Skeletal Variation among Mesolithic People of the

Ganga Plains: New Evidence of Habitual Activity

and Adaptation to Climate

JOHN R. LUKACS AND J. N. PAL

THE PREHISTORIC CULTURAL PERIOD KNOWN AS MESOLITHIC initially re-

ferred to a hiatus in the record of human technological achievement, separating
Paleolithic from Neolithic cultures in Europe. By the 1880s, as this cultural
discontinuity was slowly filling with evidence of miniature stone tools, the term
microlithic was proposed for this period of cultural decline following the Upper
Paleolithic florescence in technology, painting, and sculpture (Kennedy 2000).
Today, while some scholars question the utility of broad, culture-phase nomen-
clature, the label Mesolithic typically refers to cultures characterized by several
key technological and adaptive characteristics, including: (1) miniature stone tools
(microliths), (2) increased reliance on composite tools (sickles, bow and arrow),
(3) increased subsistence diversity, including reliance on plants, fish, and birds, (4)
larger and more sedentary settlements, and (5) enhanced regional variation and
cultural diversification.
In India, Mesolithic cultures exhibit high regional diversity as one aspect of
successful adaptations to such ecologically distinctive habitats as the arid zone of
the Thar Desert, the humid tropics of Sri Lanka, and the high rainfall, wooded
regions of eastern India. V. N. Misra (1996) notes that Mesolithic sites in India
are larger, better preserved, and more numerous than their Paleolithic ante-
cedents, in part because they are more recent in time, but also because Holocene
geomorphic disturbances have been minimal. Knowledge of Mesolithic cultural
adaptations in India is dramatically enhanced over preceding cultural phases by
the presence-for the first time in Indian prehistory-of an abundant and infor-
mative series of human skeletal remains.
The functional analysis of artifacts (Petraglia et al. 1999), spatial interpretation
of bone and stone scatters (Potts 1988), and taphonomic assessment of faunal
remains (Erlandson and Moss 2001) provide valuable insights into the activities
and life-ways of prehistoric people. However, when a site also yields the skeletal

John R. Lukacs is in the Department of Anthropology, University of Oregon, Eugene, and). N. Pal
is in the Department of Ancient History, Culture, and Archaeology, University of Allahabad, India .
.·-lsi.lIl P('/~~pf'{til'f·s. Vol. -12. No.2 \r\ 1003 by the Ullivl.-·rsity or Haw;li'j Press.
330 ASIAN PERSPECTIVES . 42(2) . FALL 2003

remams of its human occupants, an additional highly informative and unique

source of data becomes available for reconstructing past life-ways. For example,
demographic data on the age and sex structure of mortuary samples may be asso-
ciated with population density, disease load, and level of hygiene (Storey 1992),
physical attributes of individual specimens, such as stature and skeletal robusticity,
may provide anatomical clues regarding growth rates, mobility, and adaptation to
climatic extremes (Pearson 2000), and skeletal and dental evidence of disease pro-
vides insight into the prevalence of growth disruption, infectious disease, and oral
health conditions (Hillson 2000, 2001; Roberts and Manchester 1995).
In keeping with the theme of this volume-rethinking South Asian archaeology
-this contribution adopts an analytic paradigm known as bioarchaeology, and
uses it to better understand the activities and adaptations of the Mesolithic inhab-
itants of North India. Bioarchaeology is the analysis of human skeletons from past
populations within an interpretive framework that includes attention to both the
cultural and the geophysical context of those remains. An abridged summary of
bioarchaeological method and theory by Larsen includes a synthesis of primary
research accomplishments of this dynamic approach to human osteology (Larsen
1997). Typically the focus of bioarchaeological research is wide ranging and may
include analysis of mortuary sites (Gamble et a1. 2001; Robb et a1. 2001), gender
differences in dietary patterns (Lukacs 1996), variation in health status across social
groups (Walker and Hewlett 1990; Walker et a1. 1998) or subsistence transitions
(Lukacs and Walimbe 1998; Lukacs et a1. 2001). This study examines variation
in the marks that muscular attachments leave on bone (musculoskeletal stress
markers), arthritic degeneration of articular surfaces of bone (osteoarthritis), the
length of limb bones and their relative proportions to one another, and variation
in stature reconstructed from long bone lengths. Detailed analysis of human skel-
etal variations permits a clearer understanding of how Mesolithic people bio-
logically adapted to early and middle Holocene environments of the Ganga Plain.
While the human remains from Damdama comprise the primary focus of this
study, previously described skeletal series from the neighboring Mesolithic sites of
Sarai Nahar Rai (Kennedy et a1. 1986), Mahadaha (Kennedy et a1. 1992), and Lekha-
hia (Lukacs and V. D. Misra 1997,2002), serve as valuable comparative samples.
This contribution addresses several important questions about conditions of
existence during the Mesolithic. How physically "stressful" were the daily lives
of these people? Do skeletal variations among the people of Damdama reveal
clues about the nature of repetitive or forceful activities? How tall were these
foragers and how do they compare in stature with neighboring Mesolithic groups
at Mahadaha and Sarai Nahar Rai? Can the limb proportions of these people be
determined and interpreted in an adaptive evolutionary framework? Are their
skeletons robust or gracile in structure and is evidence of biological adaptation
to prevailing climate discernable? The answers to these questions will contribute
new perspectives on the past and lead to a clearer picture of Mesolithic life-ways
in North India. The rich and informative perspective ofbioarchaeology cannot be
derived from artifactual, taphonomic, or stratigraphic data alone, but must rely on
biological evidence obtained directly froni the human skeleton. Consequently, in
this approach to the past, human skeletal observations serve as an indispensable
and valuable complement to more traditional archaeological perspectives.
LUKACS AND PAL . MESOLITHIC SKELETAL VARIATION OF THE GANGA PLAINS 33 I

ARCHAEOLOGY AND CHRONOLOGY

In 1972, the University of Allahabad began excavations at the Mesolithic site of

Sarai Nahal' Rai under the direction of G. R. Sharma (1973). Subsequently, two
additional sites with human burials were discovered: Mahadaha, excavated in the
late 1970s (Pal 1985, 1988; Sharma et a1. 1980), and Damdama, excavated in the
mid-1980s (Varma et a1. 1985). All three sites are located between 40 and 80 km
north of Allahabad (Fig. 1), preserve evidence of human occupation adjacent to
oxbow lakes, and have produced the largest collection of Mesolithic human skel-
etons in Asia. Excavations document hearths and food refuse, microlithic and
bone tools, fauna and flora, as well as abundant human burials with simple jewelry
of bone and antler. Colloquium 33 of the 13th International Congress of Prehis-
toric and Proto historic Sciences is devoted to the bioarchaeology of Mesolithic
India (Afanas'ev et a1. 1996), and includes new data on plant and animal resources
at Damdama (Kajale 1996; Thomas et a1. 1996), lithic use-wear analysis (Pal
1996), a review of burial practices (Pandey 1996), rock art evidence of subsistence
practices (Varma 1996), as well as historical and synthetic perspectives (V. D.
Misra 1996; V. N. Misra 1996). In addition, the skeletal adaptations of human
remains from Mahadaha and Sarai Nahal' Rai are compared (Kennedy 1996) and
new AMS dates and carbon isotope values are reported for Dam.dama and Lekha-
hia (Lukacs et a1. 1996).
Burials at Damdama are typically shallow oblong pits, in which the corpse was
placed in an extended supine position, often oriented east-west. Position of the
corpse is variable, with a range of flexion observed at the knee and elbow, and
with some individuals buried in the prone position. Grave goods are rare and
include simple jewelry, animal bones, and artifacts. Six multiple burials were
excavated at Damdama, five double burials (Damdama 6, 16, 20, 30, and 36), and
one triple burial (Damdama 18) (Pal 1985, 1988, 1992). An innovative perspec-
tive on Mesolithic mortuary practices in the Ganga Plain provides insights into
issues regarding settlement pattern, social organization, and territoriality (Chatto-
padhyaya and Chattopadhyaya 1990).
The chronological placement of these sites is problematic. Some scholars are
highly critical of the early Holocene date from Sarai Nahal' Rai (10,050 ± 110
B.P.), which was favored by G. R. Sharma (1973; Sharma and Sharma 1987).
Charcoal is rare or absent, requiring the analysis of less traditional materials for
dating purposes (bioapatite vs. collagen; shell vs. charcoal) consequently giving
rise to questions of suitability, contamination, and methodology, which have
resulted in a controversial chronological framework. Earlier radiocarbon dates
are variable, with Sarai Nahal' Rai and Mahadaha yielding some early and some
late Holocene results (Possehl and Rissman 1992). Since 1992, several new 14C
dates have been published that suggest a mid-Holocene antiquity for Mahadaha
(6320 ± 80 B.P.; OxA-1647; Chattopadhyaya 1996) and a mid- to early Hol-
ocene age for Damdama (Lukacs et a1. 1996). Two AMS dates derived from
human bone samples from Stratum 1 (earliest) and Stratum 6 (middle) at Dam-
dama have yielded dates of 8865 and 8640 ± 65 B.P., respectively (Lukacs et a1.
1996). By contrast, three dates derived fi:om bioapatite of bovid enamel are re-
ported here for the first time and suggest a younger age, between 5550 ± 60 B.P.
 332 ASIAN PERSPECTIVES . 42(2) . FALL 2003

.r·........
"'-.·....--::----··5(.-:· Faizabad
Allahabal ........ ~',,-,
~ Archeological siles
ENLARGED
AREA
\.
.
N
• Modern cities. towns•
and villages

1 j I
0 50KM

Kilometers
~ Damdama
I
o

,L----..:::.~-----,,----------__IH26°N

Sai River
• Jaunpur

---------+-------------+-125°N

Fig. 1. Location map showing the physical setting of Mesolithic sites discussed in the text in relation
to the Ganga-Jumna River confluence and the modern cities of Allahabad and Varanasi (Benares).

and 5250 ± 70 B.P. (Table 1). While enamel is less subject to postmortem dia-
genesis, the earlier dates are derived directly from the human skeletons. Resolu-
tion of this discrepancy in radiocarbon dates is currently underway by dating
enamel samples from human remains, bone samples from bovids, as well as
chemical characterization of the sedimentary matrix.
LUKACS AND PAL . MESOLITHIC SKELETAL VARIATION OF THE GANGA PLAINS 333

TABLE 1. AMS RADIOCARBON DATES FROM BOVID ENAMEL (BIOAPATITE)

GEOCHRON ID
NUMBER DATE
SITE (SAMPLE #) STIlATUM SQUARE EXCAVATED BONE # AGE (B.p.) b 13 C %0

DDM GX-22887-AMS (1) 1 S-15 1987 378 5550 ± 60 0.5

DDM GX-22888-AMS (3) 6 N-4 1984 5250 ± 70 0.9
DDM GX-22889-AMS (6) 8 P-10 1984 126 5430 ± 60 1.0

Note: Analysis conducted at Geochron Laboratories, Cambridge, MA; for AMS dates from human
bone refer to Lubcs et al. 1996.

The remains of 47 adults are preserved at Damdama. This series is especially

valuable because it (1) constitutes the largest group of Mesolithic skeletons from
one site in South Asia, (2) comprises more than 50 percent of the North Indian
skeletal sample, (3) permits the first statistical assessment of human biological
adaptations for Mesolithic people of South Asia, and (4) yields new data that
relate to several currently contentious theoretical issues in biological anthropol-
ogy. This paper reports new data regarding the degree of musculoskeletal stress
and extent of osteoarthritis in postcranial skeletons from Damdama, as well as
variation in adult stature, and limb length and proportions. These data are used to
infer habitual activity patterns and biological adaptations to the environment.

RESEARCH CONTEXT: PREVIOUS BIOARCHAEOLOGY

Bioarchaeological analyses of human remains fron. Lekhahia and Sarai Nahar Rai
were initiated by Lukacs (1977). Subsequent comprehensive studies of series from
Mahadaha and Sarai Nahar Rai in the 1980s, revealed a tall, large-framed people,
who possessed large teeth and jaws, yet had few pathological lesions (Kennedy
et al. 1986, 1992). The dental anthropology of Damdama was preliminarily
described by Lukacs and Pal (1992, 1993). Large tooth size, low frequency of
dental abscesses, low dental caries rates, low antemortem tooth loss, and high
levels of dental attrition typify the people of Damdama, and are traits consistent
with a coarse diet and a hunting and foraging subsistence base. In dental size,
morphology, and pathology, the human remains from these three sites were
found to present a homogeneous adaptive dental pattern (Lukacs and Pal 1992,
1993). Kennedy's prior analyses of skeletal variations in the Mahadaha and Sarai
Nahar Rai series suggest a homogeneous morphological pattern in terms of
stature, robusticity, and skeletal pathology (Kennedy et al. 1986, 1992). Reassess-
ment of age at death at Damdama was conducted by Robbins (2000), who used
variations in dental microstructure to resolve questions regarding palaeode-
mography. A more recent analysis of skeletal pathology at Damdama was con-
ducted within the framework of subsistence transition theory (Lukacs and Pal
2003). This study found that indicators of iron deficiency (cribra orbitalia, porotic
hyperostosis) and nonspecific systemic infection (periostitis) were absent from the
Damdama skeletal series, findings consistent with a nomadic mobility pattern, low
population density, and a hunting and foraging subsistence system. This result
confirms prior insights regarding subsistence and diet derived from the dental
334 ASIAN PERSPECTIVES . 42(2) . FALL 2003

pathology profile and tooth size variation reported at Damdama, and further
affirms the similarity of the denizens of Damdama to their neighbors at Mahadaha
and Sarai Nahar Rai.

METHODS

The level of mechanical loading of limbs in different types of activities and

mobility patterns may directly impact the skeleton. Evidence of differential
mechanical loads may be discerned in the size and rugosity of muscle attachment
sites (entheses and musculoskeletal stress markers; Hawkey and Merbs 1995),
pathological fractures and arthritic reactions (Lovell 1994), and in the cross-
sectional geometry of long bone diaphyses (Stock and Pfeiffer 2001). Successful
osteological research on activity patterns of past people may focus on a single
individual or small sample of specimens (Hawkey 1998; Lai and Lovell 1992),
or alternatively, may involve skeletal series of moderate to large size that are
approached from a statistical or population perspective (Hawkey and Merbs
1995; Robb 1998). All postcranial skeletal elements in the Damdama series were
observed with the naked eye and under low magnification (3x) for pathologi-
cal lesions (Ortner and Putschar 1981) and peri- and postmortem modifications
(White 1992).
The analysis of entheses or musculoskeletal stress markers in the skeletons from
Damdama was conducted prior to the publication of standards for evaluating the
relative degree of hypertrophy at specific sites. Two sets of standards are cur-
rently available that use discrete categories for classifying the size and severity of
musculoskeletal stress markers. Hawkey and Merbs (1995) recommended a dis-
crete fourfold classification (0 = absent, 1 = faint, 2 = moderate, 3 = strong) that
they used to evaluate muscular origin and insertion rugosity, stress lesion size, and
ossification exostosis. By contrast, Robb (1998) used a five-scale system to classify
18 muscle insertion sites in 56 skeletons from the Iron Age cemetery at Ponte-
cagnano, Italy. Our description of variation in entheses at Damdama will refer to
approximate equivalences in the Hawkey and Merbs (1995) methodology, as their
system of classification provides more extensive photographic documentation of
skeletal variation. All muscle and ligament attachment sites were examined with
reference to their size, roughness, and proniinence.
Supplemental guidelines for paleopathological analysis follow the procedures
described by Aufderheide et a1. (1998) and Lovell (1997, 2000), while recognition
and scoring of degenerative joint disease, or osteoarthritis, follows protocols
established by Jurmain (1999). Long bone measurement followed standardized
procedures for maximal length as described by Haas et a1. (1994) and Ubelaker
(1989). Initially, only complete and well-preserved long bones were used in the
estimation of stature. To enhance sample size, long bones that had one epiphysis
preserved and were at least three-quarters complete and a few long bones with a
slight degree of deformation were included in the analysis. Maximal length was
estimated for long bones that were incomplete or warped. Estimation of living
stature from maximal long bone length was calculated according to formulae
derived from skeletal samples of American whites of European descent (Trotter
1970; Trotter and GIeser 1958) to facilitate comparison with stature data reported
by Kennedy and associates (1986, 1992) for Mahadaha and Sarai Nahar Rai.
LUKACS AND PAL . MESOLITHIC SKELETAL VARIATION OF THE GANGA PLAINS 335

RESULTS

Ellthesia! Hypertrophy, Osteoarthritis, alln Activit)' Patterns

Entheses are sites throughout the skeleton where muscles attach to bone by
tendon. In a structural-functional analytic paradigm, the extent and rugosity of
entheses is approximately proportional to the size and activity of a muscle. In the
skeletal sample from Damdama, enthesial hypertrophy was systematically eval-
uated as an indicator of skeletal function, activity level, and mobility, and as one
factor contribu ting to the perception of "skeletal robusticity." En thesial hypertro-
phy was observed at multiple loci throughout the skeleton and 57.6 percent
(19/33) of the specimens displayed enthesial hypertrophy at one or more sites.
The distribution and frequency of enthesial hypertrophy by skeletal location is
presented in Table 2, which shows upper and lower extremities equally affected.
Affected sites are distributed across six locations in the upper limbs, with common
loci of enthesial hypertrophy including the forearm, proximally near the elbow at
the supinator crest and radial tuberosity, and distally at en theses of the pronator
quadratus muscle. In the lower extremity, enthesial hypertrophy is clearly nlOst
prevalent and prominently developed at one site-the posterior proximal surface
of the tibia, where the soleal line is developed into a rugose welt-like swelling
(Hawkey and Merbs: grade 3). In more extreme instances this enthesis is hyper-
trophied into a ridge-like crest from which the soleus muscle originates (Hawkey
and Merbs: grade 4). Hypertrophy of the soleal line is a marker of frequent and
forceful plantar flexion and may be associated with high mobility, carrying heavy
loads long distances, or locomotion in hilly terrain. Enthesial hypertrophy of the
soleal line is equally well developed in the Damdama skeletal series among males
and females and among adolescent and fully adult individuals of all ages.
A single instance of ossification exostosis was observed on the posterior surface
of the right femur of specimen Damdama 23, an adult female. The exostosis is
located 10 cm below the lesser trochanter and is on the lateral aspect of the linea
aspera but contiguous with it. This area represents the proximal limit of the origin
of the biceps femoris muscle. The "bone spur" is chevron shaped with the apex
pointing proximally, the arms diverging distally, and a vascular cortical impression
trailing from the disto-Iateral aspect of the exostosis in a disto-Iateral direction.
In areal extent (3 cm long) and degree of projection from the cortical surface
(> 10 mm), this exostosis is classified as a strong expression (OS = 3) of the trait
(Hawkey and Merbs 1995). Ossification exostoses are usually due to abrupt
macrotrauma such as muscle rupture. This particular exostosis represents a serious
injury to the short head of the biceps femoris muscle (the lateral member of the
hamstring group), probably caused by a unique, extreme action, such as hyper-
extension at the knee joint.
Levels of mobility within and between groups may also be approached
through the analysis of postcranial osteoarthritis. Theory and issues relevant to
this research strategy are reviewed by Hemphill (1999), who adopts a bio-
archaeological approach to osteoarthritis in evaluating levels of mobility among
Great Basin hunter-gatherers. Three types of bone modification were recorded as
evidence of osteoarthritis: eburnation, porosity, and proliferative bone growth
(osteophytes) at joint margins (Jurmain 1999). Sample size in this analysis fluc-
tuates from joint to joint, and not all articular surfaces could be observed at each
 TABLE 2. DISTRIBUTION AND RELATIVE FREQUENCY OF ENTHESIAL HYPERTROPHY AT DAMDAMA

UPPEIl EXTIlEMITIES

CONOID DELTOID SUPINATOIl RADIAL PIlONATOIl

ENTH ESIS TUBEIlCLE TUBEIlOSITY CREST TUBEROSITY QUADIlATUS BIlACHIALIS TOTAL UPPEIl

'Xl 9.1 6.1 9.1 12.1 18.2 3.0 576

n 3 2 3 4 6 1 19

LOWEn EXTREMITIES

GLUTEAL ANTERIOR INFERIOR GREATER LINEA OSSIFICATION

ENTHESIS LINE ILIAC SPINE TIlOCHANTER ASPEIlA EXOSTOSIS SOLEAL LINE TOTAL LOWER

(% 3.0 3.0 3.0 12.1 3.0 18.2 42.4

n 1 1 1 4 1 6 14

'Yr, = frequency of hypertrophic development at a specific site in comparison with other sites of musculoskeletal stress markers or en thesis hypertrophy.
n = number of instances in which a specific site (musculoskeletal stress markers or enthesis) exhibited hypertrophic development (grade 3 or 4 of Hawkey and
Merbs 1995).
LUKACS AND PAL . MESOLITHIC SKELETAL VARIATION OF THE GANGA PLAINS 337

TABLE 3. LOCI AND RELATIVE FREQUENCY OF OSTEOARTHRITIS AT DAMDAMA

UPPER LIMBS SHOULDER ELBOW WRIST HAND

% n <Xl n % n % n
0.0 18 4.5 22 0.0 11 10.0 10

LOWER LIMBS HlP KNEE ANKLE FOOT

l).{l n (% n IX) n %l n
0.0 16 0.0 17 0.0 13 0.0 13

% = percentage of observed joints afflicted with osteoarthritis (includes marginal osteophytes, ebur-
nation, and porosity).
n = number of joints observed (one or more of the articular surfaces at the joint were preserved for
study).

joint, due to post-burial diagenetic and taphonomic factors influencing the differ-
ential preservation of skeletal parts. The age structure of the skeletal sample, on
which osteoarthritis was assessed, ranges from 16 to 52 years with, a sex-pooled
mean of 33.3 years. No significant difference exists between sexes in mean age at
death. At Damdama, evidence of osteoarthritis was present, but rare. Vertebrae
were poorly preserved and underrepresented in the sample; however, vertebral
osteophytes were observed in three specim.ens (Damdama 1, 8, 12) of seven with
preserved vertebrae (absent in Damdam.a 18a, 20b, 36a, 36b). Osteoarthritis of
the appendicular skeleton is low in frequency and was observed bilaterally in the
elbows of one individual (Damdama 1, female) and in the hand of another (meta-
carpals, Damdama 12, female). The low incidence of osteoarthritis in the appen-
dicular skeleton suggests that predisposing factors such as repetitive forceful use of
the limbs and traumatic injury were also rare (Table 3). By contrast, involvement
of the axial skeleton appears more common, and may indicate frequent bearing of
heavy loads-an observation consistent with hypertrophy of the soleal line.
Osteoarthritis of the temporomandibular joint is rare at Damdama and is related
to the biomechanics of mastication not mobility and, consequently, will be dis-
cussed in more detail elsewhere (Lukacs and Pal 2003).

Stature at Damdama: A Comparative Assessment

Descriptive statistics were computed for all postcranial osteometric variables for all
traditionally recorded dimensions. In Table 4, we present mean maximum length
of long bones of the Damdama skeletal series by sex. The data on which these
summary statistics were computed constitute the basis for the analysis of upper
and lower limb segment proportions and the estimation of stature. Maximum
long bone length was entered into regression formulae for white Europeans to
estimate stature (Trotter 1970). Estimations of living stature could be calculated
for 29 specimens from Damdama, yielding mean (± 1 standard deviation) values
of 179.1 cm (±8.1 cm) for 18 males and 173.0 cm (±10.1 cm) for 11 females.
These values are graphically compared with mean stature estimates for Maha-
daha, Sarai Nahal' Rai (Ganga Plain sites), and for Lekhahia (located in the Kai-
33 8 ASIAN PERSPECTIVES 4 2 (2) FALL 2003

TABLE 4. MEAN LONG BONE LENGTH BY SEX (IN 111111)

ELEMENT N MEAN SO MIN MAX

MALE

Clavicle 5 152.8 6.3 145.0 158.0

Humerus 7 352.4 18.3 315.0 370.0
Radius 6 276.7 26.2 254.0 325.0
Ulna 5 293.0 22.3 268.0 322.0
Fem ur 10 503.8 23.0 454.0 537.0
Tibia (, 428.7 10.7 417.0 447.0
Fibula 5 389.0 27.3 368.0 425.0

I'EMALE

Clavicle 2 134.0 12.7 125.0 143.0

Humerus 4 339.0 26.1 309.0 370.0
Radius 3 275.0 23.5 248.0 291.0
Ulna 4 284.5 22.2 262.0 309.0
Femur 3 469.3 18.9 453.0 490.0
Tibia 3 388.3 29.7 363.0 421.0
Fibula 4 385.8 29.0 353.0 417.0

mur Hills) in Figure 2, and reveal greater intersite consistency for males than
for females. The level of sexual dimorphism at Mahadaha (6.5 percent, n = 17;
male 181.1 ± 5.6 cm, n = 13; female 169.3 ± 4.8 cm, n = 4; Lukacs and Pal
1993) and at Chalcolithic Mehrgarh (8 percent, n = 20; male 171.1 ± 7.1 cm,
n = 6; female 159.0 ± 4.7 cm, n = 14; Lukacs and Hemphill 1991) is greater
than at Damdama for which sexual dimorphism is 3.4 percent [percent sexual
dimorphism = (M - F1M) X 100].
The Damdama and Mahadaha series are used to place Indian Mesolithic stature
in broader comparative context, because larger samples are available from these
sites and because the two females from Sarai Nahar Rai yielded a suspiciously tall
mean stature (188.9 ± 1.6 cm; calculated from data presented in Kennedy et al.
1986: 71, Table 6; using stature estimates from bone lengths and formulae with
the lowest standard error). Mean stature for both Damdama and Mahadaha is sig-
nificantly taller than European Mesolithic skeletal samples. Recently published
stature data for Mesolithic Europe reveals a dinal pattern from Western European
sites with short stature, to Eastern groups with intermediate stature (Formicola
and Giannecchini 1999). Figure 3 shows that individuals from both Damdama
and Mahadaha are significantly taller than Mesolithic Europeans. In Formicola's
opinion, the observation that Upper Palaeolithic skeletal series are taller than
Mesolithic series in Europe may have several plausible explanations: (1) better
nutrition; (2) retention of ancestral heat-adapted limb proportions; (3) outbreeding
mating patterns led to genetic heterozygosity; and (4) natural selection optimized
stride length and locomotor efficiency. Collectively, these factors may also have
played a role in selecting for tall stature among North Indian Mesolithic groups,
which display greater similarity in stature to Upper Paleolithic than to Mesolithic
Europeans.
 190 ---1-
_.1

~~~--1---j---
185

---E
II) 180 Male Mean

0
'-' 175
...
(1)

...
::l 170
Female Mean

...
m
(f)
165

160 •
o

155

150
Damdama Mahadaha Sarai Nahar Lekhahia
(OOM) (MOH) Rai (SNR) (LKH)

Mesolithic Site Names

Fig. 2. Mean stature for Mesolithic Indian skeletal series: Ganga Plains (OOM, MOH, SNR) and
Vindhya Hills (LKH) (error bars = ±1 standard deviation). Oata for MDH and SNR from Kennedy
et aL (1986, ] 992), for LKH from Lukacs and Misra (1997) and for DOM revised from Lukacs and
Pal (1993:76], Table 14)_

190
- Eur-male
185 W///A< Eur-female

-E ....... _....
_
_
India-male
India-female
Eastern Europe

-- 180 ---_.
(,)

x
Q) 175 ._-- --- ._....
C/) Western Europe
>- 170 f-. ... i···
.c
~
--
:::J
I II
C/)
165

160
1 1 r- ~
~.
~--_ ..
c:
I .. ~ ~
~
~

r rI
III ~
~ ~
~~
Q) 155 ~-_ ..

~ ~ ~ ~
~ ~ ~
150 ~.
~ ~ ~
~
~
~-
~
~--
~
~
~ ~ ~--
~
145
~ ~ ~ ~ ~ ~ ~ ~ ~ ~ '---
~7><:>c ~~~0 ~\7>Co 0\0~" c1>~1>
o\~ /),c, ~7> 4>c, /+-7> c7>,<:>7>
~o\7> o-1
~0V:< 1-07> o~~ 0(,0
-..1\1>
~'\ ~1>,<:>7i o1>~
"'....1> ~,0 \:>1Yo~ -..11>
,\0

Fig. 3. Mesolithic variation in stature: Europe and North India compared (error bars = ±] standard
deviation). European data frol11 FOrInicola and Giannecchini (1999: 324, Table 4a).
34° ASIAN PERSPECTIVES . 42(2) . FALL 2003

Limb Length, Skeletal Robusticity, and Climatic Adaptatiol1

With the exception of their utility in estimating stature of the deceased, data
derived from postcranial osteometry have long been viewed as esoteric minutiae
relegated to the appendices of anthropological and archaeological reports. Today,
analysis of the postcranial skeleton has taken center stage in bioarchaeological
research and currently involves biomechanical analysis of stress on the skeleton
(Ruff 2000) and utilizes a blend of traditional osteometry and innovative tech-
niques, such as CT scans and digital image analysis (Bridges 1989), which yield new
perspectives on activity patterns and climatic stresses in prehistory (Ruff 1999).
A bivariate scatterplot of lower limb lengths for individual specimens from
Damdama is presented in Figure 4. For comparison, the mean limb length values
of populations residing in warm and cold climates are included i,:- the plot (data
from Trinkaus 1981). The regression lines are clearly distinct, with warm climate
groups having longer tibias for a given femur length than groups from cold climes
(see Fig. 4 caption for group names). The specimens from Damdama fall nearest
the regression line for warm climate groups, who have relatively long limbs over-
all and especially long tibias. This pattern of distal lin,b segment elongation also
characterizes the upper extremities of the Damdama skeletal series and illustrates a
pattern of variability consistent with the ecogeographic rules of Allen (1877) and
Bergmann (1874). Long limbs and linear body builds are selected for in hot arid

460

-E
440

420
+_..
!
r = O.~O ,
Y= 0:;8_59x - 9..:.54'
..! ._. Gamdama
specimens
,
.---
._--• -_._,! ----
.
E
x 400
ro Y
-.....
Warm climate
E group mean$ .....--.41:'"".!
. r; 380 -- .':;;..---r' - -- ~-
C) ......--.. . !
c: ...<'
Q)
...J 360 ---,
ro
:0
i= 340
Cold & Warm Olimate Gro~p Means
f~om Trinkau~ (1981:19~, 202)
320

300
400 420 440 460 480 500 520 540
Bicondylar Femur Length (mm)
Fig. 4. Lower limb proportion scatterplot: tibia vs. femur. Circles = values for individual specimens
from Damdama (DDM-12, loa, lob, 23, 24, 25, 30a, 30b); triangles = warm climate group means
(Bantu, Egyptian, Melanesian) and the individual Kenyan fossil, HOII/o cl:~asfc" (WT 15,000; Walker
and Leakey 1993); squares = cold climate group means (Eskimo, Lapplander, Neandertal). Data
source: Trinkaus 1981: 199, Table 3; 202, Table 5).
LUKACS AND PAL . MESOLITHIC SKELETAL VARIATION OF THE GANGA PLAINS 341

regions because they facilitate physiological thermoregulation through heat dissi-

pation. When compared with Holliday's (1999) data on Mesolithic European
limb length, and upper (brachial) and lower (crural) limb indices, the Damdama
skeletons exhibit significantly longer limbs.
The analysis of postcranial robusticity has recently been used to resolve anthro-
pological problems of general interest to archaeologists and bioanthropologists,
such as the likelihood of continuity vs. replacement in the origin of modern
humans in Europe, and changes in the level of physical stress across the subsis-
tence transition from foraging to farming in the Southeast United States. Results
of robusticity studies have provided both surprises and insights, as illustrated
below. Here we raise the contentious question, "Do the Mesolithic inhabitants of
Damdama exhibit robusticity in the morphological structure of their postcranial
skeleton?" Skeletal robusticity in the Damdama skeletal series is evaluated using
two standard measures of robusticity traditionally employed in human osteology:
the diaphyseal robusticity index and epiphyseal robusticity index. The diaphyseal
robusticity index relates a measure of bone shaft size (circumference, transverse, or
anterior-posterior diameter) to a measure of bone length (maximal, physiological,
or articular). The epiphyseal (or articular) robusticity index relates a measure of
articular or epiphysis size (head diameter, biepicondylar, or bicondylar width) to a
measure of bone length (see Pearson 2000, for bone and index-specific formulae).
Though different investigators employ a variety of different formulae in comput-
ing indices of robusticity, we have selected two recent studies for the purpose of
contextualizing the level of robusticity at Damdama (Collier 1989; Pearson 2000).
In the first analysis, Pearson's (2000) diaphyseal robusticity index is plotted on
the horizontal axis and the epiphyseal robusticity index is graphed on the vertical

TABLE 5. ROBUSTIClTY INDEX: FEMUR

DIAPHYSEAL EPIPHYSEAL (DISTAL)

SPECIMEN BICONDYLAR BICONDYLAR MAXIMUM

NUMBER SEX AI' + ML LENGTH INDEX WIDTH LENGTH INDEX

12 F 52 459 11.3 08 405 14.0

loa M 75 530 14.2
lob M 70 498 15.3
20b M 00 497 13.3
21 M 54 448 12.1
30a F 54 483 11.2 oS 493 13.8
36b M 50 475 10.5 09 486 14.2
l1 sci l1 sci
Meal1 F 2 11.3 0.07 2 14.2 0.57
Meal1 M 3 12.0 1.40 3 14.6 0.64
Total F&M 5 11.7 1.07 5 14.4 0.57

EPIPHYSEAL (PROXIMAL)

SPEC. NO SEX HEAD MAX LN INDEX

12 F 43 405 9.2
23 M 48 50S 9.5
34 2 ASIAN PERSPECTIVES . 42(2) . FALL 2003

12..,..---------------------------,----,

'2£
~ 0)
E
c
~~
x ....
o ~
•
'- E
E::~ 11
x x
0) ro
•••• •
-g E
->0'- -- ••
• •/
~ 0)
.5:? Q)
en E • Cold climate
~ .~
o -0
0:::-0
10 •
••• / ..... Cold climate Regr
• Warm climate

.~
_ ro A Ganga males
ro 0)
0)£ • Fossil forms
lIl_
- - Fossil Reg Line
>oro Damdama~ •
"E.oE
.-
Q.O)
W:=. ~MDH & SNR mean
9+-_+-~-+-~-...___+-I__+__+-_.___t___jf___+__+-+__t_~-+-_+____j

11 12 13 14 15
Diaphyseal Robusticity Index [(ap+ml)/bicondylar In]
Fig. 5. Diaphyseal and epiphyseal robusticity: Damdama and Ganga Plains male femora in compara-
tive context. This graph is a revised version of Pearson's Fig. 4b (2000:586), with DDM and Ganga
male samples added. Data and sample names of individual cold and warm climate groups, and fossil
forms, taken from Pearson (2000:578-579, Table 4; 580-581, Table 5) Ganga males (MDH and
SNR; from Kennedy et al. 1986, 1992). (Figure redrawn from Pearson [2000; Fig. 4b, p. 586], with
DDM and Ganga males and regression lines added.)

axis (Fig. 5). Squares and the dotted line represent cold climate groups, while cir-
cles represent samples from warm climates (data and sample names in Pearson
2000). Diamonds and the dashed line represent late Pleistocene and Upper Palae-
olithic fossil samples. Note that triangles, representing males from Damdama and
fron"l Mahadaha and Sarai Nahar Rai (pooled mean) have the lowest values for
both diaphyseal and epiphyseal robusticity. In these measures of robusticity the
North Indian Mesolithic samples are gracile rather than robust, and cluster with
groups that occupy warm environments.
In an effort to clarify disagreements regarding the robusticity of Australian
aboriginal skeletons, in contrast to European immigrants, Collier (1989) analyzed
five comparative samples in terms of multiple measures of robusticity: two urban
groups (Terry Collection, Romano-Britons), and three hunter-gatherer groups
(two Eskimo: southwest Alaska riverine and northern Alaska coast whalers; and
one Native North American, Arikara). Surprisingly, Collier found that in all
measures of robusticity, rather than being more robust as expected, aboriginal
Australians were far more gracile than all comparative samples, even the urban
groups. In both articular (epiphyseal, y-axis) and diaphyseal (x-axis) indices of
femoral robusticity, aboriginal Australians are similar to the Mesolithic group
from Mahadaha, and while Damdama and Sarai Nahal' Rai are near the mid-range
LUKACS AND PAL . MESOLITHIC SKELETAL VARIATION OF THE GANGA PLAINS 343

.c 2.8 , . . - - - - - - - - - - . - - - - - , - - - - - . . . . . . , - - - - - - - , - - - - - , - - - - - - ,
C5l • Hunters & Foragers
m - - H & F Regression
-: 2.7 Ganga Mesolithic
:::l
•
A
Modern Urbanites
Eskimo: Riverine I _--..;'
E ei ~
~~
.... 2.6 ....... , ~. ., . l

:Q ::: Nativ~ Ameri~ --- Eskimo:

....
!
(;) .~
Whalers
~~.
;:,"0 i
-g ~ 2.5
..
--- . Terry
'
Briton
c::~
~ c
CO 0 2.4
"S .2
oal
:e+ I.
« .~
"0 A Sarai-Nahar-Rai
"0
III
::I: 2.2
Ai
._---
~ iDamdamai j
1::
CT
Ul 2.1 ..L-_-+ -+ -+ --I- --I- -+-_---.I
5.8 6.0 6.2 6.4 6.6 6.8

Diaphyseal Robusticity (Collier, 1989)

sqrt (AP x ML) x 100 I Femur Length
Fig. 6. Male femur robusticity: Mesolithic India in comparative context. For details on comparative
groups: refer to Collier (1989) for modern urbanites (squares; Terry collection and modern Britons);
and hunter-forager groups (circles; riverine and whaler Eskimo samples, a Native American series,
and Australian Aboriginals); and to Kennedy and colleagues for SNR (1986) and MDH (1992) data.

of values for diaphyseal robusticity (horizontal axis), they have the lowest values
for articular (epiphyseal) robusticity (vertical axis) (Fig. 6).
In sum, Pearson (2000) found diaphyseal and epiphyseal robusticity high among
people inhabiting cold climates, and low among people inhabiting warm regions.
The Mesolithic samples from North India cluster in this analysis with the denizens
of warm climates. To Collier's surprise, Australian aboriginals were much less ro-
bust in limb indices than expected, possibly due to physiological adaptation to
heat stress, but he proposed that low-stress subsistence activities may also be
involved. The low robusticity of North India's Mesolithic foragers may be due to
their tall stature and long upper and lower extremities, variations that represent
biological responses to a warm climate. Elongation of lower limbs among the In-
dian Mesolithic skeletal series may result from the combined selective influence of
heat stress and an adaptive response favoring locomotor efficiency among mobile
foragers Further study of the biomechanics oflimb structure in these study groups
using cross-sectional geometric analysis may add to our knowledge of their post-
cranial skeletal adaptations.

DISCUSSION

Musculoskeletal Stress Markers, Enthesial Hypertrophy, and Activity Patterns

Insight into activity patterns at Damdama are modest, yet informative. They are
modest in contrast with population-level studies of larger series and with the
344 ASIAN PERSPECTIVES . 42(2) . FALL 2003

analysis of unique occupations or activities in smaller series or individual skele-

tons. For example, the analysis of musculoskeletal stress markers in a large skele-
tal sample of Hudson Bay Eskimo allowed Hawkey and Merbs (1995) to discern
gender-specific activity patterns and to support a theory of substantial change in
subsistence through time with osteological data. Alternatively, Molleson and
colleagues distinguished a suite of skeletal markers associated with the habitual
and strenuous activity of equid riding, from a distinct pattern of markers that
result from routine cart or chariot riding. This permitted the identification of a
cart driver from the Royal Cemetery in the Predynastic period at Vr (c. 2500-
2350 B.C.; Molleson and Hodgson 1993), and four femora of probable bare-back
equestrians at Kish (Molleson and Blondiaux 1994). One of the probable eques-
trians at Kish is significant because it derives from the first or second quarter of
the third millennium (2750 B.C. ± 100) and thus antedates textual and archaeo-
logical documentation of horse riding (Molleson and Blondiaux 1994: 314).
Though no highly distinctive or occupation-specific activities have been iden-
tified at Damdama, two informative patterns of nmsculoskeletal stress markers have
been identified: (1) overhand throwing, and (2) long-distance or load-bearing
locomotion. The throwing complex, may be associated with the forceful launch-
ing of spears or hurling of individual projectiles or bolas, activities envisioned as
habitual among hunters, yet not without modern parallels (baseball pitching or
cricket bowling, Kennedy 1983). This complex of musculoskeletal stress markers
includes hyper-development of the anconeus and supinator muscles, and has been
previously reported in the Mahadaha (Kennedy et al. 1992) and Sarai Nahal' Rai
(Kennedy et al. 1986) skeletal series. The recognition at Daludama of hypertro-
phy of proximal and distal forearm muscle sites (radial tuberosity, pronator quad-
ratus attachments, sharp interosseous crests) reaffirm that activities involving the
brachial complex favored the right side and were an important component of the
Dam.dama activity pattern. Archaeological evidence from Damdama includes clay
balls that could have served as projectiles or bolas, and stone points that were
undoubtedly hafted to spear shafts. These arti£1cts are consistent with the activity
patterns reconstructed from the analysis of entheses and musculoskeletal stress
markers data from human skeletons at the site.
In the lower extremity, the high level of development of the soleal line-
origin of the soleus muscle-and related markers of the ankle and foot, such as
the calcaneal tuberosity, suggest forceful and habitual plantar-flexion of the foot.
The action implicated here is the propulsive "toe-off" segment of the bipedal
stride, or an exercise known as "toe-raises" in which soleus and gastrocnemius
muscles flex to raise the body and heel of the foot from the ground, displacing
body weight to the ball of the foot and the toes. The highly developed markers of
plantar flexion of the foot in the Damdama series are interpreted to represent a
range of possible locomotor stresses, including walking long distances, carrying
heavy loads while walking, or walking up steep grades. These are clearly not
mutually exclusive activities and some individuals may have engaged to varying
degrees in all three actions. Nevertheless, a differential diagnosis of relative degree
of each activity £i'om musculoskeletal stress markers size and severity in an indi-
vidual skeleton is not possible. While extensive walking is clearly indicated from
this analysis, distinguishing the osteological signature of locomotion in hilly ter-
rain from walking long distance on level plains or differentiating osteological
indicators of locomotion while carrying heavy loads (residential mobility) from
LUKACS AND PAL . MESOLITHIC SKELETAL VARIATION OF THE GANGA PLAINS 345

walking shorter distances with lighter loads (logistical mobility), may require
more refined quantitative analysis of cross-sectional geometry of bones of the
lower extremity (Ruff 2000). Only with additional data, relating geometric vari-
ation in bone shape to the stress and strain of mechanical loads can the com-
peting hypotheses of Mesolithic subsistence advanced by Sharma (1973) and
Varma (1981-1983) be tested with human skeletal evidence (Lukacs 2002).
The musculoskeletal stress marker complexes identified at Damdam.a, in both
upper and lower extremities, are not exclusively specific to one sex. Although the
throwing complex tends to be more frequent or more highly developed in males,
it occurs in both male and female specinlens. The locomotor musculoskeletal
stress marker complex is expressed to a similar degree in both sexes and in all ages
from late adolescence to mature adult.

Variation in Stature and Limb Proportions: Multiple Selective Agents

The comparative analysis of postcranial skeletal variation among Mesolithic skele-
tal series from North India demonstrates that inhabitants of Damdama, Mahadaha,
and Sarai Nahar Rai share a phenotypic pattern of body size and proportions that
includes: (1) tall stature, (2) elongation of upper and lower limbs overall, and (3)
lengthy distal limb segments (forearm and leg) relative to proximal limb segments
(upper arm and thigh). This gracile and linear phenotype was revealed by plotting
lower limb segment lengths (tibia vs. femur) and by plotting indices of femoral
robusticity for Ganga Plains skeletal series against a range of skeletal samples from
different climatic settings that practiced different subsistence and activity patterns.
The Indian Mesolithic skeletal series consistently grouped with other samples
from warm climates. Tall stature, long limbs, and relatively long distal limb seg-
ments are collectively regarded as a suite of traits that increase body surface area
and thereby promote evaporative cooling as an adaptation to heat stress. The
elongation of long bones results in a relative reduction of transverse measures of
diaphyseal and epiphyseal (articular) width, presenting a linear and gracile skeletal
pattern.
Other factors synergistically intertwine during growth and development to
influence body size and shape and may enhance or compromise the attainment
of lengthy limbs in responding to thermoregulatory stress. Positive selection for
increased energetic efficiency during locomotion would also promote elongation
of the lower limb generally and the leg in particular. In a highly mobile hunting
and foraging society, selection for locomotor efficiency may derive through the
conferral of differential reproductive success on individuals with increased stride
length. However, an adequate nutritional foundation, providing essential nutrients
for accelerated skeletal growth, must be readily available. Paradoxically, while the
dental record at Damdama reveals high rates of enamel hypoplasia-evidence that
periodic food shortages were not uncommon-catch-up growth during times of
abundance clearly permitted the attainment of tall stature and long limbs charac-
teristic of the Mesolithic inhabitants of the Ganga Plains (Lukacs and Pal 1993).

CONCLUSIONS

The findings reported here expand our knowledge of Mesolithic life-ways in the
Ganga Plains of North India. Previous bioarchaeological research has documented
 ASIAN PERSPECTIVES . 42(2) . FALL 2003

low rates of dental pathology, large tooth size, tall stature, and generally good
health among the inhabitants of Damdama, Sarai Nahar Rai, and Mahadaha. This
study examined the relative rugosity of entheses and musculoskeletal stress markers,
prevalence of osteoarthritis, long bone lengths and proportions, and variation in
stature among 47 specimens [rom Damdama. Salient conclusions derived from
this research include:
1. In size and prominence, most entheses (muscle attachment sites) display a
normal range of variation, however two areas: (a) the elbow and forearm, and (b)
the posterior surface of the tibia, exhibit enlarged and especially rugose entheses.
Functional interpretation of these hypertrophic enthescs suggest that the forearnl
was used in forceful overhand throwing with the right hand, and that the leg was
used in walking great distances, possibly with heavy loads or in hilly terrain.
2. Evidence of osteoarthritis is rare, and suggests that potential factors that
predispose onset of degenerative bone disease such as trauma and severe occupa-
tional stresses were also rare. Osteoarthritis was observed most frequently as mar-
ginal vertebral osteophytes, suggesting that activity and weight-bearing forces
such as compression and torsion influenced this region of the axial skeleton. Bones
of the hand and the elbow joint also displayed osteoarthritic modification, but few
specimens were affected.
3. The people of Damdama, like their neighbors at Mahadaha and Sarai Nahar
Rai, are tall, especially tall when compared with the stature of Mesolithic skele-
tal series from Western and Eastern Europe. While good health and nutrition
undoubtedly contributed to their tall stature, selection for a linear body size and
shape adapted to life in seasonally hot and arid environments may also have played
a role. Limb proportions display distal segment elongation, frequently interpreted
as one con'lponent of a thermoregulator response to hot climates.
4. Are the Mesolithic inhabitants of North India morphologically robust? The
answer, both yes and no, may appear equivocal, but depends upon the frame of
reference and the skeletal character under consideration. Traditional indices of
robusticity suggest that like Australian aboriginals, Mesolithic skeletal series of the
Ganga Plain are unusually gracile. Gracility of the appendicular skeleton may
result from selective influences of heat stress and locomotor efficiency. By con-
trast, if musculoskeletal stress markers and entheses are the focus of attention, a
high degree of muscularity is evident in particular regions of the body-the fore-
ann and the leg-where repetitive and powerful movements related to throwing
and walking predominate.

ACKNOWLEDGMENTS

We express deep appreciation to past directors of the Department of Ancient His-

tory, Culture and Archaeology, Allahabad University: Dr. V. D. Misra, Dr. S. C.
Bhattacharya, and Dr. U. N. Roy, for providing access to the valuable skeletal col-
lections under their care and for making research at Allahabad University both
comfortable and productive.
Financial support for this and related research on the Mesolithic skeletons at
Allahabad University was provided by the American Institute of Indian Studies
(1974-1975, 1991-1992), National Geographic Society (1988, 1993, 1996), Na-
tional Science Foundation (1994), and Wenner-Gren Foundation for Anthropolog-
ical Research (1994-1995, International Collaborative Research Award).
LUKACS AND PAL MESOLITHIC SKELETAL VARIATION OF THE GANGA PLAINS 347

Special thanks to Sveta Chekmir, Greg Nelson, and Gwen Robbins for assistance
111various phases of research on the Damdama skeletal collection, and to my wife
Shirley and daughter Sarah for their sacrifices in support of the project.

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ABSTRACT

Rethinking new perspectives in South Asian archaeology necessitates wider appre-

ciation for insights derived from the bioarchaeological analysis of prehistoric human
skeletons. Since the 1970s, Mesolithic sites near Allahabad (Uttar Pradesh) have
yielded abundant well-preserved human skeletons permitting a bioarchaeological
approach to past life-ways. Prior research on human remains from Sarai Nahal' Rai
and Mahadaha is supplemented by this analysis of human skeletal variation in 47
specimens from Damdama. This report examines skeletal variation in muscle attach-
ment sites (entheses) and musculoskeletal stress markers, prevalence of osteoarthritis,
long bone dimensions and proportions, and estimates of stature for the human skel-
etons £i'om Damdama. The objective of this study is to better understand habitual
activity patterns, variation in stature, and adaptation to climate among Mesolithic
foragers of North India. Standardized methods of paleopathology, osteometry, and
stature estin13tion were used. While most entheses displayed a "normal" range of
development, those associated with bipedal locomotion and overhand throwing
were especially well developed. Extreme hypertrophy of the soleal line indicates
repetitious and forceful plantar flexion as in walking long distances, up hills, or with
heavy loads. Hypertrophy of the supinator crest suggests forceful overhand throwing
as in launching spears or projectiles. Osteoarthritis is unusually low in frequency,
though spinal osteophytes and arthritis of the hand and elbow were observed. Stat-
ure is tall at Damdama, a trait shared with inhabitants of Sarai Nahal' Rai and
Mahadaha. Collectively, North Indian Mesolithic groups are significantly taller than
Eastern or Western Enropean Mesolithic samples. Long lower limbs may be an
adaptation to locomotor efficiency, but may also reflect adaptation to high sea-
sonal temperatnres. Indices of distal to proximal limb segments for both upper and
lower extremities conform to physiological principles of thermoregulation and sug-
gest biological adaptation to a hot arid environment. KEYWORDS: bioarchaeology,
entheses, osteoarthritis, stature, limb proportions, climate adaptation, activity pat-
tern, Mesolithic.