Site hosted by Angelfire.com: Build your free website today!

 

 

The Controversy Behind Lucy

 

            Three main controversies surround Lucy and her species, Australopithecus afarensis.  First, are finds like the 'First Family' just one species showing a great degree of sexual dimorphism, or is it two different species?  Second, was Lucy bipedal or arboreal, or both?  Third, was Lucy indeed female, or is Lucy a male?  These controversies, as well as other topics that tie them together, will be discussed throughout this paper.

 

 

Who is Lucy?

 

 

            On the 30th day of November 1974, paleoanthropologist Donald C. Johanson came upon a bone protruding out of the sediments.  These sediments belong to a vast expanse on the upper part of the Hadar Formation, central Afar, Ethiopia.  To Johanson and his colleagues, this particular region was known as Afar Locality 288.  When the bone was being excavated, other bones were discovered in the immediate vicinity which were quickly identified as a hominid skeleton.  The skeleton consists of 47 out of 207 bones, including parts of upper and lower limbs, the backbone, ribs, and pelvis (Johanson 1996: 124).  There were some important anatomical parts missing, namely the maxilla, hands, feet, and most of the skull.  The entire skeleton was 40% complete, the most of any found at that time (Fig. 1). 

To the anthropological society, the skeleton is known as A.L. 288-1.  Johanson and his colleagues celebrated the night of the discovery; a song by the Beatles played, called "Lucy in the Sky With Diamonds".  So she became known as Lucy to the entire world.  But she is also known as Denkenesh, an Ethiopian name meaning "you are wonderful" -- well deserved, since her discovery marked a milestone in the study of mankind's prehistory (Johanson 1976: 795).


Figure 1.  A picture of Lucy that was first made public in the American Journal of Physical Anthropology (Johanson 1982: 437).

 


            The stratigraphy of the site suggests that Lucy died near a lake margin, an environment that is very different than that of today.  When the fossils were dated, along with the sediments that the skeleton was found in, using K/Ar dating methods, it was dated at 3.2 million years old, during the Pliocene epoch. 

Based on the time span in which Lucy lived, and the morphology, she was classified as Australopithecus afarensis in 1978.  Australopithecus afarensis is the earliest known member of the hominid family, living from approximately 4 to 2.8 million years ago.  Lucy's species inhabited southern and central Africa.  Remains of fossilized animals and plants indicate that the environment consisted of lush grasslands, perhaps with open-savanna woodlands.  Pollen evidence proves that these hominids lived in forests of juniper and olive trees (Johanson 1996: 108). 

It is attested that she was a mature adult based upon the erupted third molar, which is slightly worn down, as well as the closed epiphyseal lines (Johanson 1996).  All the ends of her bones were fused, along with the closed cranial sutures, indicating complete skeletal development.  Her vertebrae show signs of degenerative disease, but this is not always associated with older age (Anonymous n.d.).

            Paleoanthropologists are not able to know if Lucy herself used tools or not, but there are possibilities that her species did.  There have not been any stone tools found near any of the early australopithecine sites; however, it is plausible that they may have used wood tools.  Their feeding habits may have not required sophisticated tools, but perhaps they used tools as chimpanzees do today, to get to insects, etc. Pointed or stout wood sticks would not show up in the fossil record, thus the use of tools by australopithecines would remain a theory, unless further evidence is found.    

            The use of tools would also lead a person to think that maybe they had a sense of culture.  An important part of culture is being able to communicate to one another.  The vocal apparatus of australopithecine forms, according to Lieberman (1975), does not appear to differ significantly from those of present-day apes; vocal communications undoubtedly played a part in their linguistic systems, as it does today in apes, namely chimpanzees.  Perhaps they used signaling with waves and calls to communicate to one another. 

            We do know that Lucy and her species lived in groups, perhaps ranging from 10 to 20 individuals.  In the year of 1975 a large collection of Hominid skeletons, 13 in all, were unearthed near the Hadar region.  All were identified as Australopithecus afarensis, and soon became known as the 'First Family' (discussed later).  There were many male and female remains found of varying ages.  The skeletons varied in sizes; many believe this to be as indication of sexual dimorphism in A. afarensis. 

 

 

The Anatomical Features of Lucy and Her Kin

 

 

            When reading about the various controversies that surround Lucy and her species, anatomy plays a large role in understanding them.  As a basis for the controversies, I feel it necessary to discuss the important and distinctive anatomical features of Lucy and her species as a whole.

            The skull of A. afarensis has a small cranial vault, similar to that of a common chimpanzee.  Lucy's brain was one-third the size of modern humans, measuring from 420 to 500cc.  In view of the fact that the brain case was small and the teeth moderately large for such a small brain case, a distinct sagittal crest is present on the top of the skull.  This sagittal crest is most evident in the male species of A. afarensis, and is not a prominent feature in Lucy.  The sagittal crest is present for muscle attachment in many of today's apes, allowing the chewing of tough and fibrous material such as plants.

            Under a microscope,A. afarensis front teeth show grooves and scratches made by fibrous material, suggesting that they were used to strip food off some kind of rough vegetation (Johanson 1996: 115).  Mosaic evolution is very evident when looking at Lucy's teeth, since they are between apes and humans. Lucy's teeth and jaws are very characteristic of A. afarensis.  The upper and lower tooth rows of an ape are parallel whereas those of humans diverge posteriorly (Leakey 1997-98).  The jaw of Lucy is not entirely parallel, but it does not diverge as much as humans do either (Fig. 2).  Lucy also has a small diastema on the upper jaw, a gap between the lateral incisors and the canines which humans lack.  The presence of a diastema is one of the characteristics that separate A. afarensis from Homo. 

The upper and lower canine teeth and the tooth behind the lower canine, the lower third premolar, are intermediate between those of apes and humans (Leakey 1997-98).  Australopithecus afarensis canines are intermediate in size and shape, whereas they are large and conical shaped in apes, and much more reduced in humans.  The premolar of Lucy also resembles apes in its more oblique orientation (Leakey 1997-98).


Figure 2.  The mandible from Lucy.  Note that the jaw is in between U-shaped and V-shaped (Anonymous 2001).

 


Lucy lacked feet and hand bones, but by looking at other A. afarensis skeletons we are able to reconstruct what Lucy's feet and hands looked like.  The feet and hands were similar to that of humans.  The feet have toes that are slightly longer than humans.  Some researchers believe that the big toe was positioned in line with the other toes as in humans, while others believe that it was opposable like that of modern apes (Leaky 1997-98).  The hands however, are not like humans, but rather like apes.  Long fingers with strong muscle attachment would have allowed Lucy and her kind to clasp and hold tightly onto branches.  The thumb was longer than that of humans, however the hand was not able to cup, allowing the thumb and little finger to touch, thus not being able to form a precision grip.  The anatomy of the thumb itself illustrates why A. afarensis was not able to form the precision grip.  This is due to the fact that the thumb of A. afarensis is missing three necessary muscles that help flex and oppose the thumb into different positions (Aiello 1994).  These necessary muscles include flexorpollicus longus, for flexing, and flexorpollicus brevis, for opposing and touching each finger; the third muscle is the first palmar einterosseous muscle of Henley, which connects the thumb to its knuckle (Aiello 1994).  Because of the weakness of the thumb, this enables the other muscles in the hand and fingers to allow a power grip to be used.  This would aid in gripping branches.

Australopithecus afarensis shows only a few of the adaptations that make our hands so dextrous and manipulative (Leaky 1997-98).   It is also important to note that since most researchers believe Lucy to be a female, the anatomical features would be more pronounced in the males of A. afarensis.      


Figure 3.  Illustrating the mosaic evolution of Lucy (Anonymous n.d.).

 


She has been measured to stand at 3.5 feet tall.  Looking at the morphology of the pelvis, it is believed that Lucy was a female, although that has been brought into debate (discussed later).  Johanson states that "her leg and pelvic bones showed that she walked upright on two legs" (Johanson 1996: 101).  The lower body of Lucy will be discussed later in this paper in more detail. 

 

 

Was Lucy Male or Female?

 

 

            Some people have disagreed over whether Lucy is a female or male.  This is important, because if she is female then the single-species banner would be more plausible if Lucy were discovered to be a male.  If found to be male then finds such as the 'First Family' cannot be considered to be one species displaying sexual dimorphism.  The 'First Family' would be ruled out because if Lucy were male, meaning that she is as large as her species got, then the larger individuals of the 'First Family' must not be in the same species as Lucy.  But if Lucy were female, then she would represent the small form of her species, giving credence to the idea that the 'First Family' is two forms, the small ones being female, and the large ones being male.  The discovery of the 'First Family' is discussed in more detail in the next section.

There have been many studies done on Lucy to prove that Lucy is male or female.  Swiss researchers, Häusler and Schmid, believe that Lucy's pelvis was too narrow to accommodate an australopithecine baby, thus Lucy must be male (Shreeve 1995).  Owen Lovejoy and Johanson strongly disagree with this, stating that "the Swiss study depends on estimates of neonatal head sizes in australopithecines that are themselves based on controversial estimates of adult brain sizes, all to determine whether a hypothetical infant of a vanished species could fit through a pelvis that was itself recovered in a badly crushed condition" (Shreeve 1995: 1298).  Looking at standard traits used to ascertain sex in modern humans, the ridge on the pubic bones called the ventral arc, occurring in 95% of females, and the promontorium, a protrusion at the rear of the pelvis that juts forward in males, giving the pelvic inlet a heart shape (Shreeve 1995).  Lucy's pelvis is ridgeless gives and heart-shaped, hinting toward Lucy being a male.  This could all be a manner of interpretation, but according to Häusler and Schmid, "Lucy could be female--but only if the larger fossils were another species" (Shreeve 1995: 1298). 

 

 

One Species or Two?

 

 

             Cases in which males are much bigger in height and overall stature than females are called sexual dimorphism.  When talking about sexual dimorphism in A. afarensis, Johanson stated that "many of the bones we had came from hominids of very different sizes, and some critics argued that the variations were so great at least two species of hominids must have roamed Hadar" (Johanson 1996: 101).  However, some fail to recognize the sexual dimorphism of today's apes, an example being gorillas and orangutans.  Johanson and his colleagues believe, however, that A. afarensis is simply a case where there is great sexual dimorphism within one species.

The discovery of the 'First Family' (also known as locality 333), a collection of thirteen A. afarensis individuals, has put many questions to rest, while bringing more to the surface.  What are thought to be males would have been thickly muscled, averaging 5 ft. tall and a hundred pounds, a foot taller and two-thirds heavier than females (Johanson 1996).  Dr. Todd Olson, along with Richard and Mary Leaky, claims there was far too much variation among the fossils to be explained solely by sexual dimorphism (Shreeve 1994: 34). Some protest that the disparity in size was too great and implied two separate species, with only the larger one ancestral to Homo (Shreeve 1995).   Some believe there must be two species that were buried together, possibly in a catastrophe that swept all the bodies together in one spot.   Johanson believes that the 'family' consists of all A. afarensis, of varying ages and sexes.

Geological evidence suggests that they all died because of a sudden flash flood.  The bones are fragmented and scattered because individuals fell into or were washed into a river, rapidly transported, broken up, and scattered (Johanson 1990).  This is known as hydraulic jumble.  Because of the depositional process of burial, it is impossible to date them directly; therefore, dating of the ash layers above and below the site is important.   

Looking at the skull is an important part of diagnosing hominid species.  The differences between hominid species show up distinctly there--in the slope of their foreheads, the shape of their browridges, and the degree to which their faces jut forward (Johanson 1996: 102).  As previously mentioned, there were just bits and pieces of Lucy's and other A. afarensis skulls found.  William H. Kimbel and Johanson announced that they found the most complete A. afarensis skull in 1994.  Dated to around 3 million years ago, the find from the Hadar region in Ethiopia is approximately 200,000 years younger than Lucy herself (Shreeve 1994: 34).  Based on size, it is thought to be that of an old male, because the teeth were very worn down.  This individual's canine teeth retained the large roots typical of great ape males, but the crowns-- the parts that protrude from the gums -- were significantly smaller, closer in size to female afarensis canines.  This once again leads us to thinking about how their social system must have been organized.  Since the males did not have large canines, it is presumed that there was much less competition for females among afarensis, indicating monogamy (Johanson 1996).  For this particular skull, the brain measured more than 500 cubic centimeters, on the large side for A. afarensis.  Once the skull is studied further, Kimbel and Johanson believe that it will reinforce the single-species banner. 

 

 

The Question of Bipedality

 

 

            Bipedalism has traditionally been regarded as the fundamental adaptation that sets hominids apart from other primates (Richmond et al. 2000).  The main controversy surrounding Lucy and her kind is whether they were completely bipedal.  Lucy shows a great degree of mosaic evolution, both ape and hominid characteristics.  Apelike features are seen in her long arms, which would have dangled at her side.  Yet her leg and pelvic bones showed that she walked upright on two legs (Johanson 1996: 101).  Lucy's hips and muscular arrangement of her pelvis show that it would have been difficult for her to climb trees, as is the case with modern humans (Johanson 1996).  For some researchers, the apelike features are non-functional retentions from the common ancestor of hominids and African apes (Collard and Aiello 2000: 340).  Johanson and his long-time colleague Owen Lovejoy believe that Lucy walked upright because of certain anatomical characteristics.  As stated by Johanson in 1976, "the angle of the thigh bone and the flattened surface at its knee-joint end-- so different from quadrupedal apes-- prove that she walked on two legs".  Both Johanson and Owen think that she may have climbed trees for getting to fruit or nuts, sleeping, and escaping predators.

            Before the finding of Lucy brought certain things into question, it was believed that our early ancestors came out of the trees and onto the open savanna already being able to walk totally bipedally (Catchpool 2001).  Lucy and her species may have been bipedal but perhaps still practiced arboreality.  When looking at Lucy's wrist anatomy, specialized knuckle-walking characteristics can be seen.  Wrist extension is limited in contemporary knuckle-walkers, such as chimpanzees and gorillas.  This limited wrist extension is also seen in A. afarensis.  As stated in a paper by Richmond and Strait (2000: 384), "the retention of knuckle-walking morphology in the earliest hominids indicates that bipedalism evolved from an ancestor already adapted for terrestrial locomotion".  Australopithecus afarensis had a number of postcranial traits that were related to arboreal activity, such as a long radial neck, relatively long forelimbs and long curved fingers and toes.  This could be evidence that Lucy and her species did occasionally engage in arboreal behavior.  Also, CAT scans of australopithecine inner ear canals (reflecting posture and balance) by anatomist Dr. Fred Spoor and his colleagues at University College, London, showed they did not walk habitually upright (Catchpool 2001).

 It could be that Lucy retained these arboreal knuckle-walking features but didn’t actually knuckle-walk, i.e., they are just leftovers from her knuckle-walking ancestors.  Some people have used these knuckle-walking features, as a way to say that A. afarensis is not an ancestor to Homo, rather A. africanus is.  A paper written by Collard and Aiello (2000) suggest that A. afarensis retained some knuckle-walking features, whereas A. africanus did not.  Collard and Aiello believe that Lucy combined bipedalism, knuckle-walking and climbing. 

            The majority of researchers believe that Lucy was bipedal.  The idea of bipedality is not that simple though.  A main question is what gait Lucy and her species practice.  Was it chimpanzee-like (bent-hip, bent-knee) or erect as in humans?  Studies based on these questions have been done using computer models. 

It was found that the bent-hip, bent-knee gait is much less mechanically effective, and heat generation so much greater than an erect carriage was favored (Lincoln 1998).  A study done by Kramer in 1999 states that on a mass-specific basis, the configuration developed from the fossil remains of A.L. 288-1 uses less energy to move than, and has the same cost of transport as, the modern human configuration.  Yet, Lucy's ankle and pelvis suggest that she would have walked with slightly bent legs (Johanson 1996: 114).  If Lucy did indeed walk erect, what version did they favor, short-legged or long-legged?  Lucy is believed to be short-legged with a continued reliance on arboreal resources, with no energetic compromises. 

However, some people think that if Lucy was indeed bipedal, her activity as one was not as great as that of Homo.   

            The controversy of Lucy's bipedality was put to rest, mostly, when some hominid tracks were found in Tanzania.  A short walking stride coupled with a footprint that could only belong to a bipedal organism strengthened the belief that Lucy did indeed walk upright.

 

 

The Laetoli Tracks

 

 

            In 1976, in the Laetolil Beds of northern Tanzania, a trail of hominid footprints were discovered by Peter Jones and Philip Leakey, assistants of Dr. Mary Leakey (Fig. 4).  The entire trail was 23 meters long.  Two individuals, one smaller and the other larger, made the footprints.  The footsteps come from the south, progress northward in a fairly straight line, and end abruptly where seasonal streams have eroded a small, chaotic canyon through the beds (Leakey 1976: 453).  Using K/Ar testing, the biotite from the ash was dated to 3.6 million years old.

 It is believed that a nearby volcano was erupting and spewing ash, that it was mixed with falling rain, and that the ash hardened on the ground.  According to Dr. Leakey the crispness definition and sharp outlines convince her that they were left on a damp surface that retained the form of foot (Leakey 1979: 453).


 


Figure 4.  The Laetoli tracks and one of Dr. Leaky's assistants (Ervin 2001).

 

The trail not only was made of hominid prints but also of many other organisms that were passing by that area at the time the ash was laid.  Hares, baboons, antelope, rhinos, giraffes, cats, and hyenas are just a few makers the other prints that were found, some of these animals now extinct. 

Dr. Leakey believes these hominine prints belong to a direct ancestor to man.  To leave these prints, they must have walked fully upright with a bipedal, free striding gait (Leakey 1979).  The form of the foot was no different than ours; the toe was not divergent and the pressure was equally spread out, as in ours.  The smaller prints measure 18.5 cm long, and 21.5 cm for the larger.  The stride length for the smaller was 38.7cm and 47.2 cm for the larger.  An anthropological rule of thumb holds that the length of the foot represents about 15% of an individual's height (Leakey 1979: 453).  With that it was estimated that the smaller individual must have been 4 ft tall, with the larger being 4.5 ft tall.  Because the pressure on one set of prints is heavier than the other (Fig. 5), it is theorized that a male and female were walking together with a young child.  She may have been carrying the child, or perhaps the child stepped in the mother's set of prints, simply playing.  Some think that the larger individual could be a male, and the smaller a female.  Others think that they may be of the same sex, just different ages. 


Figure 5.  The pressure that the foot made in the Laetoli tracks (left), presumed to be A. afarensis, and Homo sapiens sapiens (right) (Ervin 2001).

 


The hominid that left the tracks in the Laetolil beds, states Dr. Leakey, "although much older, relates very closely to the remains found by Dr. Johanson in Ethiopia" (Leaky 1979: 453).  For most anthropologists, the Laetoli tracks were made by the same species as Lucy, based upon the dating.  Leakey disagreed, but both of their finds broke a long-standing assumption that humans developed big brains before walking upright (Anonymous 1998). 

Another person that believes the Laetoli tracks was not made by A. afarensis, but rather by another hominid species.  Dr. Charles Oxnard has used multivariate analysis to reconstruct the foot of an australopithecine (Catchpool 2001).  He believes that the big toe is not in line like a human foot; instead, it sticks out like that of a chimpanzee.  Dr. Oxnard does not think that Lucy or her species belong in the Homo line; he believes that they are just a unique adaptation in the line of evolution.  However, I think there is one flaw to his thinking; he doesn't take into account the dating of the Laetoli tracks, which indeed show that they were made at the time that Lucy lived.   

 

 

Conclusions

 

            When Dr. Donald Johanson found a bone protruding out of the sand on that hot day in Ethiopia, he had no idea of the magnitude of his find.  Lucy has become very important in everyday society; the name Lucy is usually recognized by any common person walking on the street.  The name Lucy can also be found on trivial games and comics.  Twenty-seven years later, the name Lucy still conjures up visions of our early ancestors.

            Did Lucy belong to a species where males and females differed greatly in size and shape?  This question is still a major topic of controversy that surrounds Lucy and her species.  With the available evidence presented by Johanson and his colleagues, I believe that the 'First Family' represents Australopithecus afarensis.  Yes, there are great morphological differences between the two sizes of hominids found there, but some researchers fail to see the differences in the apes of today.  Some of the best examples of this are orangutans and gorillas.  The males are much bigger than females, why couldn't this have been the case with A. afarensis?    

            The vision of Lucy in our minds or in a drawing is usually of her walking upright as we do.  The belief of her walking upright is still a hot controversy among many anthropologists.  Lucy and her species' skeletons can be analyzed forever, yet there will always be controversy about whether she was bipedal or arboreal.  When reading the opinions of many anthropologists. I have realized that most anthropologists do believe that Lucy did indeed walk upright.  Based upon the evidence, A. afarensis is the best candidate for leaving the famed 'Laetoli Tracks'.  There is always the possibility that there was another species that left those tracks, and perhaps some day evidence will be found to prove it.  I agree with Dr. Johanson that Lucy did walk upright, and perhaps did climb trees for certain activities.

            The controversy of Lucy being a male or female is slowly dwindling from anthropological society.  It is hardly talked about, because of the overwhelming evidence that Lucy is indeed female.  Except for an occasional person who believes that Lucy is a male, most anthropologists concede to the idea that she is female, based upon the pelvis.  Even though Lucy's pelvis is not like a humans', but rather in between a human and chimp, it is still clear to most that Lucy's pelvis is bowl-shaped, like that of human females.

            As researchers continue to search for our ancestors, more evidence will be found that could strengthen or weaken our ideas of how Lucy relates to us.  Recent findings have come up with fossilized hominids that seem to be earlier than Lucy, and still be our ancestors.  We do know that A. afarensis did live unchanged for 400,000 years.  With all the faults that we find in Lucy's anatomy, by not being able to as versatile as us, A. afarensis must have been greatly adapted to it's environment for that long.  If Homo sapiens sapiens does live as long as A. afarensis, then I believe that we can scrutinize what anatomical features of Lucy are better or worse than our own.  As stated by Dr. Johanson in 1996, "At 21, Lucy has indeed come of age" (Johanson 1996: 117).

 


Figure 6.  Lucy has become more famous than her discoverer; this comic is just an example.  Drawn by Nick Downes in 1989 (Johanson 1994: 63).

 

 

 

 

 


References Cited

 

 

Aiello, Leslie C.

1994.   "Thumbs Up for Our Early Ancestors".  Science.  265 (1994): 1540-1542.

 

Anonymous.

1998.      "Johanson finds 3.2 million-year-old Lucy 1974".  <http://www.pbs.org/wgbh/aso/databank/entries/do74lu.html >.  A Science Odyssey.   Accessed April 11, 2001.

 

Anonymous.

2001.      "Australopithecus afarensis".  <http://www.robinsonresearch.com/ANTHRO/PHYSICAL/A-afarensis.htm>.  The Robinson Research World of Knowledge.  Accessed May 3, 2001. [Figure 2] 

 

Anonymous.

N.d.   "Lucy".  <http://www.asu.edu/clas/iho/lucy.html>.  Institute of Human Origins, Arizona State University.  Accessed April 11, 2001.

 

Anonymous.

N.d.   Not Titled.  <http://www.gpc.peachnet.edu/~pgore/students/f97/glenda/pelvis.htm >.  Accessed May 3, 2001. [Figure 3]

Brace, C. Loring.

1979.  The Stages of Human Evolution.  Second Edition.  New Jersey: Prentice Hall, Inc. 

 

Catchpool, David. 

2001.  "New Evidence: Lucy Was A Knuckle Walker".  <http://www.answersingenesis.org/docs2/4256news5-5-2000.asp>.  Answers in Genesis Ministries International.   Accessed April 11, 2001.

 

Collard, Mark, and Leslie C. Aiello.

2000.   "From Forelimbs to Two Legs".  Nature 404 (2000): 339-340.

 

Ervin, Stephen Dr.

2001.  "Australopithecus afarensis".  <http://erasmus.biol.csufresno.edu/HE/aafar.html>.  California State University, Fresno.  Accessed May 3, 2001.  [Figures 4 & 5]

Honda, S.

2001.  "Lucy: Australopithecus afarensis Southern Apes of Afar".  <http://www.cabrillo.cc.ca.us/~crsmith/fall99/honda.html>.  Cabrillo University.  Accessed April 11, 2001.

 

Johanson, Donald C. 

1976.  "Ethiopia Yields First 'Family' of Early Man".  National Geographic 150 (1976): 791-811.

 

Johanson, Donald C., and Maitland E. Edey. 

1981.  Lucy: The Beginnings of Humankind.  New York: Warner Books. 

 

Johanson, Donald C., C. Owen Lovejoy, William H. Kimbel, Tim D. White, Steven                                                C.  Ward, Michael E. Bush, Bruce M. Latimer, and Yves Coppens.

1982.   "Morphology of the Pliocene Partial Hominid Skeleton (A.L. 288-1) From the Hadar Formation, Ethiopia".  American Journal of Physical Anthropology.  57 (1982): 403-451.  [Figure 1, pg. 437]

 

Johanson, Donald C., Lenora Johnson, and Blake Edgar.

1994.  Ancestors: In Search for Human Origins.  New York: Villard Books. [Figure 6] 

 

Johanson, Donald C. 

1996.  "Face-to-Face with Lucy's Family".  National Geographic 189 (1996): 96-117.

 

Johanson, Donald C., and Blake Edgar. 

1996.  From Lucy to Language.  New York: Simon & Schuster Editions. 

 

Johanson, Donald C. 

1990.  "Lucy's Knee Joint -2nd Letter".  <http://www.talkorigins.org/faqs/knee-joint/johanson2.html>.  TalkOrigins.org.   Accessed April 12,2001.

 

Kramer, Patricia Ann. 

1999.  "Modeling the Locomotor Energetics of Extinct Hominids".  The Journal of Experimental Biology 202 (1999): 2807-2818.

 

Leakey, Mary D. 

1979.  "Footprints in the Ashes of Time".  National Geographic 155 (1979): 446-457. 

 

Leakey, Meave Dr.

1997-98.   "Australopithecus afarensis".  <http://www.inhandmuseum.com/LA/afarensis/AfarFrame.html>.  Ants Incorporated.  Accessed April 12, 2001.

 

Lieberman, Philip. 

1975.  " On the Evolution of Language: A Unified View".  Primate Functional Morphology and Evolution.  Chicago: Mouton Publishers. 

 

Lincoln, Tim.

1998.        "Lucy Takes A Stroll".  Nature 394 (1998): 325.

 

Richmond, Brian G., and David S. Strait. 

2000.  "Evidence That Humans Evolved from A Knuckle-Walking Ancestor".  Nature 404 (2000): 382-385.

 

Shreeve, James. 

1994.  "'Lucy,' Crucial Early Human Ancestor, Finally Gets a Head".  Science 264 (1994): 34-35.

 

Shreeve, James.

2000.     "Sexing Fossils: A Boy Named Lucy?"  Science 270 (1995): 1297-1298.

 

Stokstad, Erik. 

1999.      "Human Ancestors May Have Knuckle Walked".  Science 287 (2000): 2131-2132.