QUOTE(MooseMan @ Jul 21 2006, 11:04 PM)

QUOTE(colobus @ Jul 21 2006, 04:42 PM)

In 2003 Darris Swindler and D. Jeffrey Meldrum submitted an abstract for a paper on the Skookum Cast to the American Association of Physical Anthropologists (AAPA). The abstract was between 250 and 300 words (as required) and had no diagrams or photographs (also as required).
The proposal was quickly rejected. The basis for the rejection was that you cannot infer usable information from impressions (Have them explain that to the publishers of Ichnos).
So I stated my opinion in something less than a vacuum. I state that opinion about the difficultly in getting this topic published based on the past, and the knowledge that the peer review process is rife with politics.
Please forgive my ignorance, and the fact that I can't read the whole thread right now, but who are/is Ichnos?
http://www.tandf.co.uk/journals/titles/10420940.aspIncidently, Dr. Meldrum has had papers accepted by the AAPA with "good dialogue".
This is one:
"Midfoot Flexibility, Fossil Footprints, and Sasquatch Steps:
New Perspectives on the Evolution of Bipedalism
D. JEFFREY MELDRUM
Dept. of Biological Sciences
Idaho State University
Pocatello, ID 83209-8007
meldd@isu.edu
Abstract—The chimpanzee foot is flexible near its middle, it can bend about
the axis of the transverse tarsal joint, whereas the human foot is a comparatively
rigid arched platform. Flexion at the transverse tarsal joint—the ‘‘midtarsal
break’’—uncouples the functions of a grasping, or prehensile, forefoot and a
propulsive hindfoot during grasp-climbing on vertical or inclined supports. At
some point after the transition to habitual bipedalism, these grasp-climbing
adaptations were compromised by the evolution of the longitudinal arch, which
permits increased mechanical advantage of the flexors of the ankle and improved
endurance for long-distance walking and running.
Ape, human, and Plio-Pleistocene hominid footprints were examined for
the effects of a midtarsal break. The human footprint reflects arched-foot
architecture, combined with a stiff-legged striding gait. Pressure releases occur
at particular locations behind the ball and the great toe, or hallux. Early (ca.
3.5 million years ago) hominid footprints from the Laetoli excavation confirm
midfoot flexibility, including repeated suggestion of an associated pressure
ridge. The Terra Amata footprint (ca. 400,000 years ago), yet to be fully published,
exhibits evidence of midfoot flexibility.
Several footprints attributed to an alleged North American ape, commonly
known as sasquatch, exhibit a distinctive midtarsal pressure ridge and other
indications of midfoot flexibility. In the Patterson-Gimlin film, the feet of the
film subject correlate with the kinematics inferred from the footprints, in that
a midtarsal break is present. Additional independent examples corroborate the
consistent presence of this feature, including examples of half-tracks that record
contact beneath the foot only anterior to the midtarsus. These data provide
a fresh perspective from which to consider the pattern and timing of the emergence
of the distinctive features of modern human bipedalism and bear on the
credibility of the possible existence of sasquatch. The observed and inferred sasquatch
locomotor anatomy parallels the stable adaptations that marked the greater
span of early hominid bipedalism.
Keywords: sasquatch—bipedalism—human evolution—hominoid bipedalism
One of the hallmarks of the hominoid, or ape, locomotor system is the grasping
foot. A great toe, the hallux, functions in opposition to the relatively long lateral
digits in a pincer-like grip. This foot posture is especially evident when the ape
is climbing on vertical or inclined supports. The forefoot functions as a grasping,
or prehensile organ, maintaining a secure grasp during the contact phase of a
Journal of Scientific Exploration, Vol. 18, No. 1, pp. 65–79, 2004 0892-3310/04
65
step, while the hindfoot serves as a propulsive organ providing leverage. The
plantarflexors of the ankle elevate the heel as the power arm of a lever with its
fulcrum at the midtarsal, or transverse tarsal, joint complex. This joint is actually
a compound of the articulations between the talus and navicular on the medial
aspect of the foot and the calcaneus and cuboid on the lateral aspect (Figure 1).
Elftman and Manter (1935) first drew attention to the flexibility at the midfoot
in chimpanzees. They referred to the coordinated flexion/rotation of the talonavicular
and calcaneocuboid joints as the ‘‘midtarsal break.’’ The midtarsal break
permits the corresponding, but regionally specialized, functions of the forefoot
and hindfoot, i.e., prehension and propulsion, respectively.
During terrestrial locomotion, the midtarsal break is also evident in the
chimpanzee foot. In studies of the pressures beneath the foot, the elevation of
the heel coincides with a shift of the center of pressure to the tarsus distal to the
midtarsal joint, especially beneath the cuboid (Meldrum & Wunderlich, 1998).
In contrast, the human foot is a comparatively rigid platform, built upon
a relatively fixed longitudinal arch. This adaptation incorporates the shank of
the foot into a lengthened power arm of the lever. Elevation of the heel in the
latter part of the stance phase during human walking shifts the center of pressure
beneath the metatarsal heads, especially that of the hallucial metatarsal,
Fig. 1. Foot skeletons of human (a) and chimpanzee (

. Arrows indicate the position of the
midtarsal joint complex. CA ¼ calcaneus, TA ¼ talus, NA ¼ navicular, CU ¼ cuboid.
66 D. J. Meldrum
which now serves as the primary fulcrum of the foot lever. At some point in
human evolutionary history, the hominoid legacy of midfoot flexibility was relinquished
in favor of a striding gait on much stiffer arched feet. Selection
increased the mechanical advantage of plantarflexors of the ankle, combined
with extended legs for increased stride length, thereby improving economy of
long-distance walking and running.
Lacking the opportunity to examine directly the kinematics and plantar
pressures in early hominid feet, could any signature of this pattern of flat,
flexible foot function be identified in the footprints of fossil hominids or extant
hominoids? To investigate this question, human footprints in fine damp beach
sand were examined for a variety of locomotor speeds and directional changes.
Two subjects were particularly studied at length, an adult male and a juvenile
male. In addition, an extensive sample of spontaneous trackways made by
a variety of human subjects was examined at a public beach. Next, bipedal
footprints made in a sand track box by a captive chimpanzee were documented
and correlated with simultaneous video recordings of the kinematics of his
bipedal walking. These footprints were contrasted with stereophotos and casts of
the Laetoli hominid trackway in Tanzania, Africa (ca. 3.5 million years ago).
Finally, fossilized footprints of habitually unshod humans were examined in
Hawaii (Meldrum, 2004). These are footprints left by native Hawaiians in
historic volcanic ash deposits on Kilaeua (ca. 200–400 years ago). These provide
a natural experiment to contrast early hominid footprints with modern unshod
human footprints, both laid down in a substrate of very similar consistency.
It was hypothesized that midfoot flexion associated with a midtarsal break
would, under the appropriate substrate conditions, produce a distinct pressure
release as weight was transferred distal to the midtarsus (Brown, 1999;
Meldrum, 1999). A human foot typically produces such a release proximal to the
ball of the foot, behind the hallucial metatarsophalangeal joint. The situation in
the chimpanzee footprint is somewhat confounded by the divergent hallux,
elongated lateral toes, and high angle of gait (toe-out) associated with
chimpanzee facultative bipedalism. It was found that chimpanzee footprints in
sand do occasionally demonstrate a pressure release associated with the midtarsal
break, as indicated by a primary pressure disc (Figure 2).
After the initial transition to habitual bipedalism, the hominoid grasp-climb
adaptation was compromised by shortening of the lateral toes and reduction in
the range of divergence of the hallux. The Laetoli hominid footprints, the first
direct evidence of hominid bipedalism, exhibit these modifications to the
prehensile portion of the foot to a relatively intermediate degree. There has been
continuing debate over the extent to which the Laetoli hominids display modern
human foot morphology. Some have argued that the footprints imply a foot
essentially modern in all aspects (e.g., Lovejoy, 1988; Tuttle, 1996). Others have
pointed out features that indicate the retention of more ape-like characteristics of
the foot (Deloison, 1992; Meldrum, 2000, 2002, 2004; Susman et al., 1984). I
first drew attention to a feature evident in a number of footprints (e.g., G1-25,
Evolution of Bipedalism 67
G1-26) that suggested the retention of the capability for midtarsal flexion
(Meldrum, 2000). In a depiction of a reconstruction of the Australopithecus
afarensis foot skeleton superimposed upon a Laetoli footprint, this feature can be
seen to lie immediately proximal to the position of the reconstructed midtarsal
joint (White & Suwa, 1987). Some have suggested this is possibly the result of
termite burrowing or excavation artifact; however, the repeated and
consistent position of the feature, combined with other indications within the
footprints of a transverse axis of foot flexion (Deloison, 1992), indicates its
interpretation as a pressure ridge is justified. An exceptional example of
a midtarsal pressure release is clearly evident in the G1-26 footprint, indicated
by plastic flow of the wet ash proximal to the midtarsus (Figure 3). The
possibility of this feature resulting from exfoliation of the layers of ash beneath
the contact surface of the footprint, or other excavation artifact, is excluded upon
Fig. 2. Chimpanzee footprint left in a sand track box, demonstrating a pressure release associated
with the ‘‘midtarsal break.’’ Arrows indicate proximal edge of the primary disc.
68 D. J. Meldrum
close examination. The leading edge of the flow is continuously rounded and has
the same appearance as the ash extrusion between the first and second toes of the
very clear print G1-26. In contrast, the human foot, with a fixed arch and welldeveloped
ball, leaves a very different pressure disk located proximal to the
hallucial metatarsal-phalangeal joint. No such midtarsal pressure features were
found in the sampled fossil Hawaiian footprints.
Recent analyses of early hominid foot skeletons also indicate midfoot
flexibility in australopithecine, and perhaps early Homo, feet, such as that
represented by the OH8 foot skeleton (Berillon, 2004; Harcourt-Smith et al.,
2002; Kidd et al., 1996). At some point thereafter, in the evolution of modern
human foot morphology, changes occurred to stabilize the foot platform, increase
mechanical advantage of ankle plantarflexors, and improve efficiency
and economy in long-distance, endurance walking and running (Hilton and
Fig. 3. Cast of a Laetoli hominid footprint (G1-26) exhibiting a midtarsal pressure release indicated
by plastic flow of wet ash (arrow).
Evolution of Bipedalism 69
Meldrum, 2004). Determining the timing and pattern of the evolution of these
characteristics has remained a challenge due to the paucity of fossilized
footprints or foot skeletons from the period spanning 2.0–0.5 million years ago.
Two potentially critical specimens have remained largely unpublished. The first
of these are the hominid footprints at the Terra Amata site, in southern France
(Figure 4). The site dates to approximately 400,000 years ago. The single published
photo of one of these footprints suggests a midtarsal pressure ridge and
a lack of both a well-developed longitudinal arch and a differentiated ball
(White, 1973). The footprint also appears relatively long for its breadth, possibly
indicating an elongation of the heel segment. The relative position of the
apparent pressure ridge also suggests lengthening of the heel. This would increase
mechanical advantage of the plantarflexors of the ankle, in response to
increased body mass.
Fig. 4. Footprint from the Terra Amata site (ca. 400,000 years ago) displaying a midtarsal pressure
ridge and relatively long heel, but lacking an arch and well-differentiated ball.
70 D. J. Meldrum
The second specimen is the nearly complete foot skeleton of the Jinniushan
hominid from a site in China dating to just less than 200,000 years ago. A
photograph of the skeleton appeared in a popular Chinese magazine (Lu, 1987),
but detailed descriptions or analysis of the foot skeleton have yet to be
published. Features of the foot skeleton visible in the magazine photo suggest
that stabilization of the transverse tarsal joint had occurred by that time.
Specifically, the projecting calcaneal process of the cuboid indicates that this
joint could lock in a stable position supporting a longitudinal arch. Furthermore,
the width and proportions of the navicular are similar to those in a modern
human foot, and the hallucial metatarsal and phalanges are quite robust.
Interestingly, it appears that the heel remains relatively elongated. Therefore,
based on this admittedly limited assessment, it appears that the transition to the
modern foot form, characterized foremost by the longitudinal arch and welldeveloped
ball and hallux, occurred as recently as less than 200,000 years ago.
In that case, it should be recognized that the majority of the history of hominid
bipedalism transpired on flat flexible feet, and modern human foot morphology
was a relatively recent evolutionary innovation. This observation prompts
important questions about the nature of behavioral shifts that accompanied this
transition in locomotor adaptation.
Furthermore, one could ask whether there are any extant analogs of this
sustained, and apparently successful, strategy of locomoting bipedally on flat,
flexible feet. In October 1967, Roger Patterson and Bob Gimlin purported
to have captured on film, in a remote region of northern California, a bipedal
hominoid, commonly referred to as sasquatch (Kirkpatrick, 1968). The brief film
portrays an upright hair-covered figure, exhibiting a compliant gait (walking
on flexed knees and hips) on flat, flexible feet. A trackway of clear footprints
was left in the loamy sandbar at the site along Bluff Creek. Two exceptionally
distinct footprints were cast by Patterson, representing a right and a left foot.
These were markedly flat and exhibited little or no dynamic features that might
imply the points of flexible articulation, although the exceptionally clear outline
does preserve details of contour that appear to indicate such landmark features
as the Hallucial metatarsophalangeal joint, the navicular tuberosity, and the
tuberosity of the fifth metatarsal. Subsequent events shed additional light on the
film subject’s foot morphology.
Lyle Laverty, a U.S. Forest Service timber cruiser, came upon the site a short
time after the filming and took several color slides of the subject’s deeply
impressed footprints (Perez, 2003; Laverty, personal communication). The footprints
Laverty photographed revealed clear evidence of foot dynamics. Of
particular note is the repeated appearance of a midtarsal pressure ridge
(Figure 5).
Less than two weeks later, Bob Titmus, a professional taxidermist, visited the
site and cast a series of 10 sequential footprints, a number of which had been
covered to protect them from the elements.1 Included was the print with the
prominent pressure ridge photographed earlier by Laverty (Figure 6). This
Evolution of Bipedalism 71
sequence of casts is very informative and exhibits the qualities of variation in
apparent length, toe position, and flexibility typical of a ‘‘living’’ trackway.
From enlarged frames of the Patterson-Gimlin film, portions of three step
cycles in which the feet are visible have been examined (Figure 7). Several
features are noteworthy. First is the indication of the midtarsal break late in the
stance phase of gait. This clearly correlates with the presence of the pressure
ridge in a number of the footprints photographed and cast. Second, slight
plantarflexion recoil at the midtarsal joint can be seen in the early swing phase.
This results from the rebounding of the deep plantar ligaments after being
stretched during bending of the midfoot during the midtarsal break. The foot
flexes about an approximate transverse axis corresponding to the inferred
position of the transverse tarsal joint. Third, an elongation of the heel is evident,
especially when the calcaneal (Achilles) tendon is slack, during the swing phase.
The need for a lengthening of the calcaneus to increase leverage in a biped of
this size has been discussed at length by Krantz (1999). Its significance is further
emphasized in the context of a foot exhibiting midfoot flexibility. The
protruding heel of the film subject was taken by some skeptics as indication
of an artificial foot protruding posteriorly beyond the hoaxer’s own heel.
Alternatively, it can be seen as a sound and reasonable anatomical adaptation to
the animal’s size and foot architecture, and accords with evolutionary trends in
hominid locomotion already discussed.
In a reconstruction of the inferred skeletal configuration of the sasquatch
foot, this combination of midtarsal flexibility and heel elongation is depicted
Fig. 5. Footprint photographed by Lyle Laverty at the Patterson-Gimlin film site.
72 D. J. Meldrum
(Figure 8) based on the footprint of the Patterson-Gimlin film subject exhibiting
a distinct pressure ridge. The pressure ridge in the proximal portion of the
footprint implies the retention of the midtarsal break. Indeed, a number of
footprints attributed to sasquatch exhibit what has been interpreted as a midtarsal
pressure ridge.
Fig. 6. One of a series of 10 consecutive footprints cast by Bob Titmus at the Patterson-Gimlin film
site.
Evolution of Bipedalism 73
Another manifestation of this midfoot flexibility is evidenced in instances
where the sasquatch has run with the heel elevated off the ground. A modern
human adopts a digitigrade posture when sprinting, balancing support of body
mass over the heads of the metatarsals and the toes. This is especially
concentrated at the medial ball of the foot, under the hallucial metatarsophalangeal
joint, and is accomplished due to the stability of the longitudinal arch
of the foot. A hominoid would merely be able to elevate the heel, but not the
midfoot, because the foot would flex at the midtarsal joint, without the support
of the arch. An example of this running foot posture was first recognized in a set
of tracks I was shown near the Blue Mountains in southeastern Washington. One
very clear track, deeply impressed in mud, showed no indication of a heel
imprint. The inferred position of the midtarsal joint implied by the ‘‘half-track,’’
as I came to refer to them, agreed very well with the position of the pressure
ridge in other footprints in the series (Figure 9).
A further example of the half-track was identified in a trackway found in
northern California on the Blue Creek Mountain Road. In 1967, shortly before
the Patterson-Gimlin filming incident, a long line of tracks was found along
a logging road and investigated extensively (Green, 1978). Don Abott, an
archeologist from the Royal British Columbia Museum, Victoria, B.C., took
a series of color photographs of examples of the footprints (Green, personal
communication). One photograph shows a distinct half-track, which, when superimposed
on a complete track, can be seen to terminate at the inferred position
of the midtarsal joint.
Yet another example comes from the west coast of Washington State. Officer
Dennis Heryford, a Deputy Sheriff for Greys Harbor County, responded to a
reported disturbance at a construction site in 1982. He discovered a line of
Fig. 7. Three consecutive frames of the Patterson-Gimlin film, in which the subject exhibits
midfoot flexibility.
74 D. J. Meldrum
footprints emerging from the timber and crossing a cleared muddy landing. The
track returned in the direction of the forest with twice the step length and left
a series of half-tracks. In this case the contour of the calcaneocuboid joint is
evident on the full-length footprint, and the proximal edge of the half-track
terminates at that point. These, and other repeated examples of midtarsal pressure
ridges and half-tracks, evidence the presence of a flexiblemidfoot in sasquatch
foot functional anatomy.
Recently Haeusler and McHenry (2003) reevaluated the hindlimb morphology
of the 1.8 million year old fragmentary fossil skeleton (OH 62), attributed to
Homo habilis, and concluded that this early hominid may have had human-like
hindlimb proportions and suggested that distance travel might have evolved
early in human evolution. This, in spite of the conclusions about the primitive
morphology of the OH 8-foot skeleton, which was also attributed to H. habilis.
By 1.6 million years ago, Homo ergaster (or early African H. erectus)
unquestionably exhibited modern limb proportions, as exemplified by the
relatively complete skeleton of the Nariokotome hominid (KNM-WT 15000).
This skeleton was of a youthful hominid who was already more than 5 feet in
height when he died and would have certainly attained a 6-foot height at
maturity. Homo ergaster represents the first grade of hominid for which there is
clear evidence of expansion of its range beyond the bounds of Africa to extend
throughout much of the Old World. It has been assumed that the lengthening of
the lower extremities in early hominids was associated with the emergence of
a modern type of human walking and running, but the lengthening of the limbs
Fig. 8. Hypothetical reconstruction of the sasquatch foot skeleton (left) inferred from footprints
exhibiting midfoot flexibility and pressure release. In contrast, the human foot skeleton
(right) exhibits a longitudinal arch.
Evolution of Bipedalism 75
Fig. 9. Single footprint from a trackway photographed by the author in southeastern Washington,
demonstrating midtarsal pressure ridge.
76 D. J. Meldrum
in H. ergaster is in the extreme upper range for modern equatorial Africans.
Such limb proportions in modern African populations also reflect an adaptation
for thermal regulation in hot tropical climates (Coon, 1982). The lengthening of
the extremities increases the ratio of skin surface area to body mass and
increases heat dissipation. This is the reciprocal of a general ecological principle
known as Allen’s Rule. Thus, the increase in proportionate limb length might
reflect a response to heat stress, rather than simply an approximation to modern
human locomotor behavior.
Unfortunately, the otherwise extraordinarily complete Nariokotome skeleton
does not preserve elements of the foot skeleton, which would lend critical
insight into the specific nature of the locomotor strategy of these hominids.
Frankly, the fossil record of any H. ergaster foot skeleton is nearly
nonexistent. One exception is a single hallucial metatarsal, KNM-BK 63, of
a hominid from Baringo 500,000 years ago, which is noteworthy for its
distinctly gracile proportions as compared with modern humans (Fisher &
McBrearty, 2002). The Terra Amata footprint suggests that the older H.
ergaster foot skeleton had not yet acquired the distinctive features of the
modern human foot. Instead, this footprint appears to exhibit features of
a flexible midfoot, lacking a fixed longitudinal arch and well-developed ball.
Not until Jinniushan (H. heidlebergensis or H. sapiens) is there any skeletal
evidence implying a fixed arch and demonstrating a robust hallux. Subsequent
evidence from more recent and more plentiful Neanderthal foot skeletons
further demonstrates aspects of the modern human foot, although retaining
a greater degree of overall robusticity and relatively elongated heel segment
and toes (Trinkaus, 1983).
Should this interpretation of the pattern and timing of hominid foot evolution
be borne out by further fossil remains, then it appears that hominids were
effective bipeds for well over 3 million years, while yet lacking what has
traditionally been considered the hallmark of human bipedalism—the longitudinal
arch. The stabilization of the midfoot and associated modifications,
especially to the distal foot, i.e., shortening of the toes, increased robusticity of
the hallux, development of the ball, shortening of the heel, were relatively recent
innovations that marked a shift in hominid locomotor adaptation to skeletal
gracilization combined with endurance walking and running.
The evidence of midfoot flexibility in the foot and footprints of sasquatch, and
retention of other primitive characteristics of foot proportion, is significant, not
only for the assessment of the purported existence of this hominoid, but in that
this combination of foot morphology and locomotor behavior provides a novel
perspective on the evolution of bipedalism. The sasquatch foot and footprints
exhibit intriguing parallelism to the morphology of the hominid foot as here
presented. This bipedal hominoid may provide an enlightening analog to
hominid locomotor evolution, Ken Yielding insights and refinements to our understanding
of the pattern and timing of modern human locomotor innovations.
The inferred architecture of the sasquatch foot seems well suited to the physical
Evolution of Bipedalism 77
aspects of the terrain of its purported range. Together with observations of its
gait, its locomotor anatomy and behavior present a functionally coordinated
complex that parallels the highly stable adaptations that marked the greater span
of hominid bipedalism, modified in response to dramatically increased body
size. The combination of broad flat flexible feet, elongated heels, prehensile
toes, and compliant gait constitute an elegant adaptation for a giant terrestrial
biped evolved in a mountainous forested habitat. These observations would seem
to lend considerable affirmative evidence for the existence of an unrecognized
North American ape.
Note
1 The original casts are now housed in the Willow Creek—China Flats Museum,
Willow Creek, California. Silastic rubber molds of eight of the 10 original
casts are held by the Smithsonian. Copies of a number of these casts are held in
the author’s laboratory.
Acknowledgments
I wish to thank Gordon Strasenburgh and the Society for Scientific
Exploration for inviting this contribution. This research was supported in part
by grants from Jack Mayfield, the ISU Faculty Research Committee, and the
Leakey Foundation.
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