Leslie C.

Aiello
De@rtment of Anthropology, University College London, Gower Street, London WClE GBT, England

Allometry and the analysis of size and shape in human evolution

This contribution has two main aims. The first is to assess the applicability of allometric techniques for purposes of predication and comparison, and the second is to test the possibility that the relationship between weight and stature was more significantly different in AL 288-l (Australopithecusafarensis) and other Plio-Pleistocene hominids than it is in modern humans. It is argued that in the great majority of cases, the reduced major axis (RMA) is the preferred allometric technique. The reason for this is that the slope of the RMA is totally independent of the correlation coefficient. Both the least squares slope (LSR) and the major axis (MA) (which are both highly affected by the correlation coefficient) converge on it as the correlation coefficient approaches unity. It is also argued that the RMA provides the best estimate of the functional relation between the two variables in the analysis when the error variance is unknown, as is most frequently the case in allometric analyses. Furthermore, it is demonstrated that the MA produces a particularly spurious best-fit line when the correlation coefficient deviates from unity and the variance of the dependent variable (e.g., body weight) is particularly large in relation to the variance ofthe independent variable (e.g., stature). This has important implications not only for general allometric analysis but also particularly for the prediction of body weight from skeletal measurements. When RMA is used as the basis of inference and comparison, AL 288-l (based on the best current estimates for stature and weight) has an inferred weight for its stature that would be highly unusual for modern humans, but similar to that observed for living African apes. OH 62 is similar to AL 288-l in this respect. Journal of Human Euolution (1992) 22, 127-147

Received 8 May 1991 Revision received 28 October 199 1 and accepted 4 November 1991 Keywords:allometry, stature, weight, australopithecines, evolution. line-fitting, human

Introduction
Until recently, However, AL 288-l estimates of early hominid stature, weight and body proportions and limited evidence complete (McHenry, of relatively skeletons have had to such as

be based on rather fragmentary the discovery

1974, 1975, 1976, 1978).

of early fossil hominids

(Austrulopithecus ufarensis) (Johanson

et al., 1982), OH 62 (Homo hub&~) (Johanson

et al., 1982) and KNM-WT longer known ofNeanderthal studying the evolution through to the present day. To date the AL 288-l of 105.5 cm (Schmid, direct measurement recent stature

15000 (Homo cf. erectus) (Brown et al., 1985) together with those and early modern Homo sapiens, has opened up the possibility of period

of stature, weight and body build from the Plio-Pleistocene (Lucy) skeleton has provided reconstruction and articulated the most information

about early

hominid stature and weight. Meticulous of the reconstructed

of the skeleton has inferred a stature 1986a). This estimate is based on skeleton, and is in line with other of the femur, which range

1986; personal comm. in Geissmann, based on inferences

estimates

from the length

between ca. 104-l 13 cm (Olivier, 1976; Feldesmann & Lundy, 1988; Jungers 1988u). The inferred stature of 105 cm is equivalent to the stature of modern Caucasian 4.5 year old boys (Diem, 1962) and lies 6.5 standard deviations (S.D.) below the mean stature of 146 modern (adult) human populations (mean= 161.4 cm; S.D.=8.7) (Eveleth & Tanner, 1976). Body weight estimates for AL 288-l range between ca. 25-32 kg (Jungers, 19886; McHenry, 1988, 199 1). The most recent of these estimates, based on the size of the hindlimb
004772484/92/020127+21 $03.00/O 0 1992 Academic Press Limited

128 joints (McHenry, tive sample) between These

L. C. AIELLO

199 1)) give weights of 26.7 kg (on the basis of a modern human comparaape comparative (adult) sample). human These esti1962) and lie populations to modern Caucasian 8-10 year old boys (Diem, the mean weight for modern

and 31.8 kg (on the basis of an African below

mated weights are equivalent 3.1-3b7S.D. (mean = 57.2 kg; SD. =8.3).

data suggest that AL 288-l

might have been much shorter

than adult modern

humans, but not as correspondingly that the relationship

light. The purpose of this paper is to test the possibility different in AL 288- 1 than it ofAL 288-l as and comparison.

between weight and stature was significantly ofallometric techniques

is in modern humans. In so doing it will use the weight for stature relationships a test case to assess the applicability for predication

Methods
No modern human populations have mean statures as short, or mean weights as light, as & Tanner, 1976). In order to assess the weight for those estimated for AL 288-l (Eveleth

stature relationships of this hominid it is necessary to project the mean weight for stature relationship of modern humans into the size range of the fossil. This is done by placing a bestfit line through modern human stature and weight data and extending range. The amount by which AL 288-l deviates it into the short size to the of the deviation used statistical (or principal) each one of from this line can be compared

amount modern humans deviate from the line and from this the significance determined. The main problem in this type of analysis is that there are three commonly ways of fitting axis (MA) AL 288-l these techniques a line to the data: and reduced least squares regression analysis. (LSR), major analysis, major axis (RMA)

Unfortunately, LSR

fits a different line (Figure to modern humans.

1) and gives a different result for the position of In this example, produces a line with a the weight for stature

in comparison

lower slope than RMA, relationship Table of the fossil.

which produces a line with a lower slope than MA. The lower the

slope of the best-fit line, the closer it lies to the point representing 1 gives the deviations accepted of AL 288-l

from each of these three best-fit lines for each of in these measures (McHenry, 1988, 1991; of 146

nine weight and stature combinations. cover the currently Jungers, modern human areas oforigin 61-62, 77-78, 1988a,b; Feldesman populations 93-94). & Lundy,

The three weight estimates and three stature estimates 1988). The reference population n = 237) (Eveleth & Tanner, deviation is comprised

range of variation representing

all of the major ethnic groups and geographical 1976, Tables 29,44-45, transdevion the basis of logarithmically units. The standard

(sexes entered separately, Deviations

were determined

formed data and are presented

in the form of standard

ation unit (SDU) for each individual point is determined by the distance of that point from the best-fit line (measured in the direction of they-axis) divided by the S.D. of the distances of all of the points in the human reference sample from the best-fit line. So, for example, a SDU of 2 means that the case lies 2 S.D. from the best-fit line. Figure 2 graphically illustrates the magnitude of these deviations. If the LSR line is used as the best-fit line (and if AL 288-l is considered to represent the mean weight and the mean stature for its population) AL 288-l falls within 3 S.D. of the modern human sample for five out of the nine weight and stature combinations. However, ifthe RMA is used as the best-fit line, it falls outside the 3 S.D. limit in eight out of the nine weight and stature combinations and if the MA is used it falls outside the 3 S.D. limit for all nine of the weight and stature combinations.

SIZE AND SHAPE IN HUMAN EVOLUTION

129

l-00 I 2.00

2.06

2.12 Log,, stature

2.18 (cm)

2.24

reduced major axis slope (RMA) and least squares regression slope (LSR) for the relationship between stature (S; cm) and body weight (BW; kg) for 146 modern human populations. Sexes are considered separately (n = 237) and the slopes are computed on the basis oflog (base 10) transformed data. MA: Log,$W=3~251og,,S-5.41; RMA: log,,BW=2.691og,,S-4.18; LSR: log,,BW=2~12log,,S-2.93. Modern human populations are indicated by p). The lines marking the position ofAL 288-l indicate a stature range 100-l 10 cm and a weight range 26.7-31.8 kg.
Figure 1. The major axis slope (MA),

Table 1

Deviations in SDU

of Al

288-l from the best-fit line Weight (kg)

Stature (cm)
100

Best-fit method RMA MA LSR RMA MA LSR RMA MA LSR major

26.7 5.60 7.34 2.90 4.21 5.88 1.74 2.89 4.49 0.64 axis, MA = major

29.9 6.79 8.38 4.16 5.41 6.92 3.01 4.09 5.53 1.90

31.8 7.44 8.95 4.85 6.06 7.49 3.69 4.74 6.10 2.59

105

110

RMA = reduced re,gression.

axis, LSR = least squares

This comparison raises two major questions. Firstly, which line-fitting technique is the most appropriate technique to use as the basis for comparison, and secondly, which of the weight and stature estimates can be relied upon to be the most accurate fossil? estimates for this

130

L. C. AIELLO

MA

RMA

LSR

100 cm
26.7 kg

100 cm 29-9 kg

100 cm 31.8 kg [Stature

105 cm 2%.7 kg (cmVweight

105 cm 29.9 kg

105 cm 31.8 kg

IlOcm 26.7 kg

IlOcm 29.9 kg

IlOcm 31.8 kg

(kg) combinations]

Figure 2. The distance in standard deviation units (SDU) that AL 288-l lies from the LSR, RMA, and MA slopes defined in Figure 1. SDU distances are presented for nine stature (cm)/weight (kg) combinations. (0) indicate the cases that fall within 3 SDU of the modern human best-fit line characterizing the relationship between stature and body weight and (0) indicate the cases that fall outside this boundary.

The appropriate line-jitting technique The most appropriate difference mation stature (Hofman of AL 288-l line-fitting relation technique (Rayner, humans. & Barbour, relation for the purpose of assessing the similarity 1985; Page1 & Harvey The functional relation 1989)) or underlying can never be directly 1988, 1989a) or from modern humans is that technique which gives the best approxibetween is the ideal underlying proportional relationship

of the functional and weight relationship

in modern (Martin

biological

et al., 1986) characterizing the functional

the covariation

of stature and weight in human popuobserved and must be esti-

lations. However,

mated from a sample of data drawn from the total human population. Rayner (1985) emphasizes that the central tendency of the sample may not necessarily correspond to the functional gathering relation in the population because of both the observational in the selection of the sample. errors inherent in the data and the randomness

When comparison is the main purpose of analysis, as it is in this case, the most appropriate line-fitting technique is that technique which best represents the central tendency in the sample and provides the best estimate of the functional relation in the population. In order to decide which of the three line-fitting techniques meet both of these criteria and is therefore most appropriate, it is necessary to understand two points. Firstly, why do the three techniques produce three different best-fit lines for a given data set, and secondly, what theoretical considerations govern the choice of the most appropriate line-fitting technique? Why do the best-@ lines vary? LSR, MA and RMA produce different best-fit lines for the same data set simply because they respond in different ways to two major aspects of the data to which the line is being fit. The first of these aspects is the sample correlation coefficient, and the second is the ratio of the variances of they variable and the x variable (s*y/s*~) (Seim & Sather, 1983).

SIZE AND SHAPE IN HUMAN EVOLUTION

131 unaffected by the

The RMA is unique among the three techniques correlation deviation coefficient of they variable and the x variable

because it is completely

of the reference sample. The RMA is simply the ratio of the standard (sy/sx). As long as the ratio of the variances ofwhether the to produce the the slope of the RMA will be the same irrespective

(s*y/s*x) remains constant, data are highly correlated the RMA covariance

or not [Figure 3 (a)]. This occurs because the equation (SXJ) which is part of the formula

(sy/sx) is simply the square root ofthe variance ratio (s2y/s2x). It does not include the of the x and y variables for computing

correlation coefficient (Table 2). The specific values of both the MA and the LSR variance ratio but also on the correlation covariance Figure variance ofthese variables 3(b) illustrates coefficient.

slopes are dependent

not only on the but also the

This is because the formulae from which of the x and y variables

these slopes are derived include not only the variances variance ratio and the correlation ratio and the correlation

(Table 2). The MA and the LSR slopes do, however, react to the coefficient in very different ways. between the value of the slope of the MA, the of coefficient. The value of the MA is only independent

the relationship

the correlation correlation coefficient,

when the variance ratio is unity. In this case, the slope ofthe MA is identical to Below unity, the lower the value of the variance ratio, the more the holds.

the slope of the RMA.

coefficient will affect the value of the MA. In these cases, the lower the correlation the lower will be the slope of the MA. Above unity a different relationship coefficient, the higher will be the slope of the MA.

In these cases, the lower the correlation

This effect becomes more and more pronounced as the variance ratio increases. However, as long as thevariance ratio differs from unity, the higher the correlation coefficient the more the MA approaches the RMA. Figure 3(c) illustrates the relationship between the LSR slope, the variance ratio and the correlation coefficient. The higher the correlation coefficient, the closer the LSR slope Cy= dependent variable) is to the RMA irrespective of the variance ratio. It is also clear that the coefficient depresses the larger the value of the variance The similarities and differences are summarized in Figures ratio, the more a lower correlation

LSR slope in relation to the RMA slope. between the LSR slope, the MA slope and the RMA slope plots of stature and 1 & 4. These figures show three allometric

body weight, each with a different variance ratio. When the log ofweight (y-axis) is plotted against the log of stature (x-axis) the variance ratio is high (ratio= 7.23) (Figure 1) and the MA has a steeper slope than the RMA, which has a steeper slope than the LSR. When the log of the cube root of body weight (j-axis) is plotted against the log of stature (x-axis), the variance ratio is closer to unity (ratio = 0.80) [Fig. 4(a)] and the value of the MA is similar to that of the RMA. (ratio = 0.13) the RMA. Both techniques produce steeper slopes than does LSR. When the log of stature (y-axis) is plotted against the log ofbody weight (x-axis), the variance ratio is very low [Fig. 4(b)] and the MA approximates the LSR slope; both are shallower than different from each other is dependent

The point at which the slopes become significantly

not only on the correlation coefficient and the variance ratio, but also on the sample size. The smaller the sample size the greater will be the confidence limits for the MA and the LSR slopes, and the greater the difference in the values of the slopes before they become significantly different from each other or from the RMA. Figure 5 illustrates this for two separate variance ratios (0.2 and 3.0) at three different correlation coefficients (r= 0.5, r= O-7, r= O-9) for sample sizes of 12-500. It is important to recognize that these are not extreme examples. A variance ratio of O-2 corresponds to a RMA slope of 0.45 and a ratio of 3-O corresponds to a

32

L. C. AIELLO

0.2 (b)

0.4

0.6

0.8

l-2

I.4

I.6

I.8

2.5

0.2

0.4

0.6

0.8

I.2

i-4

I.6

I.8

2.5

0.2

0.4

0.6

0.8

I.4 Variance ratio

l-6

I.8

2.5

Figure 3. (a) The value of the RMA slope for 12 different ratios between the variance of they variable and the variance of the x variable. For each separate variance ratio the narrow bar on the left-hand side represents a correlation coefficient (I) of0.99, for the adjacent bar to the right T= 0.90, for the next adjacent bar T= 0.80, and so on until r= 0.30. Note that the RMA is unaffected by the value of r. (b) The value of the MA slope for twelve different variance ratios. For each separate variance ratio the narrow bar on the lefthand side represents the value ofthe RMA slope. The adjacent bar to the right represents the value of the MA when r = 0.90 and the next adjacent bar the value of the MA when r= 0.80. The right-hand bar for each variance ratio represents the value of the MA when r= 0.30. Note that the only time that the value of the MA is unaffected by the value ofr and equal to the RMA is when the variance ratio is one. (c)The value ofthe LSR slope Cy on X) for 12 different variance ratios. The narrow bars for each variance ratio represent the RMA and different correlation coefficients as described for (b) above.

SIZE AND SHAPE

IN HUMAN EVOLUTION

133

Table 2

Formulae for determinin g the best-fit lines and the product moment correlation coefficient1 RMA LSR3 MA JYIJ sqqs*x D=[(s2x+s~y)-4(s~sy-s?ry)ll? L= (sx+s*y+D)/2 MA=sxy/(L-s*y) sxy/ (sxS4)7

sy = standard deviation ofy, sx = standard deviation of x, sxy = covariance ofs andy, sy = variance ofy, s2x = variance ofx, s2ny= square of the covariance ofx andy, RMA=reduced major axis, LSR=least squaresregession, MA= major axis, r=product moment correlation coefficient. See Sokal and Rohlf (1981) for further discussion and derivation of these equations. ?The RMA is equivalent to the MA computed on standardized data. This is the least squares equation wheny is the dependent variable Cyon x). If x is the dependent variable the equation is sxy/s3.

slope of 1.73. The range of correlation biological data.

coefficients

is commonly

found when dealing with

What theoretical considerations should gouern the choice of technique? The best-fit lines produced correlation theoretical coefficient by LSR, RMA and MA respond in different ways to the sample (s2y/s2x), because they employ different 198 1; Harvey & Mace, 1982; Rayner, the and the ratio of the variances

criteria for fitting the lines (Sokal & Rohlf, & Barbour,

1985; Martin perpendicular the horizontal horizontal

1989). Very simply expressed, LSR fits its line by minimizing

vertical distance between each data point and that line (Figure 6). The MA minimizes the distance between each data point and the line and the RMA and the vertical only minimizes distance, 1981). distances [or the area of the triangle minimizes both of the comprised

and vertical distances and the best-fit line (Figure S)]. the vertical distance between each point and the line, and it is called an asymmetric, of LSR or Model I, regression technique in the variable is that there is only variation

Because LSR (Sokal & Rohlf,

ignores the horizontal dependent variable

A basic assumption

(they variable in the vertical plane) and not in the independent plane). An important

(the x variable in the horizontal the independent variable

aspect of LSR is that a second line can distance between the

be fitted to the data (Figure 7). If the x variable is assumed to be the dependent variable andy (x any), LSR then minimizes the horizontal data points and the line. In a LSR analysis these two best-fit lines cross at a point representing the mean ofthe x and the mean of they variables. The degree to which these lines diverge from each other is inversely proportional to the correlation coefficient (Sokal & Rohlf, 1981). The lower the correlation coefficient the more the lines diverge, and the higher the correlation coefficient the more the lines converge. An interesting point is that the RMA is simply the geometrical mean of the LSR Cyon X) and the inverse (1 /slope) of the slope of the LSR (X ony) (Hofman et al., 1986). Both MA and RMA analysis assume that there is variation not only in the dependent variable but also in the independent variable. As a result they are described as symmetrical, or Model II, regression techniques (Sokal & Rohlf, 1981). In both cases only one line can be fit to a cloud of points. If the variables are reversed and x is taken as the dependent variable

34

L. C. AIELLO

0.42

o.362M 230r
2.25 t

2.12 Log,o stature

2-18 (cm)

2.24

RMA LMSAR

2.05

I:
AL 288-l I.55 I.70 Log,, body weight (kg)

1.40

I .85

2.;0

Figure 4. (a) The MA, RMA and LSR slopes for the relationship between stature (cm) and the cube root of body weight (kg) in human populations. The variance ratio=0.80. MA: log,,BW =0.871og,,S1.34; RMA: log,,BW=O~9Olog,,S1.39; LSR: log,,BW=0,7Ilog,,S-0.97. (b) The MA, RMAand LSR slopes for the relationship between body weight (kg) and stature (cm) in human populations. Note that in this comparison body weight is the independent variable and not the dependent variable. The variance ratio = 0.14. MA: log,,S=0.3llog,,BW+ 1.67; RMA: log,,S=0.371og,,BW+ 1.56; LSR: log,,S=O~291og,,BW+ 169. Symbols as in Figure 1.

SIZE AND SHAPE

IN HUMAN EVOLUTION

135

andy as the independent

variable

(X onr), the slope ofthe line is simply the inverse of the slope techniques (RMA or

of the line when the dependency is reversed Cyon X) . There is general agreement in the literature that the symmetrical MA) are more appropriate techniques than LSR, both to determine

the central tendency in

the sample and to estimate the functional relation in the population (e.g., Rayner, 1985). However, there is considerable disagreement over which of the symmetrical techniques is the most appropriate [cf. Martin & Barbour (1989) who favour MA and Rayner difference (1985) who favours RMA]. The previous discussion the correlation coefficient. coefficient,

has shown that the fundamental

between

MA and

RMA is that the slope ofthe MA is affected by not only the variance ratio (s~/sx) but also by while the slope ofthe RMA is totally unaffected (e.g., when the variance by the correlation Under certain data conditions ratio is low) the slope of

the MA approximates the slope of the LSR ly on X) line. Under other data conditions (e.g., when the variance ratio is high and the correlation low) the MA has much steeper slope than either the LSR Cyon X) or the RMA and in these cases approximates the second LSR slope (X approaches unity do ony) (see for example Figure 7). Only when the correlation coefficient

the MA and the LSR slopes (bothy on x and x any) converge on the RMA. Ifthe LSR slope is rejected as an appropriate technique for estimating the central tendency ofa bivariate sample because its two slopes (bothy on x and x ony) only converge on each other (and on the RMA) to produce a single bisecting questioned approximate line when the correlation coefficient is unity, the MA must be and produces a single on similar criteria. Although the LSRg it is a symmetrical technique

line for the data set (MA fory on x is the inverse of the MA for x any), the value of this slope can on x and x ong. The only time it produces a clearly bisecting slope for coefficient is unity (and all the techniques result in an a data set is when the correlation

identical slope value) or when the variance ratio is one (when the value of the MA is identical to the RMA and both of these are equivalent The RMA correlation is always the geometrical coefficient) unity. Therefore, coefficients). to the geometrical mean ofthe two LSR slopes). slopes (irrespective of the on it as the correlation mean of the two LSR slopes converge

and the MA and LSR

coefficient approaches ratios and correlation the population no.

it is the only one ofthe slopes that can be consistently the question still remains whether functional the RMA relation in

relied upon to produce the best estimate ofthe central tendency ofthe sample (for all variance However, can also be relied upon to produce the best estimate of the underlying

from which the sample was drawn. The answer to this question is both yes and

The answer is a definite yes if, as is generally the case, the error variances in the data are unknown. The error variances are part of the residual scatter of points around the best-fit line. The other part of this residual scatter is biological scatter that reflects growth irregularities ever reason from the central tendency. individuals, the error variance In an intra-specific scatter (Kuhry & Marcus, 1977) or of the individual, or individual deviations for whatanalysis where the data points are

is the residual scatter of points around the best-fit line due to

measurement error (Rayner, 1985). I n an inter-specific analysis where the data points are species means; the error variance is made up of within species variation and measurement error. In both intra- and inter-specific analyses, the error variances are generally small in relation to the biological scatter and are not usually known for the type of data commonly employed in allometric analysis (but see Page1 & Harvey, 1988, 19896). The RMA assumes that these unknown error variances are proportional to the variances of the x ony variables. Rayner

36

L. C. AIELLO

r .............-_~..~
..........................,...
________----L-L~=.-,~.;-

. ,,

------G
1

53 8

8 -

-------iW .? 0

......... ................ .
N
II

...

............

z
6
0

0
9 p

OF

;s

6 .-

5
0

-+-+++4 0
m

F:

..... ..........p? Id I
----_-_----______+.L

... ...

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i5

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SIZE

AND

SHAPE

IN HUMAN

EVOLUTION

137

; ,: ..: ..,............ . . . . . . k.. .. . . .. . . . . . . . . . . . . . . . . . . . , . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . .

. . . . . . . . . . . .._

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.-2%...-_1.a.___

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az!s alduos

138

L. C. AIELLO

x Variable

Figure 6. Geometrical interpretation ofthe criteria employed by MA, RMA and LSR to fit a line to a data set. MA minimizes the squared perpendicular distance between each data point and that line. RMA minimizes the area of the triangle comprised by the horizontal and vertical distances between each data point and that line. LSR Cyon x) minimizes the squared vertical distance between each point and the line and ignores the horizontal distance. LSR (x any) minimizes the squared horizontal distance between each point and the line. The circles or ellipses correspond to the assumptions about the error variances made by each technique. MA assumes that the error variances of the x andy variables are equal. LSR Iy on x) assumes that x is sampled without error and that only they variable therefore has an error variance (represented by a very elongated ellipse). LSR (x any) assumes the reverse ofLSR Iy on x) (represented by an elongated ellipse in the plane ofx). RMA assumes that the ratio of the error variances is proportional to the ratio of the variances of they and x variables (after Rayner, 1985).

(1985) has demonstrated

that when the error variances are unknown, relation in the population

the RMA provides the

least biased estimate ofthe functional 1985). However, variances variances literature),

and should be used instead of

the MA, which assumes that the ratio is 1, or the LSR, the RMA

which assumes that it is zero (Rayner, the functional technique relation if the error

should not be used to approximate a more precise, but less known

of the x ony variables are known the general structural

are known. In the relatively relation,

rare cases where these error (in the anthropological 1966;

can be employed to fit the line (e.g., Sprent,

Rayner, 1985; Page1 & Harvey, 1989a). There is another point in favour of the use of the RMA over the MA. The RMA is totally unaffected by changes in the scale of the x or they variable (Rayner, 1985). This effect is illustrated in Table 3 which gives the deviations of AL 288-l (in SDU) from both the RMA and the MA slopes for two separate stature/weight analyses. The RMA produces an identical result for the relationship between stature and the cube root of body weight as it does for the relationship between stature and body weight. The results based on the MA are totally different for these two analyses even though the two analyses differ only in the scale of they variable, body weight. Any conclusions based on the MA slope with reference to the similarity or difference ofAL 288-l to modern human populations would therefore depend on the

SIZE AND SHAPE

IN HUMAN EVOLUTION

139

LSR x ony MA RMA LSRyonx

2.18 Log,, stature (cm)

2.24

2f30

Figure 7. For any data set the slope ofthe RMA is the geometrical mean ofthe slope ofLSR Cy on x) and the inverse of the slope of LSR (x one). For this particular example the slope of the RMA= 2.690 (. . .), LSR (J on x) = 2.120 (--) and LSR (x any) = 0.293 (--). Note that the MA ( = 3.246) (---) approaches the value of the inverse of the LSR (x ony) ( = 3.413).

Table 3

Deviations (in SDU) of AL WI-1 from the RMA and the MA based on two separate weight for stature analyses Stature (cm) 105 Weight (kg) 26.7 29.9 31.8 26.7 29.9 31.8 RMA deviation 4.21 5.41 6.06 4.21 5.41 6.06 MA deviation 5.88 6.92 7.49 3.91 5.12 5.78

Analysis Stature us. body weight Stature us. cube root of body weight

105

SDU are computed on the basis oflog,, transformed data. Variance ratios: log,,BW/log,,S = 7.229,log,,(BW) 3/log,,S= 0.803.

scale of the values used in the analysis and not solely on the differences

in inferred stature

and weight between the fossil and the reference sample. The reason this happens is because the MA slope is affected by the variance ratio and the variance ratio changes as the scale of they variable changes (body weight has a higher variance than does the cube root of body weight). The RMA has been supported as the best estimator of the functional relation by a number of authors in both the statistical and applied literature (Kermack & Haldane, 1950; Imbrie, 1956; Sprent, 1969; Ricker, 1973, 1984; Rayner, 1985; Hofman et al., 1986). In light of this support and of the strengths of the RMA technique outlined above, it is perhaps surprising

140 that this technique authors interested Martin & Barbour, & Lundy, assumption necessarily functional Particularly

L. C. AIELLO

has been dismissed in favour of MA in the anthropological literature by both in the theoretical aspects of line-fitting (Page1 & Harvey, 1988; 1989) and in line-fitting 1988). This as an analytical misconception the central tool (Olivier, 1976; Feldesman technique it appears to have arisen from the regression of the sample and the coefficient

1988; McHenry, that because produces

the MA is a Model

II, or symmetrical, tendency

a line that approximates where the variance

relation of the population. in circumstances

The above analysis has shown that this is not the case. ratio is high and the correlation

deviates from unity, the MA can deviate markedly from both the central tendency and from the functional relation. In summary, there is little to support the use of the MA over the RMA as a technique to estimate the underlying functional relation of bivariate data. The only time that it can be relied upon is under specific circumstances and/or when the variance would be identical when the correlation coefficient approaches unity ratio approaches one. In these cases the value of the MA slope slope for a given data set and there seems to be & Barbour, 1989).

to the value of the RMA

little reason not to employ the RMA

in the first place (contra Martin

The weight for stature relationship of AL 288-l

In testing the hypothesis that the relationship between weight and stature was significantly different in AL 288- 1 than it is in modern humans, we are asking if AL 288- 1 is significantly different from the functional relation ofweight and stature in modern humans. In this case, the RMA is the appropriate technique to use as the basis ofcomparison. This is particularly so because the variance the correlation However, predicted differ considerably tions for AL 288-l ratio in weight for stature comparisons is very high (ratio = 7.23) while coefficient is relatively low (r = 0.79). Both LSR and MA result in slopes that from the central tendency. question of whether the weight and stature predicare normally techniques disto by using one of the three line-fitting are accurate. Both weights and statures for the hominids

there is still the unanswered

from skeletal measurements reliability

cussed in the previous section. Therefore, the assessment of the probable stance in which the RMA

much of the previous discussion is also applicable of these inferred weights and statures. however, line-fitting

The one point that needs to be stressed in this context, may not be an appropriate

is that the only circumtechnique is when the

specific purpose of the analysis is prediction rather than the determination of the underlying functional relation in the data. In the specific cases where, for example, weights or statures are to be predicted from a reference sample, LSR might be the appropriate technique. The assumptions of LSR require that the independent variable (x) be sampled without error (error or biological scatter). This is rarely if ever the case with biological data. However Martin & Barbour (1989) have suggested that LSR can be statistically justified if the purpose of the analysis is prediction and if the data has a bivariate normal distribution (both the x variable and they variable are normally distributed). They suggest that most intra-specific data would have such a bivariate distribution but also stress that it would be highly unlikely that inter-specific data would be so distributed. Martin & Barbour (1989) recommend that LSR should be used for prediction when the reference sample is an intra-specific data set. However, if inter-specific data is to be used as the -basis of estimating weight or stature, or if there is reason to believe that the intra-specific data does not have a bivariate normal distribution, LSR should definitely not be used. In

SIZE AND SHAPE IN HUMAN EVOLUTION

141

2
E z

5432IO-

100

105

I IO

115

120

-L 125

I30

LSR predicted statures

L
135 140 145

Figure 8. The range of predicted statures for AL 288-l. Predictions are based on the length of the femur (28.1 cm) and are derived on the basis of a variety of LSR equations (Geissmann, 1986a; Feldesman &

Lundy, 1988;Jungers, 1988a). RMA is the preferred technique

these

cases,

the foregoing & Barbour

analysis (1989)

has demonstrated who recommend

that MA].

[contra

Martin

Rather

than strictly predicting of prediction

weight, a different and more general question is asked of the data. This more general question is simply, for example, what the weight for stature would be if the object corresponded to the functional relation in the reference population. It should also be stressed that even if the assumptions size range of the reference population. the more general question asked. AL 288-l staturepredictions Stature is normally predicted from the length of the long bones, and particularly, from the length of the bones of the lower limb. However, where relatively complete skeletons are the of LSR technique should only be used for prediction if the object ofprediction

are met by the data, this falls within the general

If it falls outside this range, RMA should be used and

basis of analysis, stature can also be estimated from direct measurement of the reconstructed and articulated skeleton. For AL 288-1, direct measurement has resulted in an estimated living stature of 105.5 cm (Schmid, long bone, particularly result, stature 1986; pers. comm. in Geissmann, 1986a). This estimate based on inference from skeleton is possible. As a based on LSR can act as a reference against which to assess stature measurements femur, length. of an articulated inferred from the length of the femur It is rarely the case that direct measurement is most commonly equations. prediction

From the foregoing discussion, however, LSR is only an appropriate technique for if the reference population has a bivariate normal distribution and is of the same

general size as the specimen which is the object of prediction. For stature measurements we should also add that the specimen should be of the same body proportions as the reference population (Geissmann, 19866). Figure 8 illustrates the range of statures that can be predicted for AL 288-l (femur length = 28.1 cm). This figure is based on LSR stature predictions derived on the basis of 40

142
Table 4 Predicted stature&xIq).for Data type Raw Raw Log Log Sample

L. C. AIELLO

AL 288-l LSR 108.9 110.9 108.1 110.3 RMA 106.3 109.4 105.8 109.3 MA 103.9 108.1 105.8 109.0

Human Pygmies Pygmy Chimpanzees Human Pygmies Pygmy Chimpanzees

After Jungers, 1988a (Table 1). LSR =least squares regression, RMA= major axis.

reduced major axis and MA=

different

human

populations

(Geissmann,

1986a;

Feldesman

& Lundy,

1988; Jungers,

1988a) and one population of pygmy chimpanzees (Jungers, 1988a). Only five of these LSR predictions fall in the range 100-l 10 cm suggested by the reconstructed and articulated skeleton. All of these five predictions are based (Jungers, on modern human are based on small-bodied populations that are close to the inferred size range of AL 288- 1. Two of the predictions pygmies, 1988) and one on Hindus ( 105.9) (Rosing,

( 108.1 and 108.9) Qungers, 1988) two on pygmy chimpanzees ( 1 IO.3 and 110.5)
1983). All five fall in the upper halfofthe

stature range for AL 288-l suggested by the reconstructed skeleton. However, for these stature predictions to be accurate we have to assume that the reference populations the stature populations. chimpanzee have a bivariate of AL 288-l Table population normal distribution. based on the RMA Because this is unknown it is also appropriand the more general relation population question of what to the functional of the reference and the pygmy of South African ate to look at the predictions

would be if it adhered Uungers, 1988) RMA

4 demonstrates

that for the pygmy human predications

statures for AL 288-l

also fall in the range

105-l 10 cm. This is also true for RMA

based on a population

Blacks (106 cm) (Feldesman & Lundy, 1988). It is perhaps significant that none of these analyses produce statures that are in the range loo-105 cm. However, it is also important to emphasize that RMA statures based on other, and particularly not necessarily equations North American produce Blacks inferred statures in the 105-l (1952) produce derived from the Trotter & Gleser taller, human populations For example, (ranging do 10 cm range. RMA from

data for femur length and stature in taller stature estimates

and Caucasians

117 cm based on Caucasian females to 12 1 cm based on Black males). The reason that some degree of confidence can be placed in the stature estimates ranging between 105 and 110 cm is that firstly, they are based on the most appropriate secondly, they are based on the shortest populations predictive techniques, for which data is available,

and thirdly, consistent results are produced from populations with very different body proportions (pygmy chimpanzees and pygmy humans) (Jungers, 1988a). Both of the last points are very important in matching AL 288-l to the reference populations which most closely approximate the fossil, not only in its overall small size but also in its ambiguous body proportions. Although it is definitely human-like in its bipedal locomotion, there is considerable evidence which suggests that its body proportions and approximated those of the pygmy chimpanzee differed from those ofmodern & Stern, humans 1984; (Jungers 1983; Zihlman,

Jungers, 1990). In a theoretical context mention should also be made of the difference in the estimated statures produced from raw or logged data (Table 4). When raw data are used there is a high

SIZE AND SHAPE

IN HUMAN EVOLUTION

143 variable Black and

variance Caucasian

ratio between

the dependent

variable

(stature)

and the independent North American

(femur length).

Based on data from Trotter

& Gleser

(1952,

males and females) and from Feldesman

& Lundy (1988, South African Blacks)

the ratio ranges from 7.55 (North American Black males) to 10.19 (South African Black females). These are relatively high variance ratios and they explain why, when raw data is used, the inferred statures for short individuals those based on the RMA or LSR equation. sample. When predictions changes the variance based on the MA are always shorter than High variance ratios result in steep slopes for the the

MA producing short stature estimates for individuals that fall below the mean ofthe reference are based on logged data for the same reference populations, to the RMA stature. This is because, unity. (However, MA based stature is equivalent in this case, logging it is import-

ratio to the point where it approximates

ant to realize that logging does not always cause the variance ratio to approach unity. In the case of the relationship between weight and stature, the variance ratio of the raw data approaches unity while the variance ratio of the logged data is high.)

AL 288-l weight predictions As is the case for estimated between skeletal parameters, stature, estimated weight must be based on the relationship surface size, and body weight in such as bone area or joint

modern reference samples. Accuracy can only be assessed on the basis of suitability of the reference sample, technique ofinference (line-fitting technique) and suitability ofthe skeletal measure used as the predictor. McHenry (1991) has recently argued that the preferred sample) weight for AL 288-l would be between 26.7 kg (based on an human comparative

and 3 1.8 kg (based on an ape comparative

sample). These estimates are made from hindlimb McHenry uses LSR for prediction, of the variance ratio. the

joint surface sizes (femoral head, distal femur, proximal tibia and distal tibia), the mean ofthe four analyses giving the estimated weights. Although correlation McHenry locomotion, coefficients (1991) involved would make little difference are so high in his comparisons (r=0.97-0.99) that the technique

to the results irrespective

favours the human reference

sample (26.7 kg) on the basis of bipedal

which would logically put more stress on the hindlimb joints than found in the but very low

apes and support the use of the human regression line. He supports this by arguing that the forelimb joints also suggest weights of 27-30 kg based on human comparisons, weights of 11-l 7 kg based on ape comparisons. However, Jungers (19886) has argued that on

the basis of the relatively short legs in AL 288-l and its small lumbar sacral joints, that it is half-way between humans and apes in its proportions. As a result he prefers an ape comparative sample, but the same arguments could be used to suggest that the mean of the estimates based on the ape and the human comparative samples should be used. These data [together with Jungers (19883) estimate of 30.4 kg based on the multiple regression of six joint surface areas in a reference sample based on six ape species and McHenrys estimated weight lies in the neighbourhood [based on the McHenry Weightfor earlier ( 1988) estimate of 1.8 kg 29.9 kg based on LSR ofthe area ofthe proximal femur shaft] suggests that the most accurate of 30 kg, with a possible range of ca. 26.7-3

( 199 1) predictions].
AL 288-l

stature relationshipsfor

Based on a stature estimate of 105-l 10 cm and a weight estimate of 26.7-31.8 kg, AL 288-l lies between + 2.94 (stature = 110 cm; weight = 26.7 kg) and + 6.12 (stature = 105 cm; weight = 3 1.8 kg) SDU above the human RMA. Because only 0.41 o/0of modern human populations would be expected to lie beyond the lowest of these figures ( + 2.94 SDU), there

144

L. C. AIELLO

-z
i
m 3 =; :: 52

KNM-WT 15000

I.OOl
2.00

2.05

2.10

2.15 Loglostoture (cm)

2.20

2.25

2.30

Figure 9. The relationship between stature and body weight in modern humans, wild collected African apes (n=20 individuals), AL 288-l and WT 15 000. Statures and weights for Panpaniscus are after Coolidge & Shea (1982) and for the other African apes from the records of the Powell Cotton Museum, Birchington, Kent. ( n) Modern humans; ( A) juvenile gorillas; ( 0) adult gorillas; (0) Pan troglodytes; (0) Panpaniscus. RMA for juvenile and adult African apes: log,,BW =4.0741og,,S -6.786. The lengths of the lines indicating the position ofAL 288- 1 represent the range ofinferred statures and weights for this fossil considered appropriate in this analysis (see text). The two points for KNM-WT 15 000 (0) represent the estimated stature (160 cm) and weight (48 kg) for this juvenile fossil and its projected adult stature (185 cm) and weight (68 kg) (Ruff, 1991).

can be little doubt that AL 288- 1 was unusual in its weight for stature. The unusual position of AL 288-l is underscored by its relation to African (Bayenga) pygmy populations. and -0.10 population, (Bunia) Two groups of SDU from the Of female pygmies from the Congo lie +0.78 modern human RMA, Guinea between the 56 populations

while a third very small-bodied or populations ofAfrican

Simbai females from New

(mean stature = 137 cm; mean weight = 38.9 kg) lie + 0.64 SDU from the RMA. ofAfricans, + 1.0 and -3.0 SDU from the RMA.

ancestry in the sample, 53 ofthese lie and urban females from by its

The three heavier groups are Nigerian urban

females from Lagos

Durban, The unusual position ofAL

( + 1.20 SDU) and Ibadan South Africa ( + 3.3 1 SDU).

288-l

( + 1.61 SDU),

in relation to modern humans is also underscored

similarity to the African apes (Figure 9). Based on the RMA computed for a sample of20 wild collected chimpanzees, pygmy chimpanzees and gorillas, AL 288- 1 lies between + 0.856 and - 1.648 SDU from the African ape RMA. It is similar in its absolute stature and weight to juvenile gorillas in this sample. Not only is AL 288-l similar to living apes in its inferred weight for stature but there is also evidence that AL 288-l approximated to living apes in general body proportions and in possibly being more heavily muscled than modern humans. Jungers & Stern (1983) have emphasized the ape-like short legs of AL 288-l and there is also evidence from the hand and shoulder that it had a more well developed upper limb musculature than is commonly found in modern humans (Aiello & Dean, 1990). Furthermore, if Stern & Susman (1983) are correct in their argument that AL 288-l walked with a flexed knee and hip, it would require more strongly constructed hamstrings and quadriceps to support the flexed joints than is the case in humans.

SIZE

AND

SHAPE

IN HUMAN

EVOLUTION

145

Weight

for stature

relationships

of other early hominids

Because of the fragmentary nature of the postcranial skeletons of the majority of other early hominids, there are even more uncertainties in the reconstruction of their weight for stature relationships than there are for AL 288- 1. However, (Johanson the overall similarity of the OH 62 Indeed, if skeleton to that ofAL 288-l

et al., 1987) suggests that it too would be expected to

(105-I 10 cm) as suggested by the similarity in the SDU from the modern human weight for

have a weight for its stature that would be unusual for modern human populations. OH 62 were similar in stature to AL 288-l femur lengths of these two hominids 30.4 kg (McHenry, stature RMA.

(Johanson et al., 1987) and had an estimated weight of

1988), it would lie 4-28-5.61

Like AL 288-1, it would be similar in its weight for stature to modern African weight for stature relationship but also in their body proportions 1990), suggests that OH 62 is,

apes. The similarity between these two fossils and their difference from modern humans, not only in their inferred CJohanson et al., 1987; Feldesman & Lundy, 1988; Aiello & Dean,

62, like AL 288-l) had a very different physique than is found in modern humans. IfOH in fact, a member of the genus Homo as argued by Johanson that ape-like stature/weight relationships are characteristic

et al. ( 1987)) it must be accepted not only ofsome members of the hominids suggest

genus Australofktlzecus but aIso of some members of the genus Homo. More fragmentary and problematical data for other Plio-Pleistocene the possibility that like AL 288-l body proportions ER 1500, TM uncertainties 1517) (McHenry,

and OH 62, at least some of these hominids may have had 1978; Grausz et al., 1988). However, like AL 288-l there are too many

that were quite different from those found in modern humans (e.g., KNMhominids to confidently found

in stature and weight predictions for these fragmentary

suggest that their stature weight relationships, in the KNM-WT 15 000 skeleton

and OH 62, were also signifiet al., 1985) suggest that by

cantly different from modern humans. However, the modern human body proportions (Homo cf. erectus) (Brown

approximately 1.6 m.y.a. at least some of the African hominids may have had weight for stature relationships that corresponded to the human condition. Indeed, the inferred stature (160 cm) and weight (48 kg) for this juvenile hominid and its projected adult stature (185 cm) and weight (68 kg) (Ruff, 1991), place WT 15 000 between - 1.55 (juvenilestature and weight) and - 1.99 (adult stature and weight) SDU beloze,the modern human RMA. These values are consistent sub-Saharan Africans. with the weight for stature relationships found in modern

Conclusions
Two major allometric conclusions techniques emerge from this analysis. and comparison The first is relevant to the choice of

for prediction

in the analysis of biological data. It has data to the central

been argued that in the great majority ofcases that the question being asked ofbiological is what the condition for an unknown specimen would be if it approximated

tendency of the reference sample used as the basis of analysis. For this purpose, the reduced major axis (RMA) is the clear choice above either the major axis (MA) or the least squares regression (LSR) slope. The reasons for this lie in the fact that the slope of the RMA is unaffected by the correlation coefficient of the sample. It can also be shown that the RMA provides the best estimate of the functional relation of the population from which the sample is drawn when the error variance is unknown, as is most frequently the case when analysing biological data.

146

L. C. AIELLO

The second major conclusion relationships relationship inference and comparison, for living African apes.

is relevant

to the interpretation

of the weight for stature is used as the basis of

of AL 288- 1 (Australopithecus it is demonstrated

afarensis). When the RMA

that AL 288-l has an inferred weight for stature

that would be highly unusual for modern humans but similar to that observed

Acknowledgements
Many thanks go to Laura of the theoretical I also gratefully Sigrid Bishop, Nora Giles, Danny Povinelli, Patricia Princehouse,

Melissa Remis, Eleanor development (1987/1988). Robin

Sterling

and Andrew Young for helpful discussion in relation to the aspects of this paper during my sabbatical at Yale University Dean, Martin, Bob Peter Andrews, Jan Austin, Christopher Bill Jungers, Anne MacLarnon,

acknowledge

Dunbar,

Hartwig-Scherer,

Andrew Nelson, Alan Porter, Chris Ruff, Simon Strickland and Lesley Willner for helpful comments on earlier drafts of this paper. Special thanks also are due to Derick Howellet of the Powell Cotton Museum in Birchington, Kent, for providing data from the museum records on stature and weight in wild collected African apes.

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