2 Osteoarchaeology

1.1 BONE FUNCTION

The human skeleton performs multiple functions. It supports the body and enables motion, while certain skeletal elements
protect internal vital organs; for instance, cranial bones offer protection to the brain. In addition, the skeleton stores: (1)
minerals, such as calcium and phosphorus; (2) blood-forming cells, as the red bone marrow produces red blood cells, white
blood cells, and platelets via a process known as hematopoiesis; and (3) energy via the fat (lipids) stored in the yellow bone
marrow.

1.2 BONE CLASSIFICATION

The adult skeleton (Fig. 1.2.1) typically consists of 206 bones, though this number may differ slightly among individuals
owing to genetic factors. At birth, the skeleton has many more bones (over 270) but during adolescence many separate
elements gradually fuse together.

FIGURE 1.2.1 Three-dimensional model of the human skeleton. Note: The hyoid is not visible in this figure; the corresponding arrow denotes its
location.

Based on their shape, human bones are divided into long, short, flat, and irregular bones. Long bones are elongated
elements, as their name suggests, and consist of a tubular shaft terminating in an articular area at each end. Such bones are
found in the upper and lower limbs (clavicle, humerus, radius, ulnaefemur, tibia, fibula). Their main functions include
supporting the weight of the body and enabling movement. Note that some authors place the metacarpals and metatarsals
also in the long bone category on the grounds that their length is greater than their width. In short bones all dimensions are
almost equal and their somewhat cubic shape provides them with compactness. Such bones are found mostly in the hands
and feet (carpals, tarsals). Flat bones are flat, thin, and broad elements. As such, they provide extensive areas for muscle
attachment and protect vital organs. This category includes most cranial bones (e.g., frontal, occipital, parietals), as well as
the scapulae, os coxae, ribs, and sternum. Finally, as their name suggests, irregular bones have a complex morphology,
which serves various functions. The elements of the spine (vertebrae, sacrum, coccyx) and certain cranial elements
(e.g., sphenoid) belong in this category.
According to their location in the skeleton, bones are classified into axial and appendicular. The axial skeleton has 80
bones and includes the skull, hyoid, vertebrae, sacrum, coccyx, ribs, and sternum. The appendicular skeleton consists of
126 bones, which are the long bones of the arms and legs, patellae, hand and foot bones, clavicles, scapulae, and os coxae
(Fig. 1.2.2).
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FIGURE 1.2.2 Axial and appendicular skeleton.

Two special categories of bones, which differ substantially in number between individuals, are sesamoid bones and
sutural bones. Sesamoid bones are very small-sized and found within tendons. Their function is to protect tendons from
excessive mechanical stress. Sutural bones are small-sized and found within the cranial sutures, that is, the joints that
connect cranial bones.
Details on how to identify each bone and tooth are given in the following sections. Appendix 1.I presents a recording
spreadsheet for the adult human skeleton and Appendix 1.II provides a recording spreadsheet for the permanent dentition.
Finally, Appendix 1.III presents the bone and tooth inventories implemented in Osteoware, a free software program for the
standardized recording of human skeletal remains, designed and provided by the Smithsonian Institution.

1.3 BONE STRUCTURE

1.3.1 Gross Anatomy
A typical long bone consists of (1) a diaphysis, the shaft of the bone; (2) two epiphyses, the extremities of the bone; and (3)
two metaphyses, the areas that lie between the diaphysis and each of the epiphyses (Fig. 1.3.1). The epiphyses are initially

FIGURE 1.3.1 Long bone structure.

4 Osteoarchaeology

separated from the diaphysis, as the metaphyses contain a growth plate (or epiphyseal plate), consisting of hyaline
cartilage, which allows longitudinal bone growth. Once an individual reaches adulthood and growth ceases, this cartilage is
ossified and forms the epiphyseal line. The surface of the epiphyses that articulates with neighboring skeletal elements is
covered with a thin layer of hyaline cartilage (articular cartilage) during life. The external bone surfaces that have no
articular cartilage are covered by the periosteum. The periosteum enables bone growth in thickness, protects and nourishes
the osseous tissues, facilitates fracture repair, and allows the attachment of ligaments and tendons. It attaches to the un-
derlying osseous tissues by means of thick collagen fibers, called Sharpey’s fibers. In the interior of the long bones lies the
medullary cavity, which, during life, contains yellow bone marrow. The walls of this cavity are covered by the endosteum,
a membrane that contains bone-forming cells.
Short, flat, and irregular bones have a rather simpler structure, consisting of trabecular bone covered by cortical bone (see Section
1.3.2 for definitions of the trabecular and cortical bone). The gross structure of a flat bone is shown schematically in Fig. 1.3.2.

FIGURE 1.3.2 Flat bone structure.

1.3.2 Microscopic Anatomy

Histologically, bone tissue may be woven (immature) or lamellar (mature). During periods of skeletal growth or when
certain pathological conditions have afflicted the skeleton, such as during fracture healing, the bone tissue produced
consists of irregularly shaped collagen bundles with a random orientation. This type of bone is called woven or primary
bone. Woven bone provides a means of responding rapidly to growth, disease, or mechanical loading; however, its
disorganized structure makes it ineffective in offering long-term structural support. In contrast, mature or lamellar bone is
formed in later childhood and during adulthood, and it consists of mineral crystals and collagen fibers that form layers
called lamellae. Lamellar bone gradually replaces woven bone.
Lamellar bone has two structural types, cortical (or compact) bone and trabecular (or cancellous) bone. Cortical bone is
dense and lies underneath the periosteum. Note that the cortical bone found in the joints is covered by cartilage during life
and it is called subchondral bone. Its structural units are osteons, which consist of concentric layers of compact bone,
lamellae, organized around a central canal, a Haversian canal, containing blood vessels and nerve fibers. Trabecular bone
is found at the epiphyses of long bones and in the interior of all other bones. It consists of thin bony spicules, trabeculae,
each of which is composed of a few layers of lamellae. The trabeculae are arranged irregularly, forming a honeycomb
structure. Because of this structure, trabecular bone has minimal weight but at the same time provides great strength to the
skeleton. During life, the spaces within the trabecular network contain hematopoietic red marrow.

1.4 BONE COMPOSITION AND CELLS

Bone tissue consists of inorganic (w70%) and organic (w30%) components. The principal inorganic component is hy-
droxyapatite, Ca10(PO4)6(OH)2, a mineral composed of calcium phosphate. Collagen fibers are the main organic
component, along with noncollagenous proteins. The inorganic component makes the skeleton strong, whereas collagen
offers elasticity.
As a living tissue, the human skeleton goes through a constant process of bone resorption (releasing of calcium and
phosphate from mineralized bone) and deposition (use of calcium and phosphate to form new bone). This process is
controlled by the main bone cells: osteogenic cells, osteoclasts, osteoblasts, osteocytes, and bone-lining cells. Osteo-
genic (or osteoprogenitor) cells derive from mesenchymal stem cells in bone marrow. These cells evolve into pre-
osteoblasts, which subsequently give rise to mature osteoblasts. In the marrow of growing individuals, the number of
osteogenic cells is high, but the number or potential of such cells to form mature osteoblasts declines with age. Os-
teoclasts are large, multinucleated cells, whose primary function is to resorb bone tissue by releasing lysosomal
 The Human Skeleton Chapter | 1 5

enzymes and acids. Osteoblasts facilitate new bone formation by secreting organic bone matrix, osteoid, and regulating
its mineralization. They are mostly found under the periosteum and near the medullary cavity, where metabolic bone
rates are higher. Once an osteoblast has completed bone formation it may (1) turn into a bone-lining cell (discussed
later), (2) sustain apoptosis (programmed cell death), or (3) transform into an osteocyte. Osteocytes comprise 90% of all
cells in mature bone and derive from osteoblasts that have become trapped in newly formed bone tissue. They facilitate
cellular communication and maintain the daily functions of the skeleton by transporting nutrients and wastes. Finally,
bone-lining cells derive from osteoblasts and cover bone surfaces. Their function is to release calcium when required,
participate in the initiation of bone resorption and remodeling, and possibly maintain bone fluid balance.

1.5 BONE GROWTH AND DEVELOPMENT

Bone formation (osteogenesis) may occur through ossification within a connective tissue membrane (intramembranous
ossification) or through ossification of cartilage precursors (endochondral ossification). Intramembranous ossification
gives rise to the cranial vault and face, as well as partly to the clavicle and scapula. In contrast, long-bone epiphyses,
short bones, vertebral bodies, and other elements largely consisting of trabecular bone grow by endochondral
ossification.
Most bones are formed from at least two centers of ossification. The first center that appears is called the primary
ossification center; its ossification usually begins in utero, and in long bones it corresponds to the diaphysis. Most sec-
ondary ossification centers appear after birth, and in long bones they correspond to the epiphyses. As mentioned in Section
1.3.1, between the diaphysis and the epiphyses lies a cartilaginous layer, the growth plate, which allows the bones to grow
in length. During adolescence and early adulthood, the primary and secondary ossification centers fuse, giving rise to the
complete bones.
As a bone develops, its form gradually alters to assume the final adult shape. This process is known as modeling. In
contrast, remodeling is the replacement of mature bone during the repair of bone microdamage or as part of bone
adaptation to mechanical loading (see Chapter 7).

1.6 PLANES OF REFERENCE AND DIRECTIONAL TERMS

This section presents brief definitions of the planes of reference and directional terms that are essential in the description of the
human skeleton. All definitions provided here assume that the human skeleton is in standard anatomical position, that is, standing
erect, looking forward, with the feet close and parallel to each other, the arms at the sides, and the palms facing forward (Fig. 1.6.1).

FIGURE 1.6.1 Planes of reference and directional terms for the human skeleton.
6 Osteoarchaeology

The main planes of reference for the human skeleton are used to divide the body into sections (Fig. 1.6.1). The sagittal
(or midsagittal) plane separates the right half from the left half of the body, whereas the coronal plane is perpendicular to
the sagittal and separates the anterior half from the posterior half of the body. Finally, the transverse plane is perpendicular
to the sagittal and coronal planes and it may be located at different heights.
The main directions for parts of the body are superior, inferior, anterior, posterior, medial, and lateral, whereas the
terms proximal and distal are more appropriate for the limbs (Figs. 1.6.1 and 1.6.2). Superior is toward the head,
inferior toward the feet, anterior toward the front of the body, posterior toward the back of the body, medial toward the
sagittal plane, and lateral away from the sagittal plane. For the limbs, proximal lies toward the trunk of the body, and
distal lies away from the trunk. Terms that are often used for the hands and feet include palmar, which is the palm side
of the hand; plantar, which is the sole side of the foot; and dorsal, that is, the top side of the foot or the back side of the
hand. Note that when the terms right and left are used, they refer to the sides of the individual being studied and not to
the sides of the observer.

FIGURE 1.6.2 Directional terms for the upper and lower limbs.

1.7 BONE IDENTIFICATION

The identification of individual human skeletal remains may appear to be a daunting task; however, with experience it
becomes straightforward. When the skeleton is well preserved, the figures provided in this section should offer sufficient
information to allow the identification of each bone. In cases of partial preservation, the process of elimination should be
followed. In this process, it is imperative to first determine if the elements under examination belong to an adult or a
juvenile. Subsequently, we assess if the bone is long, short, flat, or irregular; though note that juvenile bones may be
difficult to classify into these categories at certain stages of their development. Finally, we contrast the morphology of the
bone under examination with that of the elements depicted in the following figures and find the most likely match. In the
case of juvenile remains, we perform the earlier steps using as a guide one of the textbooks provided in the suggested
reading list.

1.7.1 Axial Skeleton

1.7.1.1 Skull
At birth, the human skull consists of 45 elements but many of these gradually fuse, resulting in 28 bones in the adult skull.
Fig. 1.7.1 shows the bones that are visible ectocranially on the lateral view of the adult skull. The bones of the skull may be
divided into cranial bones and facial bones, though further divisions may be used. Cranial bones include the frontal,
occipital, sphenoid, and ethmoid, as well as the right and left parietals and temporals. Facial bones include the right and
left nasals, maxillae, zygomatics, lacrimals, palatines, and inferior nasal conchae, as well as the mandible and vomer. Six
auditory ossicles (three on each side of the skull) form a separate category of skull bones.