Morphologic variability of human African populations south of the Sahara

P. Passarello F. Vecchi

Morphologic Variability of Human African Populations South of the Sahara

Institute of Anthropology, University of Rome, 00185 Rome, Italy

Morphologic relationships between 133 African populations from south of the Sahara are analysed on the basis of 7 anthropometric variables by means of correspondence analysis. As far as concerns the morphologic characteristic, the distribution of populations on the plane formed by the first two factorial axes, which account for 85% of the total variability of data, (1) shows a certain geographic pattern, (2) is continuous, without forming any particular easily circumscribed groupings, and (3) indicates an association with the habitat in which the populations live.

Received 5 December accepted 22 February

1978 and 1979

Keywords: morphologic characters, African populations, correspondence analysis, human adaptation, races.

1. Introduction Synthetic

studies

on the African

indexes have been carried sent synthesis, African

correspondence

south

of the

analysis

tables of positive

numbers.

the various

technique-the

Benzecri,

a more synthetic

simplified

used to describe

In particular,

space with only a minimum

dependence

by drawing

loss of information.

for reducing

images of a multidimensional

tion to be made of the “vicinities” analysed,

indexes,

In the pre-

between

statistical

des correspondances:

of distance

(1969).

1970a,b)-is picture

tables

and a

existing

between

which in this case refer respectively

Thus

representation reality;

or, more

up a table of numeric

the method enables the data to be represented

analysis may be seen as an algorithm supplying

relationships

a multivariate

with distance

is a method

values of n lines and m columns, in a smaller

and Schwidetzky

analysis of the data.

Correspondence generally,

Sahara,

(analyse factorielle

analysis

which gives, compared

more detailed

based upon the calculations

(1968)

with the aim to clarify the morphologic

populations

employed

populations,

out by Hiernaux

graphically

the correspondence

space of the data and

moreover

it enables a descrip-

the lines and columns

to anthropometric

variables

of the tables and popula-

tions. In order factors,

to “concentrate”

the variability

are defined computationally

persion.

The first factor is a statistically

variables

which accounts

complete

set of original variables.

for a maximum

cally independent which account variability between data.

of data,

having maximum independent amount

composite potential

linear

variables,

for describing

linear combination

of total variability

i.e.

data dis-

of the original

implied

within the

The successive factors are linear

combinations

for a proportionally

amount of the residual

decreasing

recipro-

In the resulting graph, the similarity between two populations, or the association between two anthropometric variables is in a linear relationship with their relative distance.

Moreover,

each population

will be found close to those anthropometric

variables

for which it assumes high values. The correspondence analysis also provides analytical information instrumental to the interpretation of the data, the so-called “absolute contributions” and “relative contributions”. The first express the amount of variability to attribute to a given element, either a population or anthropometric variable, within the variability explained by a factor. Journal of Human Evolution (1979) 8, 467-474 0047-2484/79/040467+08

$02.00/O

@

1979 Academic

Press Inc. (London)

Limited

468

P. PASSARELLO

AND

F. VECCHI

The relative contributions measure the fraction to attribute to a factor in the explanation of the variability of a single element. Other multivariate techniques enabling graphic representation of data to be made in uni-, bi- or tri-dimensional space have been applied to the study of variability of African populations by Hiernaux (1973) and Rightmire (1976). Figure

1. Geographic

2. Populations

location

of populations.

studied

Data refer to 133 male African populations south of the Sahara. Mean values of seven anthropometric variables, namely: stature (ST), head length (HL), head breadth (HB), face breadth (FB), (total) face height (FH), nose breadth (NB) and nose height (NH) were taken into consideration for each population. The list of populations numbered according to the geographic position from northsouth and from west-east, and the source of data, are given in Table 1. Hiernaux’s manual (1968) was used where possible to localize the distribution area of the populations (Figure 1) ; in other cases Biasutti (1967) or Murdock (1959) were used.

3. Results

and discussion

Absolute and relative contributions of variables related to the first two factorial axes which account for 85 y0 of the total variability of the data, are given in Table 2.

MORPHOLOGIC Table

1. 2. 3. 4. 5. 6. 7. 8. 9. 10. 11. 12. 13. 14. 15. 16. 17. 18. 19. 20. 21. 22. 23. 24. 25. 26. 27. 28. 29. 30. 31. 32. 33. 34. 35. 36. 37. 38. 39. 40. 41. 42. 43. 44.

1

Amhara (1) Badyar (Ful) Koniagi Baiote Balante Badyaranke Afar (2) Kasange Issa (2) Bedik (3) Bedik (3) Warsingali Sara Fali (4) Fali (4) Kaba (5) Galla Nuer Sara Majingay (6) Anuak Day (5) Ngama (5) Djedje (7) Fon (7) Mbay (5) Gio Goun (7) Bamun Bororo (Ful) (10) Banda (9) Kran Bamileke Egap Baya (9) Holli (7) Ful-South Cameroon N’Zakara (8) Mangisa Binga (pygmies) (11) Kru Nago (7) Zande Maji (12) Grebo

List

VARIABILITY

56.

57. 58. 59. 60. 61. 62. 63. 64. 65. 66. 67. 68. 69. 70. 71. 72. 73. 74. 75. 76. 77. 78. 79. 80. 81. 82. 83. 84. 85. 86. 87. 88.

POPULATIONS

469

of populations

45. Kakwa

46. 47. 48. 49. 50. 51. 52. 53. 54. 55.

OF AFRICAN

Amar Kokke (12) Arbore (12) Duala Basa Ewondo Jue M’Bimou ( 11) Mangbele Logo Acholi (13) Bangba‘ Geleba (12) Mayogo‘ ’ Mamvu Mangutu Lugbara (14) Alur Tanga Borana fl5) 01 Mold (16) Budu Nyoro Teso (131 Sab ’ ’ Lika Mbuti (pygmies) Bira-savanna Bale Fang Bira-forest Shu Mvuba Humu Konjo Toro Ganda Komo Havu Swaga Nyankole Luo Oto-Konda (18) Nyanga

89. 90. 91. 92. 93. 94. 95. 96. 97. 98. 99. 100. 101. 102. 103. 104. 105. 106. 107. 108. 109. 110. 111. 112.

Hunde Hutu-ex Congo Kykuyu (17) Tembo Tutsi-Rwanda Haya Twa-Konda (pygmies) (18) Lega Twa-North Rwanda (pygmies) Hutu-Rwanda Fuliru Hutu-Burundi Shi Tutsi-Burundi Sukuma Masai Bushong Nyamwezi Nyaturu Sandawi Cwa-Kuba (pygmies) Songye Oio (18) Swahili 113. Luba-Kasai 114. Luba-Katanga 115. Chokwe (19) 116. Ginga (19) 117. Mbundu 118. Bieno (19) 119. Luimbe (19) 120. Muila (19) 121. Humbe (20) 122. Kwanyama 123. Kwanyama (20) 124. Vatwa (21) 125. Chimba (21) 126. Kung (Bushmen) 127. Bitonga (20) 128. Chope (20) 129. Dama (22) 130. Mateve (20) 131. Auni Khomani (Bushmen) (23) 132. Nama (Hottentots) 133. Korana (Hottentots)

(1) Coon (cit. by Gschinsky, 1954); (2) Charpin & Georget (1977); (3) Gomila (1971); (4) Huizinga & Reijnders (1974); (5) Hiernaux (1969); (6) Crognier (1973); (7) Correnti ct al. (1972-1973); (8) Cresta (1965a); (9) Cresta (19656); (10) Cresta (1965~); (11) Cresta (1965d); (12) Vecchi & Ricci (unpubl.); (13) Oschinsky (1954); (14) Gruppioni & Rivalta (1977) ; (15) Ricci (unpubl.) ; (16) Corrain (1972); (17) Corrain & Capitanio (1971); (18) Hiernauxetul. (1976); (19) R&sing (1977) ; (20) Weninger & Schciber (1975) ; (21) Weninger (1965); (22) Knussmann & Knussmann (cit. by Knussmann 8s Rosing, 1974); (23) Dart (cit. by Knussmann & Riising, 1974). The other populations were taken from Hiernaux (1968).

470

P. PASSARELLO AND F. VECCHI

Figure 2. Correspondence analysis for 133 African populations and 7 anthropometric variables (ST = stature; HL = head length; HB = head breadth; FB = face breadth; FH = face height; NB = nose breadth; NH = nose height).

-1

- 0.02

1

IN n n

.‘O.02

1 I

,04

NN I

1

I

Fint factorial

Absolute ( x 100) ad metric variables

Table 2

relative ( x 100) contributions

Factorial

, Anthropometric variables 1. 2. 3. 4. 5. 6. 7.

Stature Head length Head breadth Face breadth Face height Nose breadth Nose height

ST HL HB FB FH NB NH

I

absolute

relative

24.77 5.96 19.31 21.87 2.79 21.93 3.37

97.16 38.20 76.92 82.18 20.91 65.90 15.16

\

The factor associated

of anthropo-

axes II Contributions L

I

Contributions n

/

’

absolute

relative

1.13 2.32 0.33 3.15 27.76 8,22 57.09

1.36 4.60 0.40 3.64 64.08 7.63 79.2 1

\

100~00

100-00

ponds, is characterized

.__I--

axis (65.85%)

with the first axis, to which 66% by bipolarity

between

of the total variability

corres-

the ST and the NB, FB, HB variables.

The factor associated with the second axis, to which 19% of the total variability corresponds, is characterized by bipolarity between the NB and the NH, FH variables. The HL variable

is of little importance

in the characterization

of the first two factors.

MORPHOLOGIC

The populations

are distributed

VARIABILITY

on the factorial

stretching

along the first axis (Figure

Hiernaux

(1973)

with a principal

2).

the distances ever,the

‘2, with reference between

coordinates

between

In other words, populations similar.

Various

mobility

of populations;

metric

geographic

dimensions;

(c) non-genetic

(d) sampling

to the anthropometric

tions are distributed

over the factorial

succession : (1) populations

the savanna-forest

mosaic;

grasses;

grassland;

Polunin

( 1969)].

(a) isolating

(e) lack of standardization

plane

considered

(Figure

How-

roughly.

populations scattered savanna;

more

barriers;

(b)

of the anthropoof anthropometric

and the habitat,

3) approximately

living in the tropical

bush

pattern.

is drawn

in the expression

(3) open country

savanna;

distances

for this finding:

variables

by

taken into consideration, geographic

closer are not always morphologically variability

errors;

was obtained

on the plane formed by

to a set of 32 African populations.

and morphologic

geographically

to a cloud of points

pattern

variables

reveal a certain

causes may be responsible

techniques. With reference following

data referring

to the anthropometric

471

POPULATIONS

plane according

analysis by plotting

the population-points

relationship

AFRICAN

A similar distribution

the first two latent vectors, morphological In Figure

OF

the popula-

according

to the

rainforest ; (2) populations [broadleaf broadleaf

of

deciduous trees and deciduous shrubs,

Figure 3. Correspondence analysis for 133 African populations and 7 anthropometric variables in relation to habitat in which populations live (ST = stature; HL = head length; HB = head breadth; FB = face breadth; FH = face height; NB = nose breadth; NH = nose height). Cl, rainforest; 0, savanna-forest mosaic; A, open country.

-0.04

-0.02

I

0

0.02

1

A

A

0

A

oA” A

0.04.

I

3.02

1

’

0

AA

0

A AA A A~

A A

.02

*04 First factorial axis (65.85%) Thus indicates

the distribution an association

of the variables between

and

the morphologic

the populations characteristics

on the factorial of the populations

plane and

472

P.

the habitat

PASSARELLO

AND

Populations

in which they live.

Furthermore, Analogous

relationships 1954;

tion to the climatic

affinity can be seen between

demonstrated

1968,

between

of African

1977;

climatic

populations

Schreider,

or biogeographic

(Thomson

1971;

Hiernaux

1976) ; they have in general been interpreted

& Buxton, et al.,

1975;

as a biological

adapta-

morphologic

reality

ambient.

relationships,

of Africa.

been

morphology

Hiernaux,

& Froment,

These

have

and the somatic

Weiner,

Hiernaux

in the values of the other variables.

a morphological

of the north and south savanna.

characteristics 1923;

of this association,

area are marked by low

Going from the forest to the open country

in the ST values and a decrease

on account

the populations

VECCHI

living in the rainforest

ST values and high NB, FB and HB values. there is an increase

F.

Mobility

however,

only explain

of the populations

a part of the complex

and the consequent

possibility

tween genetically

of admixture

be-

different populations are some of the most important It is possible in this way to interpret, for example, with this scheme.

factors in contrast the morphological

differences

region

observed

between

78-82-88-92-96-99) latter,

in fact,

following adapted

forest populations

which

recently

penetrated

the Ful expansion to this particular

For example,

(1) sudanid,

into the rainforest

in the Adamaoua

In order to explain the morphologic

area:

area (33-38-48-49-50-5 region

(17th

(Alexandre,

(71-75-77-

l-63-74). and

The

18th century)

1965)

might

be less

habitat.

mists usually group the populations ies or races.

of the interlacustrine

and those of the Guinean

complexity

v. Eickstedt

(2) pygmid,

of Africa

into a certain number distinguishes

(3) palaeonegrid,

south of the Sahara

taxono-

of discrete morphologic

categor-

7 racial groups in this geographical (4) nilotid,

(5) ethiopid,

(6) bantuid,

(7) khoisanid. A comparison

between

the distribution

when classified approximately races published populations

by v. Eickstedt

classifiable

of populations

in Figure 2 and their distribution

into “races” on the basis of the distribution (1934:

according

positions close to one another

see map between

to this Author

on the factorial

in a single racial group tend to occupy Populations

plane.

on the right of the graph and tend to separate particularly Populations

(1) and (6) present similar morphologic

the central

part of the graph;

and (5) tend, nevertheless, particularly

along

widely scattered

to separate

the second

along

populations factorial

the second

map of African

pp. 560 and 561), shows that the (2) and (3) are localized

along the second factorial

characteristics

axis.

and are grouped

in

(4) and (5) are found on the left side; (l-6),

along the first axis; axis.

Populations

(4) and (5) along the first and (7),

on the other

axis and are not distinguishable

hand,

are

from other African

populations. The

agreement

populations

between

as appears

is based on morphological

the above-mentioned

in Figure criteria

2, is probably

racial because

in the present study.

Hiernaux

the African populations

1973) suggested,

However

Figure

of

of v. Eickstedt

and in part utilizes the same anthropometric

as those taken into consideration (1968,

groups and the distribution the classification

characters

2 shows that, as

south of the Sahara are distribu-

ted, regarding their morphologic characteristics, in a continuous fashion, i.e. without forming any particular, easily circumscribed, groupings. For instance: (a) the Mbuti pygmies (7 1) are clearly separated from the other populations; other “pygmids” (95-97)) do not appear to be morphologically distinct from the “palaeonegrids” ; (b) the morphological characteristics of the “palaeonegrid” populations merge unnoticeably with those of the “sudanid-bantuid” populations and it is not possible to fix a morphologic limit to

MORPHOLOGIC

separate

these groups;

VARIABILITY

(c) only two “nilotid”

populations,

(20) appear to be isolated from the other populations; teristics of “ethiopid” bantuid” Ethiopia

populations

as the distance

POPULATIONS

473

the Nuer (18) and the Anuak

(d) also the morphological

charac-

from their more typical distribution

with the exception

therefore a distribution,

area, northern

increases.

In the final analysis, the morphological

less arbitrary

AFRICAN

populations tend to merge with those of both “nilotid” and “sudanid-

and the African Horn,

the Sahara,

OF

variations

of the African populations

of some special cases, tends to be gradual.

which varies basically with a continuous

number of discrete categories,

pattern,

south of

Subdividing into a more or

necessarily gives an approximation.

References Alexandre, P. (1967).

Langues et Cangage en Afrique noire. Paris: Payot. Benzecri, J. P. (1970s). Distance distributionnelle et mitrique du chia en analyse factorielle des correspondances. Paris: Faculte des Sciences. Benzecri, J. P. (1970b). Lefons sur C’analyse statistique des don&es muCtidimensionneCCes.Paris: FacultC des Sciences. Biasutti, R. (1967). Le Razre e i Popoli della Terra 3, Torino: U.T.E.T. Cavalli-Sforza, L. L. (1972). Origin and Differentiation of Human Races. (Huxley Memorial Lecture, 1972). Proceedings of the Royal Anthropological Insitute of Great Britain and Ireland 15-25. Charpin, M. & Georget, J. P. (1977). Comparaison anthropologique entre les Afars et les Issas de Djibouti. Bulletins et Memoires de la Sociiti d’AnthropoCogiede Paris 4, 113-l 19. Chopra, V. P. & Schwidetzky, I. (1970). Quantitative Rassen-Systematik. Homo 21,40-46. Corrain, C. (1972). Nuovi risultati di ricerche antropometriche ed emotipologiche tra le popolazioni de1 Kenya. Alcune tribh delle Frontiere de1 Nord: 01 Molo, Turkana, Samburu e Rendille. Archivio per C’AntropoCogiae la Etnologia 102, 3-86. Corrain, C. & Capitanio, M. (1971). Primi risultati di ricerche antropometriche ed emotipologiche tra le Archivio per PAntropoCogiae la Etnologia 101,5-36. popolazioni de1 Kenya. I Kykuyu de1 Nyandarma. Correnti, V., Vecchi, F., Capucci, E., Spedini, G. & Cresta, M. (1973). Indagine antropologica nel Basso Dahomey. Nota I-Caratteri morfologici e morfometrici degli adulti. Rivista di Antropologia 58, 39-76. Cresta, M. (1965a). Antropologia morlologica e sierologica dei N’Zakara della Repubblica Centrafricana. Quaderni de “La Ricerca Scientihca” 28, 9-20. Cresta, M. (19656). Note antropologiche sui Baya ed i Banda della Repubblica Centrafricana. Quaderni de “La Ricerca Scientifica” 28, 59-68. Cresta, M. (1965~). Contributo alla conoscenza dei Fulbe Bororo. Quaderni de “La Ricerca Scientijca” 28, 69-80. Cresta, M. (1965d). Contributo alla conoscenza dei Babinga. Quaderni de “La Ricerca Scientifica’” 28, 8 l102. Crognier, E. (1973). Adaptation morphologique d’une population africaine au biotope tropical: les Sara du Tchad. Bulletins et MCmoires de la Sociiti d’AnthropoCogiede Paris 10,3- 15 1. Crognier, E. (1977). Assortative Mating for Physical Features in an African Population from Chad. Journal of Human Evolution 6, 105-I 14. Eickstedt, E. v. (1934). Rassenkunde und Rassengeschichte der Menschheit. Stuttgart: Verlag. Gomila, J. (197 1). Lzs Bedik (Se?u!gal oriental). Barr&es culturelles et hitiroglniitc’ biologique. Montreal : Presses de l’universite. Gourou, P. (1970). L’Afrique. Paris: Hachette. Gruppioni, G. & Rivalta, G. (1977). Osservaaioni antropologiche sui Lugbara de1 West Nile (Uganda). Archivio per PAntropologia e la Etnologia 107, 293-304. Hiernaux, J. (1965). H&edit& milieu et morphologie. Biotypologie 26, l-36. Hiernaux, J. (1966). Les Bushong et les Cwa du Royaume Kuba (Congo-Kinshasa) : Pygmtes, pygmo’ides Bulletins et Mimoires de la So&% et pygmeisation; anthropologie, linguistique et expansion bantoue. d’dnthrojologie de Paris 9, 299-336. Hiernaux, J. ( 1968). La diversite’humaine en Afrique Subsaharienne. Recherches biologiques. Bruxelles: Editions de 1’Institut de Sociologic de l’universitt libre de Bruxellcs. Hiernaux, J. (1969). Investigations anthropobiologiques au Moyen-Chari (Republique du Tchad) prtliminaires a des recherches multidisciplinaires. Homo 20, l-10. Hiernaux, J. (1973). Numerical taxonomy of Man: an application to a set of thirty-two African populations. In (S. S. Sarkar Memorial Volume), Physical Anthropoloo and its extending horizon, pp. 151-161. Calcutta: Orient Longman Limited.

474

P. PASSARELLO

AND F. VECCHI

The Peojde of Af nca. . Hiernaux, J. (1974). London: Weidenfeld and Nicolson. Hiernaux, J. (1977). Long-term biological effects of human migration from the African savanna to the equatorial forest: a case study of human adaptation to a hot and wet climate. In (G. A. Harrison, Ed.) Population structure and human variation, pp. 187-Z 17. Cambridge: University Press. Hiernaux, J. & Froment, A. (1976). The Correlations Between Anthropobiological and Climatic Variables in Subsaharan Africa: Revised Estimates. Human Biology 46, 757-767. Hiernaux, J., Rudan, P. & Brambati, A. (1975). Climate and the weight/height relationship in Subsaharan Africa. Annals of Human Biology 2,3-12. Hiernaux, J., Vincke, E. & Commelin, D. (1976). Les Oto et les Twa des Konda (zone de l’equateur, Zaire). L’Anthropologic 80, 449-464. Huizinga, J. & Reijnders, B. (1974). Skinfold thickness and body fat in adult male and female Fali (North Cameroon). Proceedings of the Koninklijke Nederlanse Akademie van Wetenschappen 77C, 496-503. Knussmann, R. & Rosing, F. W. (1974). Die Ahnhchkeitsverhaltnisse im siidwestlichen Afrika nach anthropometrischen Merk malen. In (W. Bernhard & A. Kandler, Eds) Bevslkerungsbiologie, pp. 125153. Stuttgart: G. Fischer. Knussmann, R. & Rosing, F. W. (1977). The Structure of Similarities in Southwestern Africa According to Anthropometric Characteristics. Journal of Human Evolution 6, 2 1 l-2 15. Murdock, G. P. (1959). Africa. Its People and Their Culture History. New York: McGraw-Hill. The Racial Ajinities of the Baganda and other Bantu Tribes of British East ‘4ftica. Oschinsky, L. (1954). Cambridge: W. Heffer & Sons. Polunin, N.-( 1969). Introduction to Plant Geography and Some Related Sciences. London: Longmans. Rightmire, G. P. (1976). Multidimensional Scaling and the Analysis of Human Biological Diversity in Subsaharan Africa. American Journal of Physical Anthropology 44, 445-452. Rosing, F. W. (1977). Anthropometry of six tribal populations in Angola. Zeitschrift fir Morphologic und Anthropologic 66, 107-122. Schreider, E. (1950). Geographical distribution of the body-weight/body-surface Ratio. Nature, London 165, 286. Schreider, E. (1957). Ecological rules and body-heat regulation in man. Nature, London 179,915-916. Schreider, E. (1964). Ecological rules, body-heat regulation and human evolution. Evolution 18,1-9. Schreider, E. (197 I). Variations morphologiques et differences climatiques. Biomitrie Humaine 6, 46-69. Schwidetzky, I. (1969). Zur Rassengliederung dcs subsaharischen Afrika. Statistische Untersuchungen au Hiernaux’s Afrikabuch. Homo 20, 1 l-34. Thomson, A. & Buxton, D. (1923). Man’s nasal index in relation to certain climatic conditions. The Journal of the Royal Anthropological Institute of Great Britain and Ireland 53, 92-122. Weiner, J. S. (1954). Nose shape and climate. American Journal of Physical Anthropoloa 12, l-4. Weninger, M. (1965). Chimba und Vatwa, bantuide Viehziichter und nicht-bantuide Wildbeuter. Mitteilungen der Anthropologischen Gesellschaft in Wien 95, 180-I 90. Weninger, M. & Scheiber, V. (1975). Anthropologisch-metrische Untersuchungen an west-und ostafrikanischen Bantunegern. Zeitschrift fiir Morphologic und Anthropologic 67, 60-84. Weninger, M. & Scheiber, V. (1977). Anthropometrical Investigations of West and East African Bantu. Journal of Human Evolution 6, 729-732.