Ages has shown quite convincingly that individual characters cannot be reliable delimiters in assessing taxonomic affiliation. However, it seems that the presence of their complex serves as more reliable evidence of archaic (in this case, rather Neanderthal) morphology. It is noteworthy that the most significant differences between the Khvalyn individual and representatives of the Upper Paleolithic and Bronze Age populations were revealed. But, as noted above, one of the medieval samples used for comparison found an unexpectedly high, "archaic" indicator of humeral corticalization.

In what cases is it possible to observe the appearance of similar structural features, especially an increase in internal bone mass, in modern Homo sapiens? Finding out the cause of thickening of the inner walls of the cortical layer should begin with taking into account possible pathological processes. A number of pathological conditions show a general response to an increase in bone mass per unit volume. Radiologically, this is manifested in increased bone density*. Moreover, with a slow process of bone deposition, true lamellar bone is formed without visual defects. Most often, and this is exactly what we observe in the case of unexpectedly revealed internal massiveness in a medieval man (N 3), with osteosclerosis, the cortical layer thickens mainly due to the internal parts of the bone, which leads to narrowing of the bone-marrow space.

Often, it is genetic disorders that cause thickening of the cortical layer of tubular bones, including the humeral ones (Reinberg, 1964, book 2). Recall that, despite the superficial similarity in the relative level of compaction, the histological picture obtained for the Khvalynsky section is fundamentally different and reflects the consequences of active bone rearrangements under the influence of physical exertion.

Experts have discussed the dependence of the degree of massiveness of tubular bone walls on hereditary factors and hormonal status [Garn, Pao, and Rihl, 1964; Garn, 1970; Smith, Bloom, and Berkovitz, 1984]. In particular, Kennedy (1985) analyzed the causes of morphological changes characteristic of the erectus, in which, in addition to thickening of the cortex, there was a narrowing of the space of the brain canal (medullary stenosis). After discussing the relationship with increased compaction of strength adaptation, hereditary and humoral factors (including steroid hormones), and a lack of vitamins A and D in food, J. Kennedy considered the features of calcium metabolism in fossil hominids. As a result, when explaining the high degree of skeletal corticalization in Homo erectus, the researcher preferred the hypothesis of food specialization. New features of the diet and, above all, irregular, "sudden" food intake (cyclic hypocalcemia) could lead to a systemic reaction that reduced the possibility of bone resorption. It is characteristic that the massiveness of the bones of archaic sapiens and erectus was provided by accretion from the endoost, and not from the subperiost.

As for Neanderthal morphology, many researchers see the causes of hypertrophy of the cortical layer almost exclusively in the enormous biomechanical loads that occurred mainly on the right arm. One of the most convincing studies examined the issues of right-left shoulder bone asymmetry in Neanderthals in a comparative light (Trinkaus, Churchill, Ruff, 1994). In modern athletes with hypertrophied development of one arm (tennis players), an overestimation of the asymmetry of diaphyseal parameters was found, similar to Neanderthal.

Israeli anthropologists (Ben-Itzhak, Smith, and Bloom, 1988) conducted an X-ray examination of the humerus bones of Neanderthals and Upper Paleolithic humans (Aurignacians and Madeleines). The Neanderthal sample included La Chapelle-au-Seine, La Ferracy, San Cesare, Spi, Amud 1, Tabun O. The group of early representatives of Homo sapiens sapiens included Cro-Magnon, La Madeleine, Abri Pato, Roque de Serse, Le Plakard, Grotto of Children, Kafzeh, and Skhul. The bones were studied in the mediolateral and anteroposterior projections. To co-

* Usually, without quantitative histomorphometric data, it is impossible to distinguish between weight gain due to increased formation of new bone and reduced resorption of already formed bone.

Unfortunately, the authors of the publication did not provide the actual values of the thickness of the compact layer and the compaction indices for the men's series, limiting themselves to the graphic presentation of the material. This made it impossible for us to directly use the data they collected for comparison. When looking at radiographs in the anteroposterior projection, the diameter and cross-sectional modulus of the right humerus bones in Neanderthal males are significantly larger than in Cro-Magnon males (P < 0.05). On the left, the differences are not significant, but the Neanderthal numbers are again showing an increasing trend. Comparison of Cro-Magnons with the control sample from the Late Arabian Dor cemetery showed that most of the parameters were similar, but the anteroposterior thickness of the Upper Paleolithic compacts was greater.

After reviewing the images in the mediolateral projection, it was found that Neanderthals have a significantly higher cross-sectional modulus of the right humerus than Cro-Magnons; and this parameter is also larger and more compact in comparison with late Homo sapiens sapiens.

Area of the cortical layer (CA), moments of inertia around the mediolateral (Iml) and anteroposterior (Iap) The polar moment of inertia (Ip) on the right is significantly higher in Neanderthals than in Homo sapiens sapiens, both early and late. As a result, a significant asymmetry in the development of the humerus bones in Neanderthal males was demonstrated. The right bone, when measured externally, shows a more rounded shape than in the early or late Homo sapiens used for comparison in this study. At the same time, it is wider, the anterior and posterior walls are thicker than in Cro-Magnons, and the brain canal cavity has an ellipsoid shape. Sexual dimorphism based on the development of cortical layer thickness is high in Neanderthals and low in Cro-Magnons. According to Israeli anthropologists [Ibid., p. 240], this confirms the different activity patterns of Neanderthal and Cro-Magnon males. From the biomechanical point of view, the right shoulder of Neanderthals is reinforced in comparison with Cro-Magnons, which may mean hypertrophied development of musculature, primarily deltoid and large round muscles, latissimus dorsi, rotators (mm. supraspinatus, infraspinatus, teres minor, subscapularis).

If we talk about trends in the structure of the humerus bones in Neanderthals outside of Europe, we should mention the exceptional massiveness of the cortical layer and the insignificant width of the bone-brain space, identified by D. G. Rokhlin [1949, pp. 117-118] in a Teshik-tash child. The severity of the external muscle relief noted by the researcher suggests that such internal massiveness is due to both hereditary and biomechanical factors.

A computer tomography comparison of the Khvalyn and modern (Bronze Age) samples made it possible to identify two fundamentally different reactions to constant biomechanical stress (assuming that it was experienced by a Khvalyn person). It is possible that similar processes caused by different plasticity and degree of bone mineralization led to another, more well-known morphological discrepancy: the presence of a protruding pilaster on the femur of Homo sapiens and its absence against the background of an increase in femoral massiveness in Neanderthals.

According to extensive comparative data, similar features of the internal structure and bone relief are characteristic of all Neanderthal humerus bones. Thus, we are dealing with a complex, and not with individual features. It is obvious that one of the most important tasks is to identify the morphophysiological role of bone tissue rearrangements, which can indicate specific ways of adaptation that are peculiar only to this group of fossil people.

In general, the study of microstructural features of fossilized remains of ancient humans is not often carried out. This may be due to both the high labor intensity and the destructive nature of such studies. In the Russian tradition, methods of histology were developed at the Department of Anthropology, the Research Institute and the Museum of Anthropology of Moscow State University, for example, N. A. Sinelnikov used a microscopic method to find out the causes of the appearance of a tissue-like pattern on the surface of the skull cap from the Gangway [1952]. E. N. Khrisanfova published the results of a large-scale histological study of the femur bones of Sungir 1, 4, Romankovo and modern ones [1984]. The study of histological features of individuals from Sungir burials was continued as part of a comprehensive paleoecological project (Kozlovskaya, 2000).

As is well known, histological studies have proved extremely informative in working with objects of paleopathology [Ortner and Putschar, 1985; Schultz, 2001] and in the practice of age determination [Kerley, 1965; Ahlqvist and Damsten, 1969; Maat, Arents, Nagelkerke, 2005]. The question of the taxonomic value of histological features remains open. According to E. N. Khrisanfova, "to solve this problem, it is necessary to further study the variations of the osteonic structure in fossils and modern humans in the aspects of inter-population and intra-population (including constitutional) variability" [1984, p. 140]. Nevertheless, the striking differences in the size, structure, and density of the location of the osteonic systems of higher anthropomorphic apes and humans determine the absolute significance of the study.

M. Schultz presented convincing evidence of the similarity of the osteonic structure of australopithecines and higher anthropomorphic apes (Schultz, 1999). The structural features of the ulna of a mature Neanderthal individual and the clavicle of an older speckled individual described by him correspond to those of modern humans. At the same time, the experience of histological examination of the humerus of Le Moustier 1 demonstrated the activity of post-definitive bone rearrangements that are ahead of the age norms characteristic of modern humans [Ramsay, Weaver, Seidler, 2005].

The results obtained by us still allow us to assume the peculiarity of the physiological response of bone tissue to physical exertion. With active bone rearrangements (resorption) observed in the thickness of the compact layer, a significant bone relief is not formed, as it normally occurs in modern humans. The predominance of the osteoclastic reaction in areas of bone tissue experiencing stress mechanical loads from the periosteum can be considered as a disaptation or a specific local reaction aimed at improving blood supply to bone tissue.

Morphophysiological studies of modern natives of the North revealed a complex of adaptive responses of the skeletal system to low-temperature stress, almost carnivorous nutrition, limited sunlight, oxygen deficiency, and mineral deficiencies in the environment [Alekseeva, 1977; Alekseeva and Kovalenko, 1980; Dobrovolskaya, 1984, 2005]. T. P. Alekseeva, describing the Arctic adaptive type, formulated a hypothesis about the increase in the medullary channel of the diaphysis of long tubular bones as a way to increase the volume of red bone marrow and, accordingly, to intensify the processes of hematopoiesis.

The long tubular bone of the Neanderthal is characterized by a different internal structure: thickened walls and a small volume of the bone-marrow cavity. Thus, we observe two fundamentally different adaptive mechanisms that evolved in different members of the genus Homo. One approach (for carriers of archaic postcranial morphology) involves the accumulation of significant amounts of mineral substances to build a strong structure, possibly due to a decrease in the reserve of hematopoiesis; the other (for modern humans) involves the creation of a more robust scheme of the diaphysis structure (a"hollow" tube), which requires a much smaller amount of deficient minerals. Especially important is the question of mineral resources in connection with the development of ideas about the diet of Neanderthal man and modern Arctids. From ethnographic studies, it is known that the traditional diet of the aborigines of the North consists mainly of animal proteins and fats. It is formed both by the peculiarity of the local biota (a small variety of edible plants and a short growing season), and by significant energy costs during active activity under low-temperature stress.

Isotopic studies of bone collagen from a number of Mousterian samples allowed us to formulate a hypothesis about the extreme carnivory of Neanderthals as specialized hunters of terrestrial herbivores. It is important to note that this specialization is characteristic of the entire Mousterian region, on the basis of which a group of researchers led by E. Bosheren formulated a hypothesis about the change in the composition of the main commercial species during the Mousterian epoch from medium-sized herbivores (for example, various species of forest deer) to the late Pleistocene megafauna, in which the mammoth occupied a leading place [Bocherensetal., 2005].

Thus, under sufficiently similar stressors, carriers of archaic and modern morphology developed fundamentally different physiological adaptive responses. First of all, these differences are related to two different ways of building mineral metabolism. A more archaic approach involves a constant intensive accumulation of minerals and protein masses (collagen, the structural protein of bone). This mechanism is possible with more active absorption of fat-soluble vitamin D, as well as activation of the thyroid gland (calcitenin secretion), liver and suppression of synthesis or sensitivity to parathyroid hormone of the parathyroid glands.

Vitamin D performs two well-studied functions in relation to the skeletal salt metabolism that interests us. First, it promotes the absorption of calcium from food masses in the wall of the small intestine, promoting the entry of salt into the circulating blood; secondly, it helps the deposition of calcium in the bone tissue itself. A lack of vitamin D in the body, as well as its excess, can seriously disrupt the normal processes of bone formation and cause ossification pathology [Reinberg 1964, book 2].

Calcium metabolism is also directly controlled by parathyroid secretions. If vitamin D promotes the absorption of calcium salts by bone tissue, then the parathyroid hormone promotes the release of calcium from the bones-

structures. Therefore, it is an important factor in maintaining normal blood calcium levels. Endocrine disorders of various etiologies lead to fibrotic osteitis, which is also due to increased bone density, for example, in active hyperparathyroidism. A similar pattern of increased density is also observed in hypothyroidism against the background of a general decrease in the rate of bone formation and resorption, when the balance shifts towards bone formation, a denser bone appears that retains its normal structure [Ibid.].

The activity of glucocorticoids also contributes to a decrease in mineral deposition, so we can assume that paleoanthropes either have a reduced sensitivity to this group of hormones, or a reduced secretion of them. Meanwhile, representatives of Neanderthals demonstrate a clearly "androgenic" constitution, as Professor E. N. Khrisanfova has repeatedly written about [1971, 1974, 1984, 1990]. Increased androgen secretion contributes to the general intensification of anabolic processes, including in the construction of skeletal tissues. In the physiology of modern humans, it stimulates an increase in the secretion of glucocorticoids, in particular cortisol.

Conclusion

Thus, there are grounds to assume that the group of paleoanthropes to which the Khvalyn individual belonged had not only peculiar biomechanical adaptations, but also a specific hormonal status that does not find complete analogues in the normal and pathological physiology of modern humans.

Despite the external miniaturization of the examined bone, it is rather masculine. Indirect morphological criteria are used as a basis: internal massiveness at the upper boundary of individual male values in representatives of the genus Homo (including Neanderthals), as well as a sharp difference between the minimum circumference of the diaphysis and the circumference at the place of maximum development of deltoid tuberosity. These features, along with the histologically revealed pattern of active bone rearrangements, indicate an extreme biomechanical effect on the right arm, which, as is known, was highly characteristic of Neanderthal males.

The age of an individual in the most preliminary form can be estimated as mature. The histological picture is so specific that it may not meet modern standards. A similar problem was faced by researchers of the Neanderthal Le Moustier 1, whose " histological "age (40-50 years according to the sample taken from the humerus) sharply contrasts with the" dental "(15 years) and" skeletal " (9 years) [The Neandertal..., 2005; Mednikova, 2007].

Taking into account the paleontological context of the humerus from Khoroshevsky Island and the "atypical" Neanderthaloid features of the accompanying cranial cap, we can, unlike our predecessors, assume an early rather than late age of these fossils. So far, in the absence of more precise references, the fragment we examined can most likely be attributed to an early Paleoanthropus of the Late Ashelian or Early Mousterian epoch.

Acknowledgements

The authors express their deep gratitude to the staff of the Radiation Diagnostics Department of the N. I. Priorov Central Institute of Traumatology and Orthopedics: Head of the department Dr. med. A. K. Morozov, Candidate of Medical Sciences I. N. Karpov, radiologist M. Ya. Polonskaya, radiologist E. A. Gusiletova.

List of literature

Alekseeva T. N. Geographical environment and human biology. Moscow: Mysl Publ., 1977, 302 p. (in Russian)

Alekseeva T. I., Kovalenko V. Yu. Morphofunctional characteristics of the postcranial skeleton of Asian Eskimos / / Paleoanthropology of the USSR / ed. by A. P. Okladnikov, V. P. Alekseev. - Moscow: Nauka, 1980. - pp. 131-153.

Volkov M. V., Meerson E. M., Nechvolodova O. L. Hereditary systemic diseases of the skeleton. - Moscow: Medicine, 1982. - 320 p.

Dobrovolskaya M. V. Experience the quantitative determination of trace elements in the human skeleton (on the materials of the ancient burials) // Vopr. anthropology. - 1984. - Vol. 74. - p. 101-109.

Dobrovolskaya M. V. Chelovek i ego pishchiya [Man and his food], Moscow: Nauch. mir Publ., 2005, 368 p.

Kandinov M. N., Mednikova M. B., Dobrovolskaya M. V., Buzhilova A. P. Humerus of Homo from Khoroshevsky Island: history and paleontological context of the discovery. -2008. - N 4. - p. 145-150.

Kozlovskaya M. V. Femoral bone Sungir 4. Histological features of the femoral bone Sungir 4 / / Homo sungirensis: Upper Paleolithic man: ecological and evolutionary aspects of research / ed. by T. I. Alekseeva, I. O. Bader. - Moscow: Nauch. mir, 2000. - pp. 325-326.

Mednikova M. B. Metodika roentgenomorphologicheskogo issledovaniya // Homo sungirensis: Verkhnepaleoliticheskiy chelovek: ekologicheskie i evolyutsionnye aspekty issledovaniya [Methods of X - ray morphological research].

Mednikova M. B. Roentgenostructural features of the skeleton of Sungir 1 / / Homo sungirensis: Upper Paleolithic-

chesky chelovek: ekologicheskie i evolyutsionnye aspekty issledovaniya [The Russian man: ecological and evolutionary aspects of research]. mir, 2000b., pp. 218-222.

Mednikova M. B. Roentgenomorphology of children from burial 2 / / Homo sungirensis: Upper Paleolithic man: ecological and evolutionary aspects of research / ed. by T. I. Alekseeva, N. O. Bader, Moscow: Nauch. mir, 2000, pp. 286-299.

Mednikova M. B. Trepanations in the ancient peoples of Eurasia. Moscow: Nauch. mir Publ., 2001, 304 p. (in Russian)

Mednikova MB. K voprosu ob osobennostei molodezheskoi etape ontogeneza u evropeyskikh neanderthaltsev [On the peculiarities of the youthful stage of ontogenesis in European Neanderthals]. - 2007. - N3. - p. 145-153.

Reinberg S. A. Roentgenodiagnostics of diseases of bones and joints. - 4th ed. - M.; L.: Medgiz, 1964. - Kn. 1. -532 p.; Kn. 2. - 572 p.

Rokhlin D. G. Nekotorye dannye roentgenologicheskogo issledovaniya detskogo skeleta iz grota Teshik Tash, Yuzhny Uzbekistan [Some data from the X-ray study of a child's skeleton from the Teshik Tash grotto, Southern Uzbekistan].

Sinel'nikov, N. A., On the formation of a tissue-like relief on the Skhodnensky skull fragment, Uchen. zap. Mosk. un-ta. - 1952. - Issue 158: Fossil man and his culture on the territory of the USSR. - P. 175-179.

Sinelnikov N. A., Gremyatsky M. A. Bones of the skeleton of a Neanderthal child from the Teshik-Tash grotto, Southern Uzbekistan / / Teshik Tash: Paleolithic man / ed. by M. A. Gremyatsky, M. F. Nesturkh, Moscow: Publishing House of Moscow State University, 1949, pp. 123-136.

Khrisanfova E. N. Some aspects of hormonal research in anthropology. P. The importance of studying steroid hormones in functional and evolutionary anthropology // Questions of anthropology. - 1971. - Issue 38. - p. 3-14.

Khrisanfova E. N. Adaptation of the skeleton of early hominids in connection with some environmental factors // Primeval man and the natural environment / ed. by G. I. Lazukov, Moscow: Mysl, 1974, pp. 92-97.

Khrisanfova E. N. Postcranial skeleton of an adult male Sungir-1 / / Sungir: Antropologicheskoe issledovanie / otv. red.AA. Zubov, Moscow: Nauka Publ., 1984, pp. 100-140.

Hrisanfova E. N. Kontsutsiya i biokhimicheskaya individualnost ' cheloveka [Constitution and biochemical individuality of a person], Moscow: Publishing House of Moscow State University, 1990, 190 p. (in Russ.)

Ahlqvist J., Damsten O. Amodification of Kerley's method for the microscopic determination of age in human bone // J. of Forensic Science. - 1969. -Vol. 14. -P. 205 - 212.

Aiello L., Dean C. An introduction to human evolutionary anatomy. -Amsterdam: Elsevier Ptd., 2006. - 596 p.

Ben-Itzhak S., Smith P., Bloom R.A. Radiographic study of the humerus in Neanderthals and Homo sapiens sapiens // Am. J. of Physical Anthropology. - 1988. -Vol. 77. -P. 231 - 242.

Bocherens H., Drucker D.G., Billion D., Pathou-Mathis M., Vandermeersch B. Isotopic evidence for diet and subsistence pattern of the Saint-Cesaire I Neanderthal: review and use of a multi-source mixing model // J. of Human Evolution. - 2005. - Vol. 49. - P. 71 - 87.

Endo B., Kimura T. Postcranial skeleton of the Amud Man // The Amud Man and his cave site / eds. H. Suzuki, F. Sakai. - Tokyo: Academic Press, 1970. - P. 231 - 406.

Garn S.M. The earlier gain and the later loss of cortical bone in nutritional perspective. - Springfield: Thomas, 1970. - 502 p.

Garn S.M., Pao E., Rihl M. Compact bone in Chinese and Japanese // Science. - 1964. - Vol. 143. - P. 1439 - 1440.

Hambucken A. Revision des particularites de l'humerus des Neandertaliens europeens // Comptes Rendus de l'Acadernie des Sciences de Paris. Ser. II. - 1983. - Vol. 317. - P. 109 - 114.

Heim J. -L. Pes variations du squelette post-cranien: Des hommes de Neandertal suivant de sexe // Anthropologic J. -1983. - Vol. 87. - P. 379 - 416.

Kennedy J.L. Bone thickness in Homo erectus // J. of Human Evolution. - 1985. - Vol. 14. - P. 699 - 708.

Kerley E.R. The microscopic determination of age in human bone // Am. J. of Physical Anthropology. - 1965. -Vol. 23. - P. 149 - 164.

Maat G.J.R., Aarents M.J., Nagelkerke N.J.D. Age prediction from bone replacement. Remodeling of circumferential lamellar bone tissue in the anterior cortex of the femoral shaft of present Dutch population. - Peiden: Peiden University, 2005. - 67 p. (Barge's Anthropologica; N 10).

Ortner D. J., Putschar W.G.J. Identification of pathological conditions in human skeletal remains. - Washington: Smithsonian Institution Press, 1985. - 479 p.

Pearson O.M. Postcranial morphology and the origin of modern humans: Ph.D. thesis. - N. Y.: State University of New York at Stony Brook, 1997. - 783 p.

Ramsay H.L., Weaver D.S., Seidler H. Bone histology in the Pe Moustier Neandertal child // The Neandertal Adolescent Pe Moustier 1: New Aspects, New Results / ed. by H. Ullrich. -Berlin: Allprint Media GMBH, 2005. - P. 282 - 292. - (Berliner Beitrage zur Vor- und Fruhgeschichte. N.F.; Bd. 12).

Schultz M. Microscopic investigation in Fossil Homino-idea: a clue to taxonomy? Functional anatomy and history of diseases // Phe anatomical record (New Anat). - 1999. -Vol. 257. - P. 225 - 232.

Schultz M. Paleohistopathology of bone: a new approach to the study of ancient diseases // Am. J. of Physical Anthropology. - 2001. - Vol. 44. - P. 106 - 147.

Smith P., Bloom R.A., Berkovitz J. Diachronic trends in humeral cortical thickness of Near Eastern populations // J. of Human Evolution. - 1984. -Vol. 13. -P. 603 - 611.

Stirland A. Musculoskeletal evidence for activity: problems of evaluation // Int. J. of Osteoarchaeology. - 1998. - Vol. 8. -356 p.

The Neandertal Adolescent Le Moustier 1: New Aspects, New Results / ed. by H. Ullrich. - Berlin: Allprint Media GMBH, 2005. - 354 p. - (Berliner Beitrage zur Vor- und Fruhgeschichte. N.F.; Bd. 12).

Trinkaus E. The Shanidar Neanderthals. - N. Y.: Academic Press, 1983. - 502 p.

Trinkaus E., Churchill S.E., Ruff C.B. Postcranial robusticity in Homo. IP Humeral bilateral asymmetry and bone plasticity // Am. J. of Physical Anthropology. - 1994. -Vol. 93. -P. 1 - 34.

The article was submitted to the Editorial Board on 19.05.08.

Permanent link to this publication: