Dermatoglyphics of inhabitants of Alberche/Tormes valley (Sierra de Gredos-Central Spain): I. Finger pattern types and pattern intensity

AMERICAN JOURNAL OF HUMAN BIOLOGY 8:305-316 (1996)

De rm a t og I y p h i cs of I n habit ants of Al be rc h e/To rm es Val ley (Sierra de Gredos-Central Spain): I. Finger Pattern Types and Pattern Intensity

J MARTIN,l M S MESA,Z V FUSTER,2 AW P MORAL3 'Departmento de Biologre, Unidad de Gen6taca, Uniuersidad Autonoma de Madnd, 28049, Madnd, LDepartmento de Bzologia Ancmal I , Catedra de Antropologaa, Facultad de Bzologia, Unzversadad Complutense de Madrid, 28040, Madrid, 3Departmento de Bzologia Ancmal, Catedra de Antropologra, Facultad de Rrologra, 1Jniversidad de Barcelona, 08028, Barcelona, Spain

ABSTRACT The incidence of dermatoglyphic finger patterns and pattern intensity in a sample of school children from the AlbercheA'ormes valley (northern slope of the Sierra de Gredos, Central Spain) are described. Signifi- cant bilateral differences for the overall finger pattern incidence are found in both sexes. The differences between sexes are also statistically significant, except for the left hand. The Pattern Intensity Index significantly differs between hands of males only, and the sex differences are not significant. Variation ranges for fingertip patterns and pattern intensity were constructed using published data of 69 male and 62 female series drawn from the whole of the Iberian Peninsula. The overall incidence of fingertip patterns and the Pattern Intensity Index locate the studied population near the middle of the corresponding variation ranges. The only exception is the case of arches, for which AlberchelI'ormes is close to the minimum value of the range of variation. A more detailed comparison of 10 male and 10 female series from the Spanish northern and southern plateaus and from the Spanish region of Extremadura was done with R-matrix analysis. The series compared include those from the northern slope (Alberche/Torrnes valley, this study) and the southern slope (Tietar valley) of the Sierra de Gredos. The observed degree of dermatoglyphic differentiation between these two series may be explained by taking into account both the role of the Sierra de Gredos as a biological barrier limiting gene flow and the evolutionary stability of dermatoglyphics. 0 1996 Wiley-Liss, Inc

The influence of regional geography on the genetic structure of human populations is well established. For example, the studies of Piazza et al. (19881, Barbujani and Sokal (1990), Calafell and Bertranpetit (19941, among others, have shown that the genetic differentiation among European populations is influenced not only by cultural factors, e.g., linguistic boundaries, but also by physi- cal, i.e., geographic, elements.

In the Iberian Peninsula the mountainous Central System runs SW-NE across the plain that occupies most of central Spain (Central plateau) and encompasses several Spanish provinces (Fig. 1). The part of the Central System within Avila province, the

Sierra de Gredos, differentiates two clear regions, the Alberchernormes valley on the northern slope and the Ti&tar valley on the southern slope. The AlberchePTormes valley is a geographically closed region, while the southern end of the Tietar valley freely opens onto the Southern plateau.

The mountain range reaches a maximum altitude of 2,660 m and the two regions are connected only by three natural passes. It is,

Received Octoher 6, 1994; accepted August 27, 1995. Address reprint requests to Dr. Jose Martin, Depto. de Binlogfa,

Unidad de Genktica, Edificio de Biologia A-207, Universidad Au- tonoma de Madrid, 28049, Madrid, Spain.

Q 1996 Wiley-Liss, Inc.

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KEr-CODES PO FW L A T I 0 N REFERENCE

1 - A L C LA ALCARRIA (Guadalajara) Portabales, 1983

2 BAD BADAJOZ Oyhenart, 1986

3 BIE EL BIBRZO (Le6n) Cuenca, 1979

4 - CAC CACERES Oyhenart. 1986

5 - GRP. SIERRA DE GREDOS (Avxla) Fhster and Cahello. 1986

6 MAD MADFUD Martin and Castellanos. 1 9 x 8

7 MAK LA MARAGATERIA (Le6n) Car0 et al., 1988

8 - T C A L'IEKRA DE CAMPOS (Pnlencia) Martin and Portabales, 1 9 8 5

9 - TIE TIETAR VALLEY( Avila) Marrin et a1 , 1993

10 - ALI' ALBERCHE/TORhfES VALLEY (Avila) This sludy

Fig. 1. A: Location of AlbercheEormes valley within the Sierra de Gredos and within Iberian Penin- sula. Numbers within the top map correspond to geographical position of populations used for detailed comparisons. B: Key codes, names, and original references of populations used for detailed comparisons with Alberchernormes. Numbers before each key code correspond to those of Lhe top map of A.

DERMATOGLYPHICS IN ALBERCHEmORMES VALLEY (SPAIN) 307

therefore, possible that the Sierra de Gredos was an important barrier to population movement in the past, especially during the winter. If the Sierra de Gredos afforded lim- ited dispersal and migration, biological differences between populations settled on the two slopes might be expected.

In this context, a long-term anthropogene- tic study has been devised to assess the influence of the Sierra de Gredos on the genetic structure of human populations living on its two slopes. Data on dermatoglyphics, body morphology and blood markers have been collected from the populations living in this part of Spain. Data concerning marital movement have been also obtained from parish registers of the area.

The aim of the present paper is to describe the qualitative finger dermatoglyphics and pattern intensity of the population living in the AlbercheB'ormes valley (northern slope of the Sierra de Gredos in Avila province, Spain). Since similar data have recently been published for the population located on the southern slope of the Sierra de Gredos (Tiktar valley) (Martin e t al., 1993), the im- portance of the mountain range as a biological barrier to complex traits such as dermatoglyphics can be evaluated. The dermatoglyphic relationships among the present sample and other previously studied Iberian samples are also examined.

From an historical point of view, there is evidence of human presence in the Sierra de Gredos since 1,000 B.C. These human groups, the Vettones, were Celtic in origin, had tribal organization, and were mainly en- gaged in shepherding (Fernandez Gomez, 1986; Salinas de Frias, 3986). The Romans appear not to have had any economic inter- est in this land, as the absence ofthe remains of any permanent settlements suggests. Later, in the 6th century, groups of Visigoths also settled in the territory. During the first centuries of Moslem occupation, the population of the region of the Sierra de Gredos would have been sparse since it was a fron- tier between the Moslem and Christian Kingdoms and wars were frequent U'roitiiio, 1990). Repopulation of the region began in the 13th century. I t proceeded very slowly, with people arriving from important neigh- boring cities (e.g., Avila, Plasencia, Bejar). During the early Middle ages, repopulation continued with people coming from the mountains of Soria and Burgos (northern Spain). Moreover, since the area has a com-

plex geography, groups of persecuted minori- ties, such as Jews, Moors, and Mozarabs, could have hidden in the villages of the mountains of the Sierra de Gredos, occu- pying their own districts (Troitino, 1990).

The current economy and way of life of the people living in the Sierra de Gredos is clearly influenced by climate. The climate in the northern valleys is colder than in the southern valley; thus, cattle herding and a modest amount of forestry are presently prevalent in the AlbercheBormes, while the cultivation of temperate zone crops charac- terizes the agriculture of the Tietar valley.

Presently, the Alberche/Tormes valley comprises about 38 villages with a total population of 35,688 inhabitants. As with many other rural Spanish populations, an intense emigration to industrialized regions has taken place since 1960, causing a decrease in the size of the populations (INE, 1988; Troitiiio, 1990).

MATERIALS AND METHODS During the school years of 1989190-19901

91, fingerprints were obtained of 406 unre- lated and healthy school schildren (187 boys and 219 girls) from AlberchePTormes valley whose four grandparents had all been born in that valley. Although this sample is ap- proximately 1% of the total population of the AlbercheBormes valley, it actually repre- sents a higher proportion of the native population. Sampling was carried out throughout the entire area, sample sizes being propor- tional to the actual population size of each village.

Ink and transparent adhesive tape were employed to obtain the fingerprints. The methodology of Cummins and Midlo (1961) was used t o classify fingertip patterns.

Sexual and bilateral differences were assessed by means of x2 contingency tables (fingertip pattern incidence), and the Wilcoxon- signed rank and Mann-Whitney tests (Pat- tern Intensity). The normality of pattern intensity distributions was tested by means of the Kolmogorov-Smirnov one sample test. The STATGRAPHICS statistical package (PLIJS*WARE, STSC, Inc., Rockville, MD) was used for the analyses.

To parsimoniously depict the biological (dermatoglyphic) relationships between the Alberchemormes sample and other nearby previously studied Iberian samples, an R-matrix analysis (Harpending and Jen- kins, 1973) was performed, adopting the

308 J. MARTIN ET AL.

TABLE 1. Ouerall inridenre offinger pattern types in males and females from AlberchelTormes ualley (Sierra de Gredos, norfh.ern slope)

Females (n = 219) Males in = 1871 R L R + L R L R + L

Arches 2.99 3.64 3.32 3.47 5.11 4.29 Radial loops 4.60 3.64 4.12 2.01 4.11 3.06 Ulnar loops 54.76 62.89 58.82 62.47 59.91 61.19 Whorls 37.65 29.84 33.74 32.05 30.87 31.4fi

same assumptions as Crawford and Duggir- ala (1992). In this procedure, the R-matrix was obtained from the matrix of the four fingertip patterns appearing on the 10 fingers in the array of the compared populations. Eigenvalues and eigenvectors of the R-matrix were then computed, and the populations were ordered according to their scores for the first three eigenvectors. In this way, a series of plots showing the dermatoglyphic affinities o r differences among the compared populations was obtained. Fur- ther, it is also possible with this method to compute the scaled matrix of covariances among fingertip pattern frequencies, i.e., the S-matrix of Harpending and Jenkins (1973), and by working with its eigenvalues and eigenvectors in a manner similar to that with the R-matrix, a series of plots of fingertip patterns can be obtained. By comparing the plots of populations with the plots of fingertip patterns, the contributions of the different fingertip patterns to the population rela- tions can be assessed. A 0.05 significance level was adopted in all analyses.

RESULTS Fingertip patterns

Table 1 summarizes the percentage frequencies of fingertip patterns in males and females. Ulnar loops and whorls are the most frequent patterns in both sexes, while arches and radial loops are the least frequent. Bilat- eral differences in the overall incidence of finger patterns are statistically significant in both sexes (males, xa = 15.33, 3 d.f., p < 0.01; females x2 = 12.17, 3 d.f., p < 0.01), although the pattern of asymmetry is different in males and females. In males, radial loops and whorls appear more frequently on the right hand: while ulnar loops and arches appear more frequently on the left hand. In females, ulnar loops and whorls appear more frequently on the right

hand, while arches and radial loops appear more frequently on the left hand.

There are statistically significant differences between the sexes for the overall incidence of finger patterns in the right hand (x2 = 20.55, 3 d.f., p < 0.001) and for right and left hands together (x2 = 8.24, 3 d.f., p < 0.05) due to the greater incidence ofra- dial loops in males. However, between-sex differences in finger patterns €or left hands are not significant (xa = 3.68, 3 d.f., p = 0.298). It should be noted that females show a greater frequency of radial loops than males in the left hand, which is the major cause of the insignificant differences between the sexes for this hand.

Interfinger variability in the incidence of various fingertip patterns is detailed in Ta- bles 2 (males) and 3 (females). Although arches (both tented and plain) and radial loops are infrequent patterns, they appear more frequently on index fingers (right or left hands). Ulnar loops appear with greatest frequency on the little and middle fingers. Whorls appear more frequently on the ring finger (true whorls) and thumb (double loops). Very occasionally, patterns with 3 triradii or patterns consisting of two basic patterns combined, particularly on index fingers, are scored. The greater incidence of radial loops in the left index finger of females compared with males should be noted.

Pattern intensity Table 4 shows the statistical parameters

of pattern intensity. The male series tends to be leptokurtic (kurtosis > 01, except for the left hand, as well as right-skewed (skewness > 0), except for the right hand. Pattern intensity distributions in females are platykurtic (kurtosis < 0) and left- skewed (skewness < 0) except for the left hand. In no case are the coefficients statistically significant, which implies that none of

DERMATOGLYPHICS IN ALBERCHE/TORMES VALLEY (SPAIN) 309

TABLE 2. Distribulion ofpattern t,ypes in the fingers of males in = 1871 from AlherchelTormes ijalley

R1 R2 R3 R4 R5 L1 L2 L3 L4 L5

Plain arches Tented arches Total arches Radial loops Ulnar loops Whorls Double loops Composites

(3 triradii) Accidentals Total whorls

1.07 0.0 1.07 0.53

47.59 31.02 19.79 0.0

0.0 50.81

5.35 3.21 8.56

19.25 31.02 31.02 9.09 0.0

1.07 41.18

3.74 0.53 0.53 2.67 5.35 3.21 0.53 1.07 0.53 0.0 0.0 0.0 3.74 1.07 0.53 0.0 4.28 0.53 0.53 2.67 9.09 4.28 1.07 1.07 1.60 1.60 0.0 1.07 16.04 1.07 0.0 0.0

74.33 41.71 79.14 54.55 42.78 75.94 56.68 84.49 16.04 52.41 18.18 18.72 22.99 14.97 36.90 11.76 3.74 8.74 1.60 22.99 8.56 2.67 5.35 2.67 0.0 0.0 0.53 0.0 0.53 1.07 0.0 0.0

0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 19.79 56.15 20.32 41.71 32.09 18.72 42.25 14.44

TABLE 3. Dishhut ion of pattcrn types in the fingers offemales 1n=219) from AlbercheiTormes ualley

R1 R2 R3 R4 R5 L1 L2 L3 L4 L5

Plain arches Tented arches Total arches Radial loop Ulnar loops Whorls Double loops Composites

(3 triradii) Accidentals Total whorls

4.57 0.0 4.57 0.91

49.32 29.68 15.53 0.0

0.0 45.21

5.02 4.57 0.91 0.91 0.91 0.46 0.0 0.0 5.94 5.02 0.91 0.91 7.76 0.46 0.91 0.0

46.58 81.74 50.23 84.47 32.42 8.68 47.03 13.70 6.85 4.11 0.91 0.91 0.46 0.0 0.0 0.0

0.0 0.0 0.0 0.0 39.73 12.79 47.95 14.61

3.20 5.48 9.59 2.28 1.37 0.0 3.20 0.0 0.0 0.46 3.20 8.68 9.59 2.28 1.83 1.37 17.35 0.91 0.91 0.0

56.62 33.33 70.78 54.79 84.02 21.00 33.33 15.07 39.73 13.70 17.81 6.85 3.65 2.28 0.46 0.0 0.0 0.0 0.0 0.0

0.0 0.46 0.0 0.0 0.0 38.81 40.64 18.72 42.01 14.16

TABLE 4. Statistical naratneters of m t t e rn intensity in males and females from AlberchelTormes ualley

Sex Hand Mean S.D. Median 5S.E. Skewness Kurtosis D-Max. (%I

Males Right 6.77 1.72 7.00 0.29 -0.04 0.13 16.08' (n - 187) Left 6.38 1.68 6.00 0.00 0.32 -0.23 19.36'

R t L 13.15 3.17 13.00 0.29 0.01 0.10 13.44' Females Right 6.45 1.65 6.00 0.29 -0.05 -0.25 14.26'

(n = 219) Left 6.32 1.75 6.00 0.29 0.03 -0.08 14.98' K + L 12.77 3.23 12.00 0.29 -0.02 - 0.16 10.08'

'Significant at 0.05 lcvrl (Kolmogorov-Smirnov one-sample normality test).

the distributions should deviate from the theoretical values of zero for a normal distribution. Nevertheless, the Kolmogorov-Smir- nov test of normality indicates that the six distributions significantly deviate from normality (last column of Table 4).

Pattern intensity offers a quantitative measure of the general complexity of the fingertip patterns of an individual. Thus, on average, males from Albercheflormes valley have more complex patterns than females, and the right hand of males and females shows more complex patterns than the left hand (third column of Table 4). Statistical analyses of the differences in pattern intensity between hands or between the sexes are

presented in Table 5. Right versus left differences, assessed by nonparametric Wilcoxon signed-rank test, are statistically significant only in males, although they approach significance in females. Differences between sexes, assessed by means of the nonparametric Mann-Whitney test, are not significant in any of the three possible comparisons, although they approach significance in the right hand.

DISCUSSION The percentage frequencies of fingertip

patterns in the school children of the present study are consistent with those usually found in dermatoglyphic studies (e.g.,

31 0 J. MART" ET AL

TABLE 5. Pattern intensity: Bilateral and sex difrerenres, Alberche ITormes oa1le.y

1. Bilateral differences (Wilcoxin signed-rank test) Males Females

Z-test statistic 4.30 1.67 P value 1.74 X 10 0.095

2. SCX differences (Kruskall-Wallis test) Right Left R + L

Z-test statistic 3.02 0.02 1.11 P value 0.08 0.90 0.29

Steinbcrg et al., 1975; Schaumann and Alter, 1976; Jantz and Brehme, 1978; Bartsocas et al., 1982; Gomez and Martin, 1992). On the other hand, comparison of the results for pattern intensity with those of other studies dealing with pattern intensity distributions (e.g., Pons, 1958; Vrydagh, 1970; Malhotra et al., 1980; Unturbe, 1980; Martin and Por- tabales, 1985) permits no general conclusion t o be drawn regarding the shape of the frequency distribution of this dermatoglyphic trait, other than that it is not normal.

Table 6 illustrates the position of the Alb- erchemormes valley sample relative to other population samples from the Iberian Penin- sula with regard to the dermatoglyphic traits described in this report. Ranges of variation and the weighted averages of the basic fingertip patterns, and the weighted average of pattern intensity are based on relevant information from an lberian Der- matoglyphic Database, drawn from previously published studies. The data comprise a total of 69 male series and 62 female series from the whole of the Iberian Peninsula.

The frequency of arches in the Alberchel Tormes valley is lower than the Iberian average and close to the minimum values in both males and females. The incidence of radial and ulnar loops in males and females is slightly lower than the corresponding Ibe- rian averages. Regarding the incidence of whorls, males and females from the Alber- chemormes valley show values greater than the corresponding Iberian averages, especially females. Finally, the values of pattern intensity observed in the studied population are slightly higher than the corresponding average values of male and female Iberian samples.

An R-matrix analysis was done to assess the dermatoglyphic relationships between the analyzed sample and other geographically proximal samples. These include series

from the Northern and Southern Spanish plateaus, as well as from the Spanish region of Extremadura (see Fig. 1 for geographical locations within the Iberian Peninsula). A total of 10 male series and 10 female series was analyzed. Among these series, all of the available samples from the Sierra de Gredos region have been used, including those of the AlberchePTormes valley (ALT northern slope of Sierra de Gredos, this study), the Tietar valley (TIE; southern slope of Sierra de Gredos; Martin et al., 1993 j and a series from the whole area of the Sierra de Gredos (GRE; Fuster and Cabello, 1986). This series differs from the former two in that i t is composed of people whose parents and grandparents were born in any village of the Sierra, irre- spective of the slope from which they came. Thus, the genetic background of the people comprising this series may have differed from that of Alberchefrormes and Tiktar in- asmuch as the last two series differ genet- ically.

The frequencies of the four basic finger pattern types appearing on each of the 10 fingers (40 variables in total) have been used as descriptors in the analysis. About 64% and 71% of intergroup variation is accounted for by the three eigenvectors in males and females, respectively. Due to the large number of characterizing variables, it is difficult to distinguish and describe a clear pattern of variable combinations which could be responsible for the scattering of populations along the scaled eigenvectors (see Figs. 2 and 3).

Although there are various finger patterns with high scores for the eigenvectors in the male series, the simplest way to interpret the first three is by means of the dermatoglyphic patterns of ring fingers, as these score highly (with positive and negative sign) with the first three eigenvectors, and are therefore partially responsible for the dispersion of the populations along the axes. Thus, a contrast between arches and whorls can be observed in the first axis. A contrast between right radial and left ulnar loops combined and whorls, is also apparent in the second axis. For the third eigenvector, the contrast is between left ulnar loops, right radial loops and right whorls, on the one side, and right ulnar loops, left radial loops and left whorls, on the other. With regard to the scattering of the male series within the factorial space, those from El Bierzo (BIEj and La Maragateria (MAR; on the opposite

DERMATOGLYPHICS IN ALBERCHEFORMES VALLEY (SPAIN) 31 1

TABLE 6. Finger dermatoglyphic variables in males and femolrs rrom Alberche 1 Tormes valley compared uith Iberian Peninsula series

AlbercheEormes Males Females

Arches 3.32 4.29 Radial loops 4.12 3.06 Ulnar loops 58.82 61.19 Whorls 33.74 31.46 Pattern Intensity Index 13.15 12.77

Iberian male series Min. Max. Average

2.18 10.28 6.27 2.50 9.37 4.59

48.59 67.19 59.05 21.58 40.20 31.01 11.50 13.59 12.63

Iberian female series Min. Max. Average

2.06 10.10 6.86 2.38 7.20 4.07

52.20 73.80 61.38 14.40 38.20 27.67 10.56 13.22 12.13

side) are the most extreme along the first axis, and the Caceres (CAC) and Badajoz (BAD) series are the most separated along the second axis (Fig. 2, left upper side). The third axis separates the series from the whole region of the Sierra de Gredos (GRE) from all other series (Fig. 2, right upper side).

In females (Fig. 3, bottom), the first eigenvector is characterized by a contrast between whorls in general and a combination of ulnar loops and arches. Ulnar loops (especially those appearing on left middle ILU31 and little fingers [LU5]) and arches, in general, are separated by means of the second eigenvector. The third eigenvector is more difficult to define, but a contrast between pattern types on thumbs and middle fingers is ob- servable (e.g., ulnar loops of right [RUl] and left thumbs [Lul l versus radial loops of left thumb [LRl]; radial and ulnar loops of right TRR31 and left middle ILU31 fingers, respectively, versus radial loops of left middle fingers [LR31). The compared female series are arranged within the factorial space of the first and second eigenvectors in a band along the first axis, except the series from Badajoz (BAD) and the whole region of Sierra de Gredos (GRE) which are clearly separated by the second axis (Fig. 3, left upper side). The third axis (Fig. 3, right upper side) separates the series from El Bierzo (BIE) and Madrid (MAD) from all other series.

No clear geographic pattern in the ar- rangement of population samples is observed either in males or females. Rather, the samples tend t o be scattered within the factorial space according to the similarity of the frequency of finger patterns or combinations thereof. For instance, the male series of Tietar valley (TIE) and La Maragateria (MAR), which are, respectively, in the southern and northern central plateaus, appear close to each other in the factorial space de- fined by vectors 1 and 2. This is largely due to the low value of the A/W ratio for the

ring fingers which is explained by their high frequency of whorls (see Fig. 2, left side).

The dermatoglyphic relationships of the male series from Alberchemormes (ALT) to all other series (Fig. 2 upper side), reveal it to be close to the Caceres (CAC) series and to that drawn from the entire Sierra de Gredos region (GRE) for the first two eigenvectors (Fig. 2, left upper side). Nevertheless, the third eigenvector clearly separates the entire Sierra de Gredos series (GRE) from those of both ALT and CAC (Fig. 2, right upper side).

It is difficult to discern which dermatoglyphic patterns are responsible for these relationships. According to the scores of those populations and the variables with the three scaled eigenvectors (see Fig. 2), whorls of right little finger (RW5) parsimoniously characterize both the ALT and CAC series, as do whorls of left thumb (LW1) in the entire Sierra de Gredos series (GRE).

It can also be observed in Figure 2, that the southern series from the Sierra de Gredos, Tietar valley (TIE), is distinguished to a cer- tain degree from the northern series Alber- che/Tormes (ALT). Considering the scores of the variables (finger patterns) with the three eigenvectors, this different position within the factorial space is specifically due to the different incidence of whorls in right little fingers (RW5), which characterize the ALT series, and of whorls in left ring finger (LW4), which characterize the TIE series. The scores of these patterns with the first three eigenvectors are those which more closely correlate with the corresponding scores of the previously mentioned series. Notwith- standing their different locations in the factorial space, statistically significant differences between the ALT and TIE male series are not evident in either the incidence of fingertip patterns on the 10 fingers consid- ered as a whole, or in the incidence on each finger separately.

In spite of this statistical similarity, a com-

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J. MART" ET AL.

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Scaled eigenvector I 128 87%/

Fig. 2. R-matrix analysis of finger pattern types in 10 male series from the Spanish Central Plateau and Extrcmadura. Distribution of populations (A) and variables (B) along the first three scaled eigenvectors. Codes used for populations are those of Figure 1B. Those of

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Scaled eigenvector I (28.81% I

the northern and southern slope series from Sierra de Gredos are within boxes. Codes for variables: 1st letter: Hand, R: Right, L: Left; 2nd letter: Pattern type, A: Arches, R: Radial loops, U: Ulnar loops, W: Whorls. The number refers to the finger (1, thumb; 5 , little finger).

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DERMATOGLYPHICS IN ALBERCHEflORMES VALLEY (SPAIN)

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Fig. 3. K-matrix analysis of finger pattern types in 10 female series from the Spanish Central Plateau and Extremadura. Distribution of populations (A) and variables (B) along the first three scaled eigenvectors. Codes used for populations are those of Figure 1B. l'hose of

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the northern and southern slope series from Sierra de Gredos are within boxes. Codes for variables: 1st letter: Hand, R Right, L: Left; 2nd letter: Pattern type, A Arches, R: Radial loops, U: Ulnar loops, W: Whorls. The number refers to the finger (1, thumb; 5, little finger).

31 4 J. MART" ET At

parison of the results of Martin et al. (1993) for the incidence of finger figures in Tietar valley with those obtained in the present study reveals that the asymmetry (bimanual differences) differs between the two male series. This is a measure of which hand, or finger of a hand, has a greater incidence of a given fingertip pattern than its homologue. For instance, considering the overall incidence of digital pattern types, while arches are more frequent in left hands and radial loops are more frequent in right hands in males from the ALT series, the opposite be- haviour is observed in males from the TIE series. The asymmetry pattern for ulnar loops and whorls is essentially the same in both ALT and TIE valleys. It is important to point out that bimanual differences are statistically significant in both valleys (ALT, x2 = 15.33,3d.f.,p = 0 . 0 0 1 5 ; T I E , ~ ~ = 8.92, 3 d.f., p = 0.03). Likewise, a more detailed analysis of the pattern of asymmetry in ALT and TIE male series for the incidence of fingertip figures (i.e., bimanual differences in each pair of homologous fingers) shows that the asymmetry pattern is different in the two series and for all pairs of fingers. Several studies suggest that dermatoglyphic asymmetry has a genetic component (Jantz, 1975, 1979; Martin et al., 1982). If this is so, the different pattern of asymmetry for the incidence of finger pattern types observed in the male series of the two Sierra de Gredos samples (ALT and TIE) could be a reflection of slight genetic (dermatoglyphic) differences between the two male series.

The dermatoglyphic relationships among the female series (Fig. 3) clearly differ from those found in the male series. In this case, the AlberchePTormes sample (ALT) does not clearly cluster with any other female series, although the UPGMA cluster analysis of the matrix of Prevosti distances (Prevosti et al., 1975) among populations (data not shown), locates the ALT and Badajoz (BAD) series in their own separate cluster. This relationship (ALT and BAD), based on the plots of the finger pattern dispersion along the three eigenvectors, could be ascribed to the whorls of right and left index fingers (RW2, LW2) and to the whorls of left ring fingers (LW4; see Fig. 3).

The divergence between the series from the northern (ALT) and southern (TIE) slope of Sierra de Gredos in the factorial space of females (Fig. 3) is clearer than that observed in males. In fact, the overall incidence of

finger pattern types differs significantly between ALT and TIE series (X2 = 8.78, d.f. = 3, p = 0.03). Using the plots of finger pattern dispersion to assist in the interpre- tation of the dermatoglyphic differences between the female series from northern (ALT) and southern (TIE) slopes of Sierra de Gredos (compare the plots of the top and bottom of Fig. 3) , it can be seen that the Alberchernormes (ALT) series is distinguished by the frequency of whorls in left ring (I,W4) and in right and left index fingers (RW2, LW2), while the series from Tietar valley (TIE) is distinguished by the frequency of ulnar loops, especially those of left thumbs (LU1).

Although it is difficult to account for all of the dermatoglyphic divergence in males and females of the ALT and TIE series, such differences would be expected from the pattern and extent of sexual dimorphism of finger patterns among populations. The different distributions of pattern types usually observed between the sexes would lead us to make different interpretations for males and females when using dermatoglyphics t o investigate population relationships (Meier, 1991). Moreover, the observation that in the present study only male series exhibit dermatoglyphic similarities for finger pattern types could partially be due to a small proportion of males (2.5%) born in the Alberche/ Tormes valley that marry and make their home on the southern slope (Fuster, personal communication, data from parish marriage records, Sierra de Gredos 1850-present). As the samples analyzed in the present study and in that ofMartin et al. (1993) from Tietar valley consist of school children, then those males who are sons of fathers who had mi- grated from the northern slope carry the Y chromosome characteristic of the northern slope population. Jantz and Hunt (1986) have demonstrated the influence of the Y chromosome on pattern intensity, a conve- nient measure ofvariability in finger pattern types. Such influence could explain, in part, the similarity between the male series from both slopes of Sierra de Gredos observed in this study.

It has recently been proposed that the Si- erra de Gredos could have acted as a biological barrier limiting gene flow between the populations living on its two slopes (Alber- chelI'ormes valley and Tietar valley). A study of marriage movements in the Sierra de Gredos region (Fuster et al., 1993) indicated

DERMATOGLYPHICS IN ALBERCHEKORMES VALLEY (SPAIN) 31 5

results compatible with the existence of a degree of reproductive isolation between the populations settled in Alberche/Tormes valley and in Tietar valley. Furthermore, Mesa et al. (1994), using blood group traits, have shown genetic differentiation between the two populations living in the northern and southern slopes of the Sierra de Gredos. The amount of genetic differentiation found by Mesa et al. (1994) is not easily comparable with that found for the same populations in the present study. However, given the widely held belief of the evolutionary stability of dermatoglyphic markers due either to poly- genic resistance to drift (Rothhammer et al., 1977; Froehlich and Giles, 19Sl>, or to the effect of stabilizing selection acting on mor- phogenetic processes (Jantz, 1987), a lesser differentiation between populations would be expected for dermatoglyphics than for simple genetic traits such as blood group markers. In any case, the results obtained in the present study are compatible to some extent with the existence of dermatoglyphic differentiation between the samples from the northern (ALT) and southern (TIE) slopes of Sierra de Gredos, especially for females. The dermatoglyphic differences for finger pattern types found between Alber- che/Tormes and Tietar populations, therefore, tend to support the prevailing hypothe- sis concerning the role of the Sierra de Gredos as a biological barrier to gene flow.

ACKNOWLEDGMENTS This paper was financially supported by

a DGICYT grant (PB88-0119, Ministerio de Educacion y Ciencia, Spain). The dermatoglyphic database used in this paper was com- piled with the financial help of DGICYT grant PB89-0220, Ministerio de Education y Ciencia, Spain. Thanks are also due to Dr. Philip Mason who reviewed the English ver- sion of the manuscript, and t o two anony- mous reviewers who made helpful comments.

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Documents

Dermatoglyphics of inhabitants of Alberche/Tormes valley (Sierra de Gredos-Central Spain): I. Finger pattern types and pattern intensity