Journal of Biomechanics 32 (1999) 163—164
Neandertal knees and ankles:
a comment on Miller and Gross
Erik Trinkaus
*
Department of Anthropology, Campus Box 1114, Washington University, St. Louis, MO 63130, USA

U.M.R. 5809, du C.N.R.S., Laboratoire d+ Anthropologie, Universite& de Bordeaux I, Avenue des Faculte& s, 33405 Talence, France
Received 23 July 1998; accepted 24 September 1998
I was pleased to see the application of current bio-
mechanical analytical techniques [SIMM (Delp et al.,
1990)] to the interpretation of past human locomotor
efficiency by Miller and Gross (1998). The appropriate
application of such methodologies will greatly assist
those of us who study human paleobiology to compre-
hend the adaptive significance of both past changes
in human biology and the nature of modern human
variability. However, I have difficulty accepting some
aspects of their interpretation of Neandertal locomotor
efficiency.
They used mean percentage differences in the ratios of
knee and ankle moment arms to tibial length to adjust
a modern male based model for muscle moment calcu-
lations. At the knee, patellar thickness and tibial tuberos-
ity projection (anterior tuberosity to mid-condyles) were
used as measures of the quadriceps femoris moment arm,
and at the ankle the posterior pedal moment arm (mid
talar trochlea to calcaneal tuberosity) was employed to
approximate the triceps surae moment arm. Although
the patellar, proximal tibial and posterior pedal skeletal
measures are reasonable approximations (on dry skeletal
remains) of the actual moment arms (or a portion thereof
in the case of the patellar thickness), scaling them to tibial
length has consequences. Tibial length, except in unusual
athletic activities, does not represent the actual load arms
for these muscles.
The normal locomotor load arm for quadriceps
femoris is the perpendicular from the knee axis of rota-
tion to the line between the hip and ankle joints. Since
Neandertals had hyperarctic body proportions with rela-
tively short tibiae (Holliday, 1997), tibial length is a poor
* Correspondence address: Dept. of Anthropology, Campus Box
1114, Washington Univ., St. Louis, MO 63130, USA. Tel.: 001 314 935
5207; e-mail: trinkaus@artsci.wustl.edu.
0021-9290/99/$ — see front matter  1999 Elsevier Science Ltd. All rights reserved.
PII: S 0 0 2 1 - 9 2 9 0 ( 9 8 ) 0 0 1 6 3 - 8
surrogate measure for this load arm across ecogeographi-
cally diverse human populations. Across two temperate
climate recent human samples (US Amerindians and
Euroamericans), this moment arm is highly correlated
with femoral length (r"0.952), tibial length (r"0.971)
and femoral plus tibial length (r " 0.999). Yet, Nean-
dertal leg proportions make them highly significantly
different (P(0.0001) from both recent human samples
in each comparison, with their load arms being longer
than predicted by their tibial lengths. When patellar
thickness and tibial tuberosity projection are divided by
the resultant load arms, the Neandertal ratios do remain
significantly higher than those of the recent human sam-
ples (0.156 $ 0.016 versus 0.125 $ 0.011 and 0.128
$ 0.010 for the patella, and 0.307$0.034 versus $0.271
$ 0.024 and 0.234 $ 0.025 for the tibia).
However, the ultimate issue is whether the Neandertals
had a more effective arrangement for their knee exten-
sors, and that involves assessing whether the required
contractile force of quadriceps femoris for a given body
size was significantly different from those of recent hu-
mans in similar activities. In a clinical context, body
weight as the external load to a lower limb joint is
normally constant. In a comparative framework, that
cannot be assumed. Indeed, the same hyperarctic body
proportions which gave the Neandertals short tibiae also
gave them broad trunks and high lean body masses for
their statures (Ruff, 1994; Ruff et al., 1997; Trinkaus,
1981). Incorporating the differential body masses to stat-
ure (or leg length) into the analysis may well eliminate
most if not all of the differences between the samples.
The normal locomotor load arm for triceps surae is
the distance from the talocrural axis of rotation to the
region of ground reaction force, normally around the
metatarsophalangeal articulations during the primary
propulsive portion of the stance phase. Consequently, the
anterior pedal moment arm (mid talar trochlea to distal
164 E. Trinkaus / Journal of Biomechanics 32 (1999) 163—164
metatarsal 1 head) best approximates this value skel-
etally. When the ratio of the posterior pedal moment arm
to the anterior one is compared between Neandertals
(0.444 $ 0.034) and recent humans (0.430 $ 0.038 and
0.387 $ 0.028 for prehistoric Egyptians and US Amerin-
dians, respectively), the difference between Neandertal
plantar flexor mechanical advantages and those of the
recent human samples is greatly reduced from the 13 and
30% provided by Miller and Gross (3.3 and 14.7% on
average; P" 0.305 and 0.003, respectively). Incorpora-
tion of the greater relative body masses of the Neander-
tals would further reduce the difference.
Furthermore, Miller and Gross (1998) compared the
Neandertal mean values only to a single modern (Euro-
american?) male. There is increasing evidence that
modern industrial-society populations are experiencing
significant postcranial gracilization relative to pre-indus-
trial human populations (Anderson and Trinkaus, 1998;
Ruff et al., 1993; Trinkaus and Hilton, 1996). Conse-
quently, however relevant models based on living Euro-
americans may be for clinical analyses aimed at treating
those populations, they represent only a microcosm of
both living humanity and the human condition over even
the past few centuries and an extreme pattern of appendi-
cular hypotrophy. Such models therefore are seriously
limited beyond providing a heuristic model for assessing
the biomechanical affinities of diverse and past human
populations. Moreover, use of an individual model and
mean sample values removes any possibility of compar-
ing populations is an appropriate statistical framework.
Consequently, it remains unclear whether the Nean-
dertals had significantly greater knee extensor and ankle
plantarflexor moment arms, at least through the ranges
of normal locomotor knee flexion (usually )40°)
and ankle rotation ($20°). Reanalysis incorporating
appropriate load arms, full body proportions and differ-
ential body masses may yield significantly different
results.
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