Beals Brain Size and Climate PDF

Brain Size, Cranial Morphology, Climate, and Time Machines Kenneth L. Beals; Courtland L. Smith; Stephen M. Dodd Current Anthropology, Vol. 25, No. 3 (Jun., 1984), 301-330. Stable URL: htp//links jstor.org/sici?sict=0011-3204%28 198406%2925%3A3%3C301%3ABSCMCA%3E2.0,CO%3B2-7 Current Anthropology is currently published by The University of Chicago Press, ‘Your use of the ISTOR archive indicates your acceptance of JSTOR’s Terms and Conditions of Use, available at hhup:/www.jstororg/about/terms.hml. JSTOR’s Terms and Conditions of Use provides, in part, that unless you have obtained prior permission, you may not download an entire issue of a journal or multiple copies of articles, and you may use content in the JSTOR archive only for your personal, non-commercial use. Please contact the publisher regarding any further use of this work. Publisher contact information may be obtained at hup:/www jstor.org/jouralsuepress.himl, Each copy of any part of a JSTOR transmission must contain the same copyright notice that appears on the sereen or printed page of such transmission, STOR is an independent not-for-profit organization dedicated to creating and preserving a digital archive of scholarly journals, For more information regarding JSTOR, please contact jstor-info@ umich edu, upslwww jstor.org/ “Tuo Jan 13 13:49:17 2008Cconnenr aNTinorOLoGY Vol. 25, No 3, June 1988 Brain Size, Cranial Morphology, Climate, and Time Machines! by Kenneth L. Beals, Courtland L. Smith, and Stephen M. Dodd INCREASING CRANIAL CAPACITY has historically been associated with increasing complexity of society. The resultant ten- ddeney has been to think of humans with larger brains as mentally more capable. Gene-pool (racial affinity) and somatic (body size) explanations have also been advanced to account for the braincase variation. ‘We offer an alternative hypothesis that suggests that hominid ‘expansion into regions of cold climate produced change in head. shape. Such change in shape contributed to the increased era: nial volume. Bioclimatic effects directly upon bady size (and indirectly upon brain size) in combination with cranial glob- Uularity appear to be a fairly powerful explanation of ethnic soup differences. Within this hypothesis, the evolutionary tends of brachycephalization and encephalization are consi cred as functionally connected. This thermoregulatory model 7 Evaluation of data was supported by research funds from the Or- ‘gon State University Computer Center. David Fulrer and Robert ‘MeNaughton afed in cretion of the mapping program. Davi Frayer And Gerald Brash assisted in preparation of is. Clare Younger prov ‘ded invaluable asistance wit the suanuscript, We are indebted 10 Bennett Blumenbers for detailed and useful critique Keywert L, Beats & Professor of Anthropology at Oregon State University (Corvalli, Ore 97331, USA). Born in 1940, he was fcueted atthe University of Oklahoma (B.A. 1966) M-A., 1968) Sind the Univer of Colorado Ph-D.. 1971} His research terest ‘Shuman vartation. His publieatonsipclade “Head Form and Clr mati Sires" American Journel of Physical Anthropology 37:85 52) with A. J. Keo, “Genetic Diversity and Cultural Evolution” (American dnibropologit 7756679) and, with Timothy Baugh, ‘Biocuturl Bvolution Mlnneapolis: Burgess, 1978). CCounruanp 1, Sirti alco Profesor of Anthropology at Oregon State Univers: Horn in 1989, he received his BME from Bens. selaer Polytecnic Institute in 1961 and his PhD. trom the Ua erst of Arizona in 1968." His research interests are human ‘Sitpltion and computer-aided instruction, Map oulines and plot. ting of points for Bguresineuded inthis paper were adapted from Instructional computer programs developed at Oregon State. Tis Publications incude Salmon Fishers of he Columbia (Corals: Gregan State University Pres, 1979) and “Human Behavior In- arporation Into Ecological Computer Simulations” Emeivonmen- tat Aanogement 6:251-00) ‘Stari M. Dopp was both in 1947, After receiving hie B:Se from Oregon State University in 1975, be graduated from the Unt ‘vers of British Columbia (MCA, 193) His researc specialties le linguistics and computer modeling He is currently engaged in Seldwork with urban Arabic and Muslin populations i the United States and Canada, “The present paper was submited in final form 24 X 83 Vol. 25 » No, 5 > June 1984 is taken not as exclusionary or competitive with other approaches but rather as an adjunct toward understanding the Aistibution of cranial morphology over time and space. Anthropometric distributions are importantly affected by li- matic adaptation. Examples of investigations, reviews, and discussion include Thomson (1913), Roberts (1953, 1978), Wie- ner (1954), Coon (1983, 1968), Newman (1953, 1961), Baker (1960), Schreider (1964), Hiernaux (1968), Wolpotf (1968), Steegman (1970, 1975), Beals (1972), Koertvelyessy (1972), and Crognier (1981), Traits with thermoregulatory associations in- June 1984 Beals, Smith, and Dodd: CRANIAL CAPACITY AND CLIMATE rubber bladder into which air could be pumped. Endocranial volume, then, isthe result not simply of body size but also of cranial shape. ‘To our knowledge, the first suggestion that such morphology isa reflection of thermoregulation was given by Coon (1955:296): “Ttis easier to keep a small head cool than a large one. Witness the extreme dolicocephaly of hot-country peoples. In regions of great cold a large head is at an advantage from this point of view, as isa round one” From a geophysical perspective, the energy available to flora and fauna basically depends upon the earth's inclination to the sun. As high-energy photons of| solar radiation decrease, the body and cranium must become ‘more energy-conserving, Innovations such as specialized tools and controlled use of fire permitted occupation of areas of lower solar radiation and thereby set in motion a series of physiological and anatomical changes. Such trends of ecotypic differentiation should be observable inthe fossil record—at least since the first significant exposure to winter frost (approximately a half-million years ago. “The selective mechanism capable of producing the required sifferential reproduction is a thermodynamic hfe isis, Tis not fa matter of day-to-day comfort. The most obvious causes of ‘death are hypothermia (exposure”) and heat stroke. These have probably always been relatively infrequent as a percent- ‘age of the total death rate. Thermoregulation, however, plays ‘contributory role within a spectrum of criss situations such asshock, drowning, and traumatic injury: The same inventions (4, reindeer herding) that allowed occupation of regions other than the tropical savannah of origin may also have increased ‘the probability of death in which thermoregulation plays a part (Steegman 1975). ‘This brief summary of explanatory models cannot convey more than a general outline. Critiques of the use of brain size in typology have been offered by Gould (1978, 1981). Tobias (1971) has reviewed the evolutionary evidence. Brengelmann and Brown (1965) have summarized physiological aspects of thermoregulation. General treatments of human bioclimatol- ‘gy occur in Coon (1965) and Flach (1981) ‘Our focus is the bioclimatic model, and the investigation suggests that approximately 30~40% of the variance in population means can be attributed to thermoregulation. The obvious question is, what explains the remainder? Part of the complexity is that all of the explanatory approaches (including. ‘our own) involve elements that produce nonsystematic variance and therefore complicate any general interpretation, Among, them are statistical “noise” from measurement and sampling error, local circumstance (ie.,a famine that affects body size), stochastic genetic events affecting geographical distri and inventions that alter relative death rates, CRANIAL CAPACITY AND CLIMATE, ‘A summary of data on endocranial volume is given in table ‘The distribution forms a normal curve that is mesokurtic and slightly negatively skewed, Our averages for volume are some- ‘what less than the 1,400 cm frequently cited as typical of modern humans. The latter figure historically derives from not Adjusting the shot method and often considering Europeans ot ‘males as the model, We mention this because the magnitude fof the difference is sufficient to affect interpretations of the rate of change over time. As in physiology, itis convenient to have a standard of an “average” person. The physiologists’ standard human is a re- ‘ection of their most common research subject—an adult male ‘of European descent, with a weight of 70 ky and surface area ‘of 1,73 m'—who generates energy at the approximate rate of 885 keal/hour when sitting. This is an output similar to that of 305TABLE 1 Mean Caaniat Caracrriss (6m!) FoR 122 BrNic GuovPs Sours 3 Rance Maes war Females, un Combined 1389 1085-1881 Dimorpiem iss 20-276 ook B Smewi-s) ms 10 01 0s sto 4902 00. bundred-watt light bulb (keal = 1.16 watt). Normal daily heat loss is 16 kwh. OF this, only 4 kwh is replaced by food ‘metabolism, and the remainder must be met by some combination of insulation (clothing) and atmospheric energy. The amount of atmospheric radiation available in combination with ‘worldwide temperature and humidity is largely’ funetion of latitude and varies from 557 eal per em! per min between 0 and 10° N to 310 cal per em? per min between 60 and 90° N (Flach 1981), Our heterographic (in contrast to physiological) standard “human” represents the sex-combined world average under all types of climate, with each climatic zone given equal weight.” ie/she weighs 54.1 kgand hasa stature of 187 em and asurface area of 1.525 m*. This corresponds to a mass-area ratio of 35 kg per m:, with an endocranial volume of 885 em’ per m of surface atea, 24.9 cm’ per kg of weight, 8.6 cm per cm of stature, 17-8 em! per unit of cephalic index, and 32.4 em’ per unit of ponderal index. The typical buman has for each cubic centimeter of brain mass 11.45 cm of total body radiation! conduction/convection surface. (Dural contribution is approx- ‘mately 50 em» but is closely matched by shrinkage ofthe dried cranium.) Table ?tabulates heterographic data in traditional fashion— by continental area. If one merely lists such means by ge0- eraphical region or race, causes of similarity by genogroup and ecotype" are hopelessly confounded. To illustrate, the percentage (FCIN) i given of samples within each continental area which also happen to be exposed to significant winter frost (Wemperate, wet cold, and dry cold areas). For example, 73% ‘of the samples from Asia are native to areas of winter frost, ‘compared with 100% of those from Europe. The correlation is (0.91 # 0.08. This simple factor alone statistically explains 839% ‘ofthe variance in capacity between major geographical regions. For the last column of table 2, we have used the resulting regression to predict the continental means. Comparisons are close; the average difference from actual observations is only V7 em’ ‘The implication is that any effort to attribute racial or cognitive significance to brain size is probably meaningless unless the effect of climate is controlled. For example, the endocranial ‘volumes of Europeans and Africans differ little from what one would expect given the difference in their respective winters, RELATION OF CRANIAL CAPACITY 70 I$ COMPONENTS ‘The volume of the brain container is obviously a function of its dimensions and geometry. Increasing vault height and breadth Grade averages, such as means of data tables, are usually dispro- postionaely representative of particular regions or groups. For in Stance, the cranial fle disproportionately represents North America because of the exhaustive catalogs of Frdigka. In order to have consistent and objective standard of comparison, we calculate sex ‘combined means for each climatic zone and then give equal weight to tach zone. "The result c's average morphology under ll conditions ‘otelmate ™ Beotypes are statistical agereates associated with particular en vironmental conditions, such ts climate. Genogroups are populations Classted by common genetic heritage. ‘The distinction 1 aml to that between analogous und homologous Variation, 306 relative to length thus increases capacity, Empirical relations between external dimensions and container volume relate to the time-machine project, since, if partial data are availabe, more reference points through time may be determined by prediction. The climatic file was used to correlate data with ‘composite means of length, breadth, height, and module. A discriminant function indicated that the greatest contribution to the volume derives from breadth, followed by length and height. Intercorrelations are shown in table 3. ‘The matrix illustrates the differing geometries of cranial size (module) and brain size (endocranial volume). The latter is primarily determined by breadth. To simplify, the proximate reason some srroups have larger brains is that their heads are broader. While some of the increase in volume is due to a larger head (which in tum is due to a larger body—which in turn is partially due to thermoregulation), another portion derives from increased slobularty of the container, again partially attributable to ther ‘moregulation, with breadth playing the primary role. In one sense, a larger brain can be explained geometrically ‘One might speak of brain size as being biophysical, while brain function is biocultural. In a larger perspective, there is no single cause of hominid encephalization, but rather an interplay of total ecology involving the magnitude of solar radiation, the principles of thermodynamics, and cultural innovations which Jed to adaptation within new econiches ‘A multiple regression was calculated between volume and external measurement, em! = 403.9 + (80.6 B) + 42.81) ~ 0.3), ‘hich has a multiple X of 0.82 and applies to sex-combined TABLE? Sex-ConpineD MEAN CeaNtat CaPactrits (4) rox ‘CONFINENTAL AREAS CoMPank rr PmEDeT# VALUES Basko on Cuistarte Zowt* Recon N 2 @ Tom Papo Non America = Tom 1,66 Asi % 07s 361 Europe 10 351001304 South America 2 420801388 Oceania a oso 39 Aiea 0 010 1st 22 + aren, TABLES Copnsiarions arrwkEN Contnostr® MEANS oF CRANtal Lexcri, Bueabrn, Hiscitr, MODUL, ax CaPactr Taare Tenor BaEapra Hrcnr Moour Baeanre on Hecnr os 0 Moouue os: Os? Ob Cancrry 036 ome 8contemporary ethnic groups, Interestingly, when climatic zone ‘was incorporated into the analysis, it made a greater contribution to the variance than either length oF height. Forthe time machine, more reference points may be obtained by satisfactorily predicting the cranial capacity of fragmentary specimens (if breadth is known) by including the climate from Which the specimen originates. Beyond this, the extent of geometric influence upon volume leads us to reconsider the generally presumed taxonomic significance of brain-sze difference between contemporary hominids, such as H.habilis and Aus- tralopithecus. The question is whether this difference isa variation that has behavioral significance (which in turn may or may not have reproductive-solation meaning) ofa slight variation in cranial geometry. Among present-day groups, large differences inthe capacity of the container are known to have no reproductive-isolation consequence, They result instead from small differences in absolute dimensions ‘Correlations were calculated amang all the climatic and an thropometric variables; a summary of linear relations between capacity and other traits is shown in table 4. Overall patterns between the size (volume) and shape (cephalic index) ae vie- ‘ually identical; they inerease together, increase with weight and surface area, deerease with nasal index, and are only weakly associated with stature (CRANIAL CaPacttY AND CLimarie ZONE ‘The basic test of bioclimatic theory is comparison of population ‘means in regions exposed to winter frost (temperate, wet cold, and dry cold regions) with those from regions of dry or wet heat. Table 5 contains the summary from the cranial file. The ‘Beals, Smith, ond Dodd CRANIAL. CAPACIEY AND CLMATE slobal mean for populations in temperate and cold climates is 1,386 + 6.7, while that for hot-climate populations is 1,297 = 10.5. The absolute difference of 89 cm is highly significant (= 7.5, 9 = <0,0001), The lower variance of temperate/cold soups is also significant (F = 1.69). The same pattern of the ‘means oecurs within each continental area; there are exceptions: tothe rule with individual groups, but the means are invariably’ higher for temperateicold cases within each geographical di vision, Figure 4 shows climatic zones as based upon general ized, predominant types of thermal stess. Figure illustrates ‘the trends which result from plotting the means for each cl ‘matic zone separately. Table 6 summarizes correlations of eli ‘matic zore with 11 morphological traits. Head morphology in size, shape, and nasal form is more closely related to climate than is the body as a whole. (CRANIAL Caractry AND LatrTupE Grid coordinates in the hominid file are supplied for each ste. By taking selected segments of time, its then possible to eval uate spatial trends which may be’ helpful in predicting the required data points for the clinal maps. This allows spatial ‘comparisons Detween the past and the present. A general fea ture of hominid evolution has been occupation of the globe beyond the tropical savannah of origin, Te bioclimatic model predicts that cranial capacity will increase with distance from the equator—latitude being correlated with a decrease in solar radiation. Latitude is actually intercorrelated with a number of climatic conditions, relationships of which produce a high TABLES Constaarions nerwees: Composite Mraxs oF Castat Caracrry, Weten, SFATURE, SURFACE AREA, CRANIAL INDEX, AND NASAL INDEX! Crane Caeser Waonr 068 oo oon o.00 “oe ‘ooo 0.024) Senrace Anea, CCaastat. sex Srarone Figures In parents are snfiance levee Sunrack Grainy Nasat Wercnr AREA” Isa Tbe coon 056° 0.26 (008) (0%) “oss 085) 0.40 (oo 00 (000 001) 90) 0.362) (1008 TABLE s DDisrausuvios oF Sex-Coumiven MEAN CRANIAL Caracrriss (68) FOR ETHNIC (GzoUrS BY CONTINENTAL AREA IN Winter Frost Cumarie Zone Wel or Dy Heat “Teperate or Cold Recios wt «of Nk of Oia Won Europe 10 son Africa ° as as Asa 7 aw sie Ocean 18 ow 8 a 8 "Total o mo 38 oon New World "North America 0 sows ss 10 South America. ‘ Bis 6 21s "Total 16 no 3 39 Grand Total 50 m0 2 a7 Vol. 25 + No. 3 + June 1986correlation with capacity (7 = 0.62, 6 = 0.00001). On a global scale, each degree of equatorial distance adds 2.5 cm’ to the volume, Volume as a funetion of latitude is shown in figure 6. ‘This scattergram also indicates one of the reasons the validity ‘of the record low report among the Akka is statistically ‘questionable Since Oceania, the New World, and the Old World have had different oceupation patterns over time, latitude associations ‘within them were examined. Some of the comparative data are shown in table 7, As anticipated, rates in different parts of the ‘world vary according to their culture histories. The highest slope of 3.1 cm" per degree of distance from the equator is found within the Aftican-Burasian landmass, which has long been occupied by hominids. As also expected, the association israndom within Oceania, where occupation is recent and there is little cold stress ‘The Americas provide a unique test ofthe theory, since there is a known point of origin (the Bering Strait), a known period ‘of adaptation, and a known direction of dispersion toward the ‘equator and through a funnel of tropical forest in Central Amer: ica). These circumstances predict that the mean of the trait will be higher in America, the point of regression origin from the equator higher, and the slope of the regression lower, All three of these are empirically observable ‘American data also indicate that braincase volume can change rapidly in response to climatic conditions, The slope from the equator to a distance equivalent to the Bering Strait (65°) amounts to an average difference of almost 100 em’. Assuming Beal, Smith, and Dodd: CRAWtAL. CAPACITY AWD CLIMATE ‘occupation of the various ecozones, then the systematic adaptive pattern observed in the clinal maps isa reflection of only 530,000 years" development. Tis implies a rate of change greater than 3,000 em? per million years CRANIAL MORPHOLOGY AND CLIMATE During the general course of hominid evolution, the cranium tends simultaneously toward both larger size and rounder shape: Even if size remains the same, volume increases as the ratio of length to breadth decreases. The two trends together ean be considered a trend toward globularty. Reduction of the browridges may also be a part of the process—large ridges increase surface area. The overall effect produces a simpler land more regular cranial topology, along with a more pedo- morphic appearance. Perhaps the morphological complex sometimes attributed to neotenous mutation may be more a {question of biophysics, ‘Atany rate, cephalic index and cranial capacity are expected to have interactive effects. If 0, the correlation of both with climate should be higher than with each separately. Their mul tiple regression is, CZ = 16.4 + 0.0088 (CO) + 0.091 CD, and there isa significant additive effect, with a multiple R of an nial entry of 95,000 Br and a 3000," pertd for 069 ed | | Intercept = 1257.3 | t b> 2.501 : i Standard ercor of b = 0.286 | wenn f Sigatiicance of © = 0.00001 . ! melt or ee | io. took ut i ” | . aan . re . { io | rel : t } Fic. 6, Didribton of sexcombined mean canal capaci smong 122 population as a fanction of distance fom the equator, Axe ar canal capacity (cm) and absolute degrees north or south latitude. Question mak refers to Akka report. Numbers on scattergram are multiple cases, TABLE? (Crania CaPAcrrY (Ch) AND LATITUDE WITHIN MAjon Wonk REGIONS Rocron w New World s Oceania, 21 1d Wortd 6 (Old World + Oceania or o > ? 12328 5.059 0.76, 1asst2aee 0.68 [Nore N, numberof mens; 0, on a eqtr 8 slaps frm equator: 7 cote of Vol. 25 + No. 3» June 1988 309While multiple regression allows for interaction between size and shape, itis desirable to plot a composite value which at least crudely corresponds to one's visual perception of a spec imen. We are, however, limited to the cephalic index and vol- lume as imperfect measures ofthe actual morphology: Asa point of reference, we return to our heterographic standard human. With each climatic zone having the same weight, the global mean cephalic index is 78.0. The cranial and cephalic indices are not, however, identical, again primarily because of ge- fometz: Since the head is oval rather than spherical, removal ‘of approximately equal tissue from the circumference lowers the length:breadth ratio (Krantz 1980a). On the average, the cranial index is about 1.5 units lower. Subtracting this mag: nitude leaves 76.5 as the standard for comparison with the fossil record. The analogous reference point for volume is 1,383 Visual appearance is a matter of cognition. For simplicity wwe give size and shape equal weight and calculate a coefficient of eranial morphology COM = Ys ((obs.CC11,383) + (obs.C1176.8)] =1 When length:breadth ratio is combined with volume, their in dividual conteibutions to the visually perceived morphology are lost; individual components are therefore included in the data files. The coefficient is standardized to a mean of zero, and differences from the mean are percentage values from & typical modern human under all types of climate. Positive val- lies should be astociated with cold environments, negative val tues with hot ones, The coefficient can also be used as simple description without any climatic implications. For instance, a baseball has a coefficient of 0.27, a slow-piteh softball 0.01, 8 volleyball 1.45. More tothe point, the lowest coefficient from 4 Pleistocene adult for which we have information is Stezk fontein § (—0.88) and the highest Grotte des Enfants 4 0.13), ‘This merely indicates a range for purposes of comparison; any climatic implication requires matching with reasonably’ corresponding time. Tn the paleontological record as a whole, more relative change thas occurred with volume than with shape. The lowest cranial index for an adult known to us is Sangiran 4 (62.8). It has a ratio to the heterographic standard human of 1.22, whereas the corresponding ratio for cranial capacity (Sterkfontein 60, 428 em/1,383 em’) is 3.16, For some reason, however, there 60S 25W is a reversal of these relationships in the comparison between the Upper Paleolithic and the heterographic standard human, in which volume deereases while roundness increases dramat. ically. For modern populations, the coeficient should vary by ecological adaptation, The range is from the Vedda (—0.10) to the Buriat (0.11). Figure 7 illustrates the distribution, [Bran Size AND CLIMATIC VARIABLES Highs and lows (normal and means of the seven climatic vari ables for the 82 populations in the climatic fle are shown in table §. Kikuyu data are selected as representative of a current tropical savannah, Tropical savannahs are relatively uniform, with more of a wet/dry seasonal difference than a summer! winter one, They represent the climatic ecology of the ancestral hominid homeland. Most present-day populations are exposed to lower winter temperature and lower vapor pressure, and these latter two factors might be anticipated to have the highest correlations with contemporary anthropometric means. The correlation matrix is given in table 9 ‘Climatic influence on relative brain size is likely to be more interesting than that on the absolute value. Table 10 surne rizes the available data on the distribution of braincase volume relative to weight, stature, and surface area. Groups with large volumes per unit of mass include San (33.4 em’ per kg), An- damanese (27.6), and Bengali (27.8). Groups with small vol umes per unit of mass include Choctaw (20.8), Freneh (22.0), ‘Mapuchi (22.1), and Maori 22.1). An overall relation between cranial capacity and body mass is clear from these examples, Brain size in relation to weight follows the mammalian pattern AAs previously mentioned, the greater the body weight, the ‘smaller the relative volume of the cranium." The linear eor- relation of weight with cranial capacity is 0.63 0.10; the corzelation of eranial capacity with cranial capacity: weight ra tiois ~0.16 = 0.16. Incidentally, greatest mass is not an aretic phenomenon. The empirical model for extreme cold is mod trate weight, moderate stature, moderate nasal index, mod: erate brain size per unit of weight, but large absolute cranial capacity, large cranial capacity per unit of stature, round cra- 7 Conventional interpretation of brain weight (2) to body weight (his te llometrie relation £ = KP. However, Martins (981) ‘work indicates taxonomically variable slopes. For placental mammals, Fisregression itlote/) = 0.76 logy) 1.77, with En miligrams snd Pin prams Fic, 7. Coofcient of cranial morphology at heterographic preset, Cs intervals are percentage of difference from world mean set at 20 Black, 10-148; checkerboard, §-98 cosshatchi si0 (0-45; horizontal siping, O-(~A¥e} dots, S(O) sole, — 10-4 ~ 10)TABU Vauues or Cuisearte Vans Ba ues ron Porutarions i rie Cemtaric Fite Vantante Solar radiation teal per em 220 (Nubians) Sunshine annual hous) 4,200 (Nubians) ‘Winter vapor pressure (mb) 132 Veda) Summer vapor presure (mb) 53 (Maya) “Anmual precipitation (em) 450 (Andamanese) ‘Winter temperature C) 29 Tucano) Summer temperature °C) 35 (Papago) Mian For Grour —_‘Trorieat Nowseat Cosirostra® SAVANNA Mirae rs 750 70 (Yabsan) 2268 2.000 1,500 Mapuche) 15 25 (Eskimo) is 2 (Chaiken 18 109 Sinan) . 18 = 38 (Yakut) 2 26 6 (Siberian Yuit) ‘Ta Connetarions nerweew InpicaTors oF CxaNtat, MonoiioLoc avn Ivnivinuat CLasaric VARIABLES yee SR ARS WP SOP SCSPRESSCWTM~SCSTM CS ce @ oes —08%3 08% —0sI0 040) 0597 ——OGH aR? OD (oon (00H) (002) oOo (008) @.0) OOOH cost or ‘owes oss) ower owoe “ost aris, ot! oat! 0.836 (001) 00.001). OD OKO OD!) oH cowe sr 017s ‘ost “0380° ‘0.29902 a2s ‘atta 0.208, (0158) (006) (005) (Oost) (04s) (O42) 284) 19H (UBB) cess, sr ‘os ‘o'o6o—‘o.u29, ‘aortas ‘aso 0.218, 0.006 (0.068) 365) 0225) (0358) (0.20 (Ost) ION) 0.286) com 6 ‘oss, © 0610" 0.395 oss) 0482 018? 0.623" 0.426 oon (00) ‘o.oo 000.072) D000) cn ‘ase “os8 “ozs 028 ‘oa 0.390 ‘outst (001) 001) 0.028) (005) (aor (0.399) 00) _—_—@.10H terse C, canal capaci Sy, slture; We, web. SA, sure ares nium, and low surface area:mass ratio, Brain size relative to stature has significant associations with all of the climatic variables. It also has the highest corcelation with winter temperature (r = ~0.64 * 0.07) of any ofthe six cranial variables. Groups with high ratios include Aleut (0.8 em’ per em), Es- kimo (9.8), Yakut (9.6), and Yukaghir (9.6), Groups with low ratios include Australians (7.7), Nubians (7-4), and Sinhalese (7.8) The ratio isa good indicator of climatic conditions, and Wwe assume that a large endocranial volume in combination with moderate to short stature would be particularly indicative ‘of cold adaptation during the Pleistocane—as is indeed observed among Glacial Neandertals. Figure & depicts geographical variation for the heterographic present. The Old World has a striking: southwest-northeast cline, while New World variation is more regular with distance from the equator. Ex- tremely low values around the East African Hor are consistent with the world's greatest physiological heat stress. limate is a multivariate phenomenon, and questions arise with regard to the relative importance of its components, In brain size normally refers to brainbody-weight ratio. a5 used here includes a greater numberof comparisons, Le We to surface area, weight, and statureeach of ble group data for diecly measured surface area are witlly Bon {xistent. Toles of brain-body relations that we consider beyond out present scope include lean body mas, iferental body compotion, brain weightendoerania-vlume coreation, and funcional signi tance of the neurology. Discusion of surface area calculation, mets Sole rate, ad lean body macs in given by Broven and Brengclmann (96s) Vol. 25 + No. 3 + June 1988 (Coa, cuter of eral morphology CI, cephalic inex " the general summary of table 9, volume (CC) has higher correlations than shape (CD, but there i sufficient interaction to produce the highest associations known in the coefficient of, cranial morphology. Generalized classifications (climatic zone and isothermic zone) tend to have higher correlations with the ‘morphology than do individual climatic variables. Climatic zone produces the highest correlations with the traits and is also the most applicable to the fossil record. With respect to Doth temperature and vapor pressure, winter conditions are ‘more important than those of the summer. We assume that annual precipitation has no morphological effect in itself and that the occasional significant correlations with the anthro- pometrics are attributable to synergistic relationships between precipitation, temperature, and vapor pressure. There is litle Alfference in'the associations between vapor pressure of the summer and winter, As one would anticipate from the dispersion pattern of hominids, major types of adaptations are to TABLE 10 (CaantaL Caractry (CO) RELATIVE T0 WEIGHT, STATURE, AND ‘Somrace Anza Recaro N 8 Rave @ oP Cowes Wwa7e tosase aS 0a CCStature oko 7498 as od CCsSurlae ate. S7815.59 769.0-1,0000 5406.2 Data base on popalation means * Cate of waiaton snreduced energy from the sun, lower absolute humidity, and the rigors of a cold winter. SUMMARY 1. Variation in endocranial volume among ethnic groups is partially explicable by thermoregulation. Its significantly as- Sociated with every climatic variable examined and has the highest correlations of any single morphological trait consid- cred, Furthermore, the mechanism of thermodynamic life crisis relates the biophysics to differential reproduction, whichin part explains not only the present variation but also the trend of encephalizaton, 2. Average cranial capacity is not as great as is generally assumed. There ate historical reasons for ths; the larger figures of the past result primarily from not adjusting for the overestimation of Broca's measurement procedure. The world mean ‘depends on how one chooses to weight reports. We suggest 1,383 em? as an appropriate estimate. This reflects sex-com: bined ethnic groups under all conditions of climate 3. From a structural perspective, the greatest contribution to volume is from breadth. Different populations have diferent cranial geometries. Most simply stated, some groups have larger brains than others because their heads are rounder, Arctie peoples obtain large capacities not so much from large heads as from a more globular shape. The high correlation between breadth, climate, and absolute volume leads us to believe that if breadth can be obtained from fragmentary fossil specimens, cranial capacity can be reasonably estimated. ', As anticipated from conditions of solar energy, the brain container volume and latitude are highly correlated. The world average slope is 2.5 em? per degree of latitude, but the slope is substantially sharper in the Old World. Latitude associations are supported by the culture history of each continental area, 5. The evidence suggests that thermoregulation has more effect upon the cranium than upon the body as a whole. The highest correlations aceur with the coefficient of cranial morphology, absolute volume, and capacity relative to stature, Lower correlations are observed with surface area:mass ratio, cephalic index, nasal index, and ponderal index. Lower yet (Gut stl significant) are the correlations with weight and body surface area. Stature and cranial capacity relative to weight 60S 25W Fic. 8, Distribution of cranial capacty relative to statute (cn per cm). Black, 9.5-9.9; checkerboard, 9.0-9.4 rosshatching,8.5-8.9; hriontal striping, $0-8-4 dots, 5-79. 32 and surface area appear tohave but negligible associations with climate 6. Generalized climatic classifications usually have higher associations with anthropometries than specific variables. The strongest individual effects occur with solar radiation, winter temperature, and vapor pressure. Winter conditions are more important than those of the summer. The overall pattern fits ‘with hominid dispersion from a tropical savannah, 7. Wefind little support for the use of brain size in taxonomic assessment (other than with paleontological extremes overtime) ‘Racial taxonomies which include cranial capacity, head shape, ‘or any other trait influenced by climate confound ecotypic a phyletc causes, For Pleistocene hominids, we doubt that the ‘volume of the braincase is any more taxonomically “valuable” than any other trait. Ecotypic differentiation (fig. 9) appears sometimes greater than average taxonomic difference. A slight Increase in head size combined with a rounder cranium has a disproportionate effect upon volume. Even with absolute capacity difference, a connection to reproductive isolation is ques tonable given the lack of such connection among modern peoples. '8, The bioclimatic model provides a fairly powerful expla nation of several morphological traits. It likewise accounts for ‘4 portion of the trends toward brachyeephalization and en cephalization, We suspect that it may play’ a role in browridge reduction as well as, certainly, in the evolution of body size. Itis not, however, a full explanation of the paleontological trends. In the frst place, adaptation to cold is limited to approximately the last half-million years. Second, crania become ‘more capacious and rounder even among fossil ecotypes not exposed to winter frost (table 11, fig. 9), Climatic adaptation is apparently superimposed upon other causal mechanisms. It is possible that cognitive and somatic factors could account for 1 portion of the unexplained variance. If so, itis likely that the weight of climatic, somatic, and cognitive effects varies over time, We conjecture that prior to around 200,000 .P., encephalization was primarily the result of a combination of Selective advantage in mentallinguistic capacity and larger body size with associated energy efficiency. We further conjec: ture that within our own species (including Neandertals) cli- ‘matic factors have become the principal source ofthe variation 9, The explanation of human brain size difference has his torically been colored by a search for “the cause." This tradi-tionally focused upon difference in mental ability oF ace, Neither has been shown to have any significant direct effect. The dis tribution indicates that racial means are actually reflections of| secondary'correlation with climate. For example, Native Amer- icans have a common ancestry but almost the entire range of variation in cranial capacity. The cognitive model requires that mental function change not only the internal organization of the brain, but alo its absolute sie. It is not supported by any preponderance of direct evidence from either psychology oF ‘ethnology. Interpretations have more recently turned to body size, but ‘no measure of this explains more than 40% of the variance. “Metabolic rate as “the cause” cannot be directly evaluated for lack of ethnic group data. Vet given the association between ‘capacity and shape, the need for a multiple-factor interpreta: tion remains evident. Heterographie evidence supports Thom- son's (1903) almost ignored experimental work. ‘With an ever broader perspective, cognition is part of the ‘answer in an indirect manner—through cultural inventions Fic. 9, Postulated approsimate elfect of occupation of temperate and cold regions an coefficient of cranial morphology: Data to gradvalist snd alternative attribution ‘weighting systems generally produce less diferentiation. The bioclimatic model produces ambiguous interpret fable 11 and give equal right to ecotypie means. Letters 4 and B ‘Beals, Smith, and Dod CRANIAL CAPACITY AND CLIMATE. Which led to occupation of the world’s diversity of ecological zones. “The cause,” in short, does not exist. Explaining the variation in human brain size requires a synthetic theory, portions of which best apply to given particulars of time and space APPLICATION TO THE TIME MACHINE: HYPOTHESES AND INTERACTIONS, ‘To investigate the paleontological evidence, the combination of data processing technology and the unique format of CUR RENT ANTHROPOLOGY permit an interactive feedback with re- spondents, Within this section we attempt an experiment in Which the respondent is invited to select a problem, data set, and type of analysis. Within limits of response space, we wil apply files to the requested description, analysis, hypothesis, ‘or map—including whatever additions or corrections to the ‘reneRATE/cOLD Plotted from ‘a5 listed in appendix. Other jons for H. erectus and early ‘modern H-sapiens. Increase through tne occurs within the tropics as well as intemperate and cold regions and indicates that nonclimate factors are also required to explain the evolutionary trend. TABLE, Crain Monenotocy ay Taxon aNp Ecorvre tk Eanty Homo ‘Grau cree r00 a8 28S 1018 y0.0 1,200 ‘Temperate pa 133 Ghacal ma 00 ‘Bary moder H. sapiens “Tropica. tua art ‘Temperate my iam Glacial 75.0 1306 Vol. 25 + No. 3 + June 10988 ‘Move AureRNaTive MODEL car cr cc cM -020 8 997-017 Tos 735 99 os soo 1273-001 oot -o.01 313appendix might be obtained from feedback. We have misgiv- ings concerning limitations of funds, time, response length, and state of project development. Nonetheless, we consider it as a practical and interesting possibility to be explored. ‘There are severe limits on the nature of the evidence. Het- cerographic interpretations can be based upon thousands of specimens within an approximate 100-year span of time. Pa- leontological interpretations must be made upon scarcely 100 ‘eases spread over more than two million years. In addition, there are complications of reliability which result from recon struction, estimation of adult capacity from subadults, post- mortem deformation, dating error, and sexing error. Reliability is, however, a matter of degree and sometimes subjective judg- ment.” As a practical matter, the summaries and illustrations that follow are “total body of reported evidence.” All the cases in the appendix are inclided, since any particular inclusion’ ‘exclusion set may be specified. Some of the major questions of reliability are briefly noted in the appendix ‘Any particular taxonomic rearrangement may be chosen. In| table 11, morphology is tabulated by taxon and ecotype as a basis for comparison, There are two models. The first is "grad: Al nvestizations of cranial capacity including this one have re lability problems, «zy sampling and measurement error General, tania capacity value is more reliable than brain svlght (Bes 197 ‘There are time and location diferences between the skeletal obser ‘ations and the anthenpometrcs. All reports of eranial capacity ean bevregnded © population estimates only. To our knowledge, these fetors do not produce a systemati fleet upon the averallsaisteal Conclunons. Almajor facor limiting the rellabilty of paleontological ‘onclunons i smallness of sample sz relative to total population. For ‘ample, Westing’s (1981) estates imply thatthe entre hominid fle inthe appendix represents only one individual per $0 milion bor up {010,000 years ago. For the fesdback experiment, any static ‘weighting stem can be spesied ualisti" in the sense thatthe chronological sequence correlates more closely with taxon. The second model is derived from the ‘most common alternative attribution among disputed specimens. Major differences occur with a broad oF narrow concept of Neandertals, the antiquity of H. sapiens, and H. habilis as ‘a taxon separate from Australopithecus and H. erectus ‘The hominid data generally support the conclusions drawn {rom the study of ethnic groups. In table 11, a pattern of larger, rounder crania in colder climates is observable for both tax: ‘onomic models. The evidence is strongest among Neandertals, ‘more ambiguous among #1. erectus and early modern H. sa. pens. In figure 9 the coefficient of cranial morphology i plotted {or Tropical compared with Temperate/Glacial forms. Data are not adjusted for sex proportion or collective difference in time; however, this may be statistically corrected if desired. Figures 10 and 11 scatter cranial index and capacity by time without regard to taxon. For consistency, each is graphed on the same logarithmic scale of time in thousands of years 8.” ‘The resulting regression data are included in the iMustrations Lines of regression are omitted since they are not necessarily the best it for selected periods of time—within which rates of evolution vary. The time machine uses selected time segments rather than overall rates. In figure 12, the mapping program fs used to illstrate limits due to lack of data, unoccupied regions, and glaciation ‘Maps or associations may be taken from any of the files ‘mentioned (cranial, climatic, hominid, or HRAF). A variable Uist not within the files may’ be added but requires a convenient tabulation from the respondent. Funds are not available for analyses beyond programs to which we have access. Resources are presently lacking to provide analyses or maps beyond those associated with the present paper Besta Standatd error of b = 0.672 Fic. 10. Evolution of cranial index. The overall tend is geometec, with ahigh rate of increase during the Holocene, The heterographie compte i: 76.5, Among contemporary groups, the index has a lower association with climate than does cranial capacity. The converse may be true with fossll forms (able 11), With regard to climatic influence, the data for erectus follow the expected direction lower indies inthe opis), but ‘means are not significantly different. We assume that litle climatic deretiation with morphology had occurred at such an early date. The ‘model has no applicability to Lower Pleistocene forms, confined tothe tropics, The greatest ciference is observed between Glacial and Tropical [Neandertals n which the index-~adjsted by appropriate regression for Ume—is approximately units higher. The evidence indicates a decrease inthe index between the Middle and the Upper Paleolithic, and we have bec unable to explain this without a gene-Row model in regard to the "Neandertal Problem,” As with cranial capacity, climatic adaptation isfy succesful in explaining variation anong contemporary humans bat less so in explaining the phyetic te sia CURRENT ANTHROPOLOGY180E WH ts i pgGAPPENDIX: HOMDAT ‘The following isa list of hominid specimens, in chronological corder, for which values are available for either cranial index (Ci or cranial capacity (CC). Estimated dates are in thousands of years, Presumed sex (S)is indicated where possible. Climatic zone is coded as tropical (7R), temperate (FM), or glacial (GL) ‘Taxonomic codes are 4A, Australopithecus africanus; AR, A robustus; HH, Homo habilis; HE, H. erectus; N, Neandertal, Neandertaloid, archaic 1. sapiens; MM, early modern H. sa” ‘iens. Sources are coded as follows: A73, Aigner and Laughlin 1973; 476, Alexeyev 1976; B35, Von Bonin 1935; B50, Briggs 1950; B70, Brain 1970; B72, Brace, Nelson, Korn, and Brace 1979; B80, Billy 1980; Do2, Dart 1962; Dos, Day 1965; Ds0, Day, Leakey, and Magori 1980; #78, Frayer 1978; H151, Howell 1981; H72, Holloway 1972; 173, Holloway 1973; 178, Hol- loway 1978; 80, Holloway 1980; 130B, Holloway 19808; Hs1, Holloway 1981;.166, Jacob 1966;.J73, Jacob 1975; K70, Kelso 1970; £70, Leakey 1970; 172, Leakes, Mungai, and Walker 1972;173, Leakey 1973; 174, Leakey 1974;175, Lestrel 1975; Moz, MeKern and Kozlik 192; M74, Mann and Trin kaus 1974; ND, Neumann n.d.; N70, Newell 1979; 052, Oakley 1952; 067, Oakley and Campbell 1967; 071, Oakley, Campbell, and Molleson 1971; 075, Oakley, Campbell, and ‘Molleson 1975; P72, Phenice and Saur 1972; P73, Parenti 1973; P74, Protsch 1974; P7S, Protech 1975; R74, Rightmiee 197 ‘954, Singer 1954; 877, Siemon 1977; S80, Smith 1980; 771, “Fobias 1971; 781, Thorne and Wolpott 1981; V49, Vallois 1949; 175, Vallois and Vandermeersch 1975; H39, Weidenreich 1939; W5, Weidenteich 1945; W45B, Weidenreich 1945b, W58, Woo 1958; 71, Wolpoff 1971; 30, Wolpoff 1980; W30B, Wolpoft 19808. Pull references are available upon request, ‘We will be grateful for readers’ attention to errors or omissions. SPECIMEN == cI oC DATE S LOCATION C2 TAX oosr FoRa-732 [STEREFOUTEDX1 [STERKFOUTEZI-S ‘STERKFONTEDN-T 500 2500 F OOMNOSTE TR AR 435 2500 | 0265027 TR AK BS 2500 F O26SOZTE TR AK m 6.5 m 500 2500 | O26S027R TR AA m nm [STENKPOUTEIN-B 530 2500 02650278 TR AK ‘STERKPONTEIN-19 436 2500 O26S027E TR AA ‘STEBKFONTEIN-60 4128 2500 O26S027E TR AK ‘STERKFONTEINT! 428 2500 O26S027R TR AL oMD~L 3381-6 8 2100 M OO5NO36E TR AA ‘SUANTEGANS-46 73,0” 2100 M 0265028E TR AR VOMDRART-B 650 2000 | 02650288 78 AR SANGTRAN<4 62,8. 908 1900 M OOTSIIIE TR HE KoOBT FoRA~1870 "752 1800 OOMNOSTE TR AA-HH NARAPANSOAT-37 495 1800 02850298 TR AL oupuvar-24 590 1800 00350356 TH Ad-t ‘SHARTKRANS-59 500 1800 02650288 TR AR SSMARTKRANS-1585 530 1800 0265028E TR AR OOBY FORA-106 510 1700 M oOnNO3TE TR AR KOOBI FORAW1805 $82 1700 ODINDSTE TR AA-Hit RoOBT FORA-1813 509 1700 OO4MOSTE TR AA-HH oupivar7 687 1700 0035035 TR AA-HH YOOBI FORA-3733 72-3 800 1700 oOwNOSTE TR HE SAYBUNEMACHANG "1038 1500 M OOTSITIE TR HE OLDUVAT-5 67.0 530 1500 M OO3S035E TR AR OLDUVAT-9 674 1067 1300 M 003S035E TR NE oLDUVAI=16 {680 1250 00350355 TR AA-HH (HEsoWRNIRA1 ‘550 1150 _ 003H033E TR AR oLDUvAr-13 650 1000 F 00350358 TR HE-HH SAYGIRARSA2 830 1 OO7StT1E TR HE SSAVGIRAN= 830 F OO7SI11E TR HE ‘un. 800 O26s028E TR ALAR LawTTan-2 175 F O34NTO9E TH HE SANOIRAN-2 110 F OOTSIIE TR HE SANGIRAN-3 THO | oo7SHE TR HE SANGIRAN-17 710 M oO7SI11E TR HE oubuvar~i2 50 F 00380356 TR HE TTRINTL=2, 650 F OoTS1i2e TR HE verreszzéi.bs 500 ONGNOTAE GL HE SALDAMHA 500 M O335018E TR HE-W (GrouxoorreN-3 300 | ONONTISE TH HE. ‘HoUKOUTIEN-10 300 m ONONTISE TH HE (cHOUKOUTIEN=11 4300 F otowti5E TH HE. cuouxour1eN-12 4300 M OR0MII5E TH HE cawDoNG-1 250 F OO7SII26 TR Mat GANDONO=5 250M OOTS112E TR N-HE caNDONG-7 250 F 00781128 TR N-HE cANDONG=10 250 F OOTSI12E TR NAHE oaNoowa=11 250 F OOTSII2E TR NAHE 316 NOTES AND SOURCES eR 732¢172-73)¢L72)(08) (P13)(k80) PL TRANSYRALENSIS"(O67)(P72)(P73)(8T9) (P73)(K80) (73)(R60) CConbosiTe oF 19/58(w72-73)(88) (rr2=13) (078) (72-73) (78) JUVENILE) Co 453. (061) (v0) (879) (os1)(P7) DUETIS, HOLLOWAY REVISION (379)(D65)(075) ER 1870 (H78)(LT3)(B79) ‘ConPasITE OF 37/38 (067)¢H72-73) (era) care) crco73) (HT) (081) (870) (on) (72) 078) ER 406(H72-73) B79) 178) ‘Be 1805 (L7H) (979) (878) em 1813 (174) (979) 878) (0880) P13) (H72=73)(B79) 1 3733(879) DATE UNCERTAIN, (weO(OTS) NEINSANTHROPUS"(O67) (#7273) (B79) OHELLEAN MAN"CBT9) HTB) (13) (72-73) (879) (rays) CINDERELLA, (P73) (H72-75)(879) (80075) HOLLOWAY REVrsroH (¥80)(373)(075) ADULT ESTIMATE OF cc (178)(067)(BT9) (80) 473) C80) (P72) (O75) HOLLOWAY REVISION (065) 075) SUVENILE, "Co FOR ADULT (X80) (065) (075) {809 (373) (181) (065) (075) FRacMenTane, (W60)(H78) (x0) 075) (er20urt) REVISED DATING (879)(061))(P72)(S58) ADOLESCENT co +26: (WO) (D65)(O75) (80) (065) 075) (#80) (065) (075) (80) (055) (075) HOLLOWAY REVISION, SOLO-1 (x80)(272)(879)(0T5) HOLLOWAY REVISION, SOLO-5 (80) (P72)(B79) OTS) HOLLOWAY REVISION, SOLO-6 (x80)(P72)(B79)(OTS) ‘5040-9 (80)(P72)(B78) (OTS) HOLLOWAY REVISION, SOL0-10 (x80)(P72)(879)(OT5)noanpowa-12 12.0 1090 STEIMEIN 72:6 1860, BHRIGSDORF- 74.0 1450, ‘SMINSCOME 78.0 1250 Thal 1120 ont 68.6 ono? 61.8 1435, LAETOLI-18 68.3. 1200 FOUTECHEVADE 78.9 1350 RRapmii-c 83-7 1200, RRapmnnD 85.5 1850, ‘aNovce 78.9 1320 KANJERA-1 6.1 KANER3 67-3 GIBRALTAR 16.8 1200 ‘SACCOPASTORE-1 78.4 1200 Uk FERRRSSII went Le MousTIER 1352 Nowne crRceo 1552 NEANDERTAL 1852 Sr-1 1525, sere 185, PETRALONA 1220, ‘TESHIK-TASH 1565, INOWAVONAS1 150 SHARIDAR- 1600 Le QUINA-HS 1385, ce quimacats D.TRHOUND=1 120 irRounD=2 SvBALTUE TABUKA1 wn BROKEN HILL-1 1280 Lk CHAPELLE 1600 FLORISBAD~1 (QUAFZEN-S 1568, FISH HOBR=1 20 1600 CHATELPERRON 85.5 BYSI-1 m3 DES ENEANTS-¥ 76.3 1715 GGibes BUFANTS=5 68.6 1375 IDES BIFANTSA6 69.3 1580 ‘iL 118 1558 SLs Ths 1520 set.-9 68:1 1590 MeaDee-5 Ba amt Ta rho PREDHOST-3. 71.3 1580, PREDwast-t 70.2 1250 PREDHOST-9 1555, PREDeosT=10 152 COMBE CAPELLE 65.7 1340 cho-tucHos 73.8 1590 Manin cork 71-5 CAP BLANC 16.3 STAROSELYE 73.1 (CHOUKOUTIEN-101 70.2 1500 (GHOUROUTIEN=102 69.3 1380 ‘GHOUKOUTIEN-103 71-3 1300 BARMA GRANDE 11.6 ARMA GRANDE 76.3 BARMA GRANDE 72.2 BRN-1 59:0 1600 Lm PIGUER = T4.7 OLDUVAT-1 66.0 WaDdAK-2 1650 CAPE FLATS 69.0 1230 CHEDDAR To. GABLES cavE=n 70.8 GHBLE'S CAVENS 13.7 LAUGERIE mh CAVGERTE ma Lu xvano-1 75.1 1880 OBERCASSEL = 7.6 1500 OBERCASSEL 10.0 SPRINGBOR-1 73.8 150 CHANCELADE 72.0 1530 KETLOR=1 2.6 1593, Vol. 25+ No. 5 + June 1988 oorsti2e otsHo09E ostnon ie ‘510008 osNMtOTE (051036 05n035E otso3tE (0860008 ON6HOT6E 086016 otgyoz0E 00180355 0150355 035N005H OioNoT3e nS KOO1E ‘onsnoote ontnotae ostNoo7 ‘0501005 50n005¢ owoNoZ3E 038n067E oa7s032e o37HoRnE i6nO0oF 6no00E os2no09 32n009M ouenozie 03300358 o1as028E nsnoo2e oe9s026e 033n035e O3ASO1E oiunoote ooaso35e onanoee uaNDOBE ‘onnncoge 033N035E 0330035 033n035e o4gHO17E 033N036E ‘onpn017E ougnor7e ongnot7e ougnore ousno03e onsno0i 0510039 OksKOOIE OugnO3 KE oon 5 oom 5 oon SE onanoose eno08e banoo8e onguorre ‘swoon 00350358 ooast2e 03550185 ost1003¥ oorso36e 00150368 otsnoorw oasnooiw O2kMt09E ostn007E ostHoo7E (02550298 ‘obsn0O1E 03851855 m ™ ™ ™ ™ ™ ™ m ou ou m a. o a ou au a cL au a a Tt ™ aL a ™ ™ a. ™ a ti m m a t a a a nm m4 7H a mm a a a a a a ou a a mm ™ ™ a ot a or aL ™ m a tt aL ™ c. a a m wene. Hew tt 38 a a8 epesppeseEapessaaee= z rt gE = B Ess EESESSESESEEREEESESES5535935553555--223, 22 HOLLOWAY AEVISrON, SOLO-11 (x80) (P72(B79)(OT5) DISTORTED (P72) B}9) (Ht) (WBOB) DATING REVISED (WE0)(B79)(H51) (O71) [ESTIMATED Cr (W8O)(P72)(B79) (052) (D65) (w0) KIBISH, UNCERTAIN DATE (x60) R74) KIBISH, UNCERTAIN DATE (180)(RTA) (P80) ‘SPECIMEN MUDER UNCERTAIN (360) W60)(B79) (CL QUESTIONED (WB0)(B79)(S80) ‘cL qussrroneD (w80)(H79) (80) 071) RECONSTRUCTED, REDATED (L.70)(W8O)(PTS) SEE. ABOVE (180) (680) (P75) (379) H51 (580) (#80) (79) (H51)(S80) (379)(S80) ‘SEX DOUBTFUL (879)(H51) (79851) (380) (879) C51)(380) (07990851) (079) (951) (0180) (579) (W808) ADOLESCENT, CO +58. (HT) HS) BORDER CAVE, DATE REVISED (067)(PT2)(PTS) (0) (x70) (79) (075) ‘SEX DOUBTFUL, ADULT (879)(H51) cum (879)(51) (061) 80) (P72) (879) (067) (0) (P72) (879) CHILD(T-9 TEARS) (A76) (a0) (879) TH) CHS) RHODESIAN MAN” (067)(PT2)(B78) (879) (851) (580) (os1)(Pr2) (879) 975) JEBEL KAFZEN (MBO) (VTS) (oon) (Pray Fr) @r) ‘SEX DOUBTFUL (067) GRIMALDI (F78) aN) cere) can) (F718) (m0) (279) (P72) (951) (TH) (O75) (ai51) (879) (P72) (TH) (075) (078) P72) 951) C879) (075) [tus (W80) (879) (071) RECORD CRANIAL VOLUME, (w80)(B70) (O75) (379) (0) (279) 000) (0) (mo) (Pra)(a19) (Pr2)(879) (065) ROSTENKT (476)(071) MAGDALENIAN, ABSOLUTE DATE UNCERTAIN (VR5)(OT1) CHILD, C1 ESTIMATED FOR ADULT (A76) uppén’ cave (W38)(W0) (B79) UPPER CAVE. (WBO) (879) (438) (075) UpPEn cave (W80) (579) (W38) (O78) GRIMALDI, DATE UNCERTADH (XD) GRIMALDI, POSTHUMOUS DEFORMATION (K80)(mND) GRIMALDI, MENTONE (80) (HD) (80) (P72)¢579) ‘curLD( 880) {Low CX DUE TO POSTHORTUM DISTORTION (P14)(06T) (480) (065) coon (ere) (78) (aN) cos tur0) THOUTURE, MALE? (067)(170) Le BASSE, SEX UNCERTAIN (F78) (10) BASSE’ (P78) (HD) Liuxrana (075)(P72) (878) xp) (879) (78)(90) (879) MAY BE LATER BURIAL (067)(P72) (rr) (858) (075) a7TaLGAT=1 13.4 137012 4 o77st50E TR MY onwer n2.1" 80.5, 11-H O4gNOOE GL He onWeT 2.1" TTT 11 F O¥9NO10E CL HL ORWET "3.1" 88.9 11 F ORgHOIOE GL He ORWER mali" B62 11 F O8gHOIOE GL HH OnWeT msc14" 77.0 11 F ORQNOIOE GL HL OnWET "BI" 83.3, 11 F OR9NOIOE GL HO conver "tits 78:7 11 on9NOIOE GL He onwer niast" 75:7 11 F O4gHOIOE GL HOE onwer tits 7207 11 F Ot9NOIOE GL HO ower 16.8 11 F onguOIOE GL Ho onuer 78:9 11 F ORgNOIOE GL HB ‘onweT 70:5 1500 11M ONGNOIOE GL Mt OFWET "2k.1" 73.7 182011 M ONONOTOE CL mM ORNET "25:1" 1.2 11. F O¥9NOIDE GL He comune 65.8 1260 10 M O36S1NKE TR He. Kou sunmr-1 67.0, 10 M O36S1mRE TR Me ow SuiMr-5 72.8 10 03651008 TR HH Fou SuiMP-in 58.6 10 M O36S1mRE TR 1 TOE YANG-1 77-4 121010 F O30NIOSE TH HL WADSAR-1 72:5 1550 10 F 00851128 TR Mt Comments by J. Lawnence ANGEL Department of Anthropology, Smithsonian Institution, Washington, D.C, 20560, U.S.A. 9 x11 88 [As a student in the mid-1930s at Harvard I learnt how Berg- man’s law of surface-mass relation applies to human pops lations, with exceptions for recent migrants from another climate and for culturally protected groups in the last few millennia, ‘We were also taught that cranial eapacity was greater in cold climates and cranial and cephalic indies lower and nasal index higher in tropical ones, apparently from natural selection, T ‘taught these things in tuen with pleasure because they explain ‘modern brain-size differences in terms of climatic determination of mass-surface relations rather than intelligence. No one except Nazis or White supremacists could then see biological differences in intelligence between any surviving groups. By now it seems lkely that there has been no meaningful increase in brain size since the Homo erectus phase fully ended (cf Howells 1973) about 100,000 B.¢. Coon reemphasized much ofthis in hooks from 1939 to 1965 (e,g., Coon 1962) thank Beals, Smith, and Dodd for their painstaking gath- ving and analyzing by computer of all the rather patchy data ‘on endocranial volume, body size, body surface area, and head form. It is vastly more useful than in the form of the earlier ‘maps (e.g, those of Biasutti or elines. The authors have had to assume correct and unbiased meastrements and have made proper adjustment for the overestimation of cranial capacity ‘when lead shot is used rather than mustard seed, They also assume genetic determination of the variables with minimal disturbing ontogenetic effects of nutrition, often considerable, oF af artificial head deformation. ‘The reason the authors find the culture—intelligence—brain “size” feedback less effective than climate in explaining brain Increase in human evolution is that increasing intelligence re: lates to silent or redundant cortical surface with its subcortical inks) as well as increasing sensorimotor cerebral and cerebellar cortical surface for better-controlled actual and imagined ac- tion, achieved in the pongid-to-Homo sapiens contrast by more and deeper surface folding without a proportional increase in ‘mass. As Hebb (1949) points out, there is an upper size limit forefficient brain function in terms of cell numbers, interaction, arrangement, and blood supply—size of female pelvis related to newborn head is rvelevant a aimiter. Human beings reached this upper limit of efficiency about 100,000 years ago. Hence to say that “one would expect to find supporting evidence among present-day groups” ereates an absurd straw man: 4,000 gen- exations is time enough for selection to iron out kinks and to 318 apovescent, co +55 (wise) OFMET REDATED FROM MESOLTTHIC-NEWELL=79_(¥79)(HHD) 1D MOMBERS FROM NEURARN CATALOGUE. (79) (1D) (m9) 10) (0179) (a0) (0879) 0) (79) (1n0) (0879) (0x0) (079) (100) (0179) (oD) (779) 100) ‘08799 (x0) (79)(100) (ar79) (1D) (79) (on) (P12) (075) (x81) (075) (181)(075) (ran) (ors) (P12) (058) (075) (812)(879) (W458) (065) 075) equalize intelligence (and approximate neuronal surfaces—not exact mass) in all surviving Rroups ‘Minor criticisms: (1) “Encephalization” means putting some- thing inside the head (brain, ar fui or blood) without implving brain-size increase. (2) The New World is not a foolproof test of climatic selection over 30,000 years, since the latest arvivals, the Inuit, came after the Pleistocene from ancestors adapted to Siberian cold with large Arctic endocranial volume. (3) Neo- teny is not a mutation, but a phenotypic result of the slowing ‘of some, but by no means all, relative growth rates (usually by selection fora numberof rate genes, rarely by environmental malnutrition) () Figures 10 and 11 may confuse some readers, since they reverse the usual left-to-right reading usage in We em culture by ESTE ARMSTRONG Department of Anatom, Louisiana State University Medical Center, 1901 Perdido St., New Orleans, La. 70112, U.S.A 1s x 83 Beals, Smith, and Dodd's bioclimatic model suggests that, among modern human populations, increased cranial capaci reflects increased brain size and that the latter is the result of selection pressures working to inerease brachycephals, itself a thermoregulatory adaptation. An increase in brain size is thus seen as a side effect of thermoregulation. While I concur that the variables of brain size and thermoregulation are associated, do not think that they are causally related in the manner suggested. My hesitation is as follows. The brain is a metabolically very expensive organ (Armstrong 1983, 1984), Al though the human brain represents about 2% ofthe total body ‘mass, the brain continuously uses about 20% ofthe body's total supply of energy (Sokoloff 1981), This large use of energy normally undergoes no significant alterations or cycles such as during normal sleep-wake cycles oF during increased mental Activities (Sokoloff etal, 1955, Sokoloff 1981, Mangold et al 1985). The assignment of a high percentage of energy to the brain distinguishes us fom other known animals (Armstrong 1983, 1984). It is hard to think that such a metabolically expensive organ would enlarge passively from selection for Drachycephaly While the differences in cranial capacities between the wit ter-frost and dy/wet-heat ethnic groups are statistically significant, theyare also small, 89cm. The small differences means that the data on which the interpretations are based must be very clean, particularly with regard to the populations’ nutti- tional and disease states, While overall brain growth is somewhat protected from malnutrition (compared with that of othertissues, such as muscles and skin), the brain also has diminished capacities for recuperation (e.g., Dobbing and Sands 1973) unless a protein-enhonced diet becomes avaitable (Angulo-Col- rmenares, Vaughn, and Hinds 1979). Pre- and postnatal malnutrition produces lowered brain weights in both human and laboratory animals (e.g, Dodge, Prensky, and Feigin 1975), Changes in brain weights have also been noted for populations. Miller and Corsellis (1977) observed an inerease in the mean adult brain weight (52 g for men and 25 g for women) of people dying inthe London Hospital from 1907 to 1977. Teis thought that most of this increase is the result of changed nutrition How much of the difference between ethnic populations may represent differences in nutritional standards? Perinatal dis- cases can also influence brain size directly or indirectly by retarding overall fetal growth (e.g., Myers etal. 1971), a con- dition which produces children with smaller cranial capacities (Leutenegger 1982). Again itis not clear how much catch-up srowth is possible (Roche 1981), Perhaps in the future skeletal ‘markers can be used to estimate disease and nutritional status of the populations used in studies of cranial capacities ‘The high and provocative associations between climate and ‘ranial capacities should also be examined for non-neural as sociations. Cranial capacities include the brain plus the me: hinges and cerebrospinal fluid. Changes in cranial capacities may be associated with increases in the latter two features, particularly the meninges. ‘The meninges are connective tissue land are thus not as metabolically expensive as nervous tissue. ‘While the suggestion that meningeal thickness and volume vary among ethnic groups is speculative, itis testable, With the advent of worldwide use of computerized tomography scanning. it should also be possible to determine whether the ventricles (the brain's internal containers of cerebrospinal uid) vary in size among different ethnic groups. by BENNETT BLUMENBERG Faculty of Sciences, Lesley College, Cambridge, Mass, 02238, U.S.A. 10 x1 83 ‘This is an innovative and broadly conceived study. Aspects of it that rest upon a solid methodological and analytical foun- dation include (a) the cline maps, (b) the revised estimate for present-day worldwide mean endocranial volume, (€) the description of the overall heterographie human (present-day), (athe empirical description ofthe “average” model for different climate zones, (e) the presentation of variate change over time (figs. 10 and 11), in which regeession lines describing illusory central trends are omitted, and () the statistical correlations reported in table $ and figures § and 6, This body of material is provocative and thought-provoking. [A number of questions are raised, however, by the analytical protocols and the conceptual framework within which the statistics and cline maps are interpreted. Why were the spatial autocorrelation algorithms of Sokal (Matula and Sokal 1980; Sokal and Menozzi 1982; Sokal and Oden 19780,5) ignored in favor of an “in-house methodology” whose theoretical foun- dation is obscure and that docs not provide discrimination Statistics? As the authors point out, considerable measurement terror exists in many of thelr parameters. Why were nonpara- metric statistical methods neither considered nor used? “The taxonomic assignments that underlie table 12 and figures 10 and 11 need considerable discussion, Issues that are under serious debate in the geochronological literature include the dating of the Djetis, Trini, Vértessz66s, Saldanha, Chou- koutien, and Ngandong hominid material (Beaumont, de Vil- liers, and Vogel 1978, Jacob 1972, Ninkovich and Burckle 1978, Pope 1982). Furthermore, whether or not any European hominid specimens ean be taken to represent HT. erectus is a problem under intense scrutiny (Cook et al. 1982, Howells Vol. 25+ No. 3 + June 1984 ‘Beals, Smith, and Dodd: CRANIAL CAPACITY AND CLIMATE 1980, Stringer 1981), If H.evectus never occupied Europe, the hhypothesis cannot be investigated for this taxon. ‘Taxonomic schemes must be decided upon and adopted as a methodological device to provide an appropriate context within which to in- ‘vestigate certain evolutionary questions. However, such choices ‘and their associated calibration must be explained and refer- ‘enced. A lack of such discussion assumes consensus where none exists, ‘The data base for figures 10 and 11 is dominated by late Middle Pleistocene and Upper Pleistocene hominids. A regression analysis will therefore be biased towards illustrating late Quaternary H. sapiens population variability and against high- lighting the evolutionary trend(s) that characterize (2) the last 2 million years. Artificially low r values will also result from this approach. It might be better to select a data set in which individual points are as evenly spaced over time as possible in ‘order to maximize perception of the long-term trend in endo- Ctanial-volume evolution rather than uncrtially submit to a discovery bias towards the late Quaternary. Furthermore, on the basis of statistical criteria itis impossible to choose between several bivariate models, both Tinear and nonlinear, that describe long-term trends in hominid endocranial-volume evolution (Blumenberg 1978, n.d.o; Godfrey and Jacobs 1981). Is the very complex question of evolution over time best left for ‘a separate thorough presentation? Likewise, interpretations of the scaling coefficient in allometric relationships might best be considered beyond the scope ‘of the paper. Its not at all clear whether a particular body size and metabolic rate entrain a specific brain size or vice versa (Armstrong 1983, Blumenberg n.d.e) Cognition is suggested asa critical variable for Australopith- cus, H. habilis, and H. erectus (endocranial volume?) but not for the encephalization of later hominids, in which climate assumes the status of priority selection pressure. T disagree in the sense that behavior follows from a particular type of brain and cannot de novo create a new type of nervous system, although it can certainly foster biases in potentials actually realized (Blumenberg 1983). Yet, once a critical threshold (? H. habilis) is crossed, gene-culture coevolution might well be responsible for augmenting early Homo brain size, The Lum den (1983)-Lumsden and Wilson (1981, 1983) model deserves comment here ‘Throughout the text, an increase in cranial size is seriously considered as an important influence upon increasing endocranial volume—surely the cart before the horse! The brain is the active functioning organ that generates (adaptive) behavior; the cranium is but its protective housing. Might not a hypoth: cxis about the coevolution of brain and cranium be more appropriate? Might not cranial morphology be mandated to. large extent by changes in brain anatomy and endocranial volume? Many shapes can contain identical volumes: indeed, cranial morphology reflects important proportional (allometric) relationships among brain parts (Baron 1979, Passingham 1973, Stephan and Andy 1974). Several early crania (ER 1470, ER 1805, ER 1813) are considered globular when compared with contemporaries (Howell 1978), An evolutionary trend towards the domed cranium reflects @ progressive enlargement of the neocortex ‘Modality of evolutionary change is confused with phylogen- «xis. Why isan evolutionary madel gradualistic because chronological sequence correlates with a particular view of systematics? Such a correlation in the evolution of a non- branching lineage does not comment upon rate of change, As Thave said elsewhere, I disagree that endocranial volume and taxonomy bear litle relation to one another (slumenberg 1983), Overall distribution characteristies show statistically signifi cant differences, and all but one taxon are characterized by a distinctive coefficient of variation for endocranial volume (Blu- menberg n.d.6), Within a single species (taxon), variation in 319endoctanial volume may be due to both bioclimatic parameters fnd the range characteristic of stochastic genetic processes. relationship between endacranial volume and reproductive io- lation would be very difficult to demonstrate and likely does not exist. On the other hand, such a hypothesis may be enter- tained for between-species comparisons, wherein the object of study is not simply population-level variation. It is important not to confound the legitimately different levels of the evolutionary hierarchy (Arnold and Fristrup 1982). Furthermore, endocranial volume may be a valuable window that allows critical parameters of brain reorganization to be examined and Interpreted (Blumenberg 1985; Jerison 1973, 1977; Hofman 1983, Passingham 1975), AS the authors observe, the majority of the variance in era: nial morphology is not explained by their model. Statistical noise is certainly present, but I do not believe that all attempts to explain this unexplained portion of the variance are, of necessity, futile. There is an important contribution toe made from the realm of evolutionary’ genetics, ‘The cranium is not a tabula rasa subject only to environ- ‘mental influence. Many components of the cranium have significant heritability coefficients with values that approach 0.5 (Bernhard et al. 1980, Cheverud 1982, Susanne 1977, Torger- sen 1981), The large variance unaccounted for by the model fs likely genetic variance. The University of Michigan group that has been studying Amerindian genetic architecture for over 20 years has established that tribal village gene pools may be ‘considered demes as defined by classe population geneties and are quite distinct from one another. Furthermore, their mode of evolutionary change is dominated by drift stochastic events, 2 punctuational modality, frequent departures from Hardy Weinberg equilibrium, and a fssion-fasion pattern of demic spread (Neel 1978, Smouse, Neel, and Liu 1983). The unexplained variance in this study may well reflect the present-day distinctiveness of gene pools whose evolutionary histories are very different and dominated by such processes. The general ‘model for the hominid eranium is likely adaptive to all ecozones. I suspect thatthe unexplained variance reflects not dif ferences in cranial morphology that are specific adaptations but the range inherent in this suite of stochastic genetic processes Because of their molecular-level genetic mechanisms, such processes do not respond to selection pressures except fortuitously; they are random in design and effect and do not result in ‘obvious directionality and adaptive significance (Barigozzi 1982, Dover and Flavell 1982, Milkman 1982). Nonetheless, in this particular case, T wonder if sexual selection (mate choice) might be an important directional selection pressure with specific cultural boundaries that modifies the stochastieity inherent in the genetic realm T have focused upon, ‘As with all ground-breaking endeavors, this study raises more questions than it answers, and several potentially valu ‘able avenues within which to widen the model and conduct future research are suggested. The authors are to be congratulated for introducing cline maps and historical biogeography into this diseussion and for broadening the conceptual frame- ‘works within which endocranial-volume and eranial evolution may be investigated. They are also to be highly commended {or offering their computer services to other workers invest ating similar problems. by Faxsry G. Girots and SPENCER TURKEL Department of Cell Biology and Anatomy, Cornell University Medical College, 1300 York Ave,, New York, N.¥. 10021, U.S.A. 8 x11 83 ‘The ecological fallacy results from accepting mere astociations as causative relationships. Even if we accept the data as reasonably representative of the groups included and the groups included as representative of the variety of the world’s popt- lations, we are still eft with the question of whether the thermoregulation hypothesis explains a large part of the dis 320 tribution. Although a thermoregulatory mechanism that in- volves the skull does appear to exist, it i not clear that cranial capacity per se is affected by the evolution ofthese mechanisms. Recent studies on other mammals suggest that brain temper” ature is controlled by regulating the venous return from the brain. Some mammals have a “carotid rete" in which the small arterioles course through the venous sinuses of the brain, al- lowing for countercurrent heat exchange between arterial and vvenous blood as well as heat exchange between the blood and cerebrospinal uid, In humans and other mammals the cerebral rete is absent. Nevertheless, in humans the internal carotid artery courses through the cavernous sinus, This sinus is connected to both the internal jugular venous return, via the pe- trosal sinuses, and the external jugular venous return of the face via the ophthalmie vein and its anastomoses. Changes in the ambient temperatures of the face produce changes in the tonus of the smooth muscles inthe venous drainage of the face land, hence, the drainage direction of the venous blood in the cranium. This, in turn, affects heat exchange among the fluid spaces of the cranium (see Winquist and Bevan 1980). There are a number of other areas in which the internal and external jugular drainage systems anastomose, Most notable isthe nasal ‘eavity, where there is a complex system of arterial and venous plexi for emperature and humidity exchange (see Negus 1958), ‘The evolution of this mechanism may indeed effect changes in cranial shape by its effect on the cranial base, Conroy (1980) has discussed the relationship of cerebral venous patterns on the size and shape of the cranial base foramina, This is im: Dortant because there is ample evidence thatthe size and shape fof the cranial base determine the configuration of the cranial vault and of the face (Taylor and DiBennardo 1982, Bjork 1950), Thus, although the authors assume that the measure- ‘ments of the cranial vault have some functional significance, studies on the growth and development of the skull indicate that the size and the shape of the cranial vault may be the result of the way in which various factors are resolved at the cranial base. Since the indirect methods for the estimation of cranial capacity are all based upon measures of the cranial vault itis possible that such methods ae telling us more about the growth dynamics of the skull than about its volume. The authors frequently appear to equate eranial capacity with brain size, which gives the impression that cranial capacity reflects the number of neurons within the skull. Given the thermoreg- Uulatory mechanism cited above, the sizeof the fui spaces in the cranium may be of greater importance, and if there is any increase in cranial capacity due to climatic adaptation, it may be the result of increasing the size of these spaces. In addition, the metabolic role ofthe neuroglia i till unclear, and itis also not clear whether a real increase of brain size occurs primarily by increasing the number of neurons (see Holloway’ 1968). Thus, the authors may have found an actual indicator of brain ther. ‘moregulation, Dut it may be independent of brain size. ‘What do we do withthe additional argument thatthe colder «limes were not inhabited until proper artificial protection was acquired? Hats, or their equivalents, certainly must have replaced a great deal of whatever other thermoregulatory mech= anism previously existed; shouldnt this have buffered variation somewhat? ‘The above comments notwithstanding, we believe that the authors have presented a very important paper. The use of the ‘computer for mapping and analyzing worldwide trends in biocultural relationships will eventually lead to important insights ‘The various difficulties in method will certainly not prove long- lasting. We applaud their efforts by KATHLEEN R. Ginson University of Texas Dental Branch, P.O. Box 20068, Hous- tom, Tex. 77225, U.S.A, 9 X10 83 ‘This paper dresses up old-fashioned physical anthropology with nnew-fashioned computer techniques. One would expect thestrength ofthis approach to be more accurate and rapid analysis fof metrical traits and their diswibution, while weaknesses would Tie in the lack of sound biological theory that has frequently characterized such “trait-plotting” methods of anthropological analysis. For instance, the authors assume that statistical correlation implies causation via natural selection, Causal analysis, how: ever, requires in-depth study not only of potential selective agents, but also of developmental and clinical data, all of which are ignored. These additional data suggest that brachycephalization, rather than being a metabolically adaptive event which permits increased brain size, is a developmental by-product of ‘many interacting variables, some of which may themselves be correlated with climate, Supporting evidence for this interpre tation comes from data indicating that, developmentally, the skull is a highly plastic entity. Cultural practices influence final hhead shape, as do a variety of functional matrices which exert their influence during the maturational period, e.g, the brain, the oral-masticatory apparatus, and the respiratory tract (Moss 1968). The coneept that the brain expands to fill ts container, the skull, is untenable on developmental grounds. Rather, brain growth creates tension on eranial suture lines. This tension initiates bone deposition and grovth of the skull. The causal relationship of brain size and shape with respect to skull form 's obvious from the anomalies of the skull that result from hhydrocephalus and inherited microcephaly as well as from the craniofacial asymmetries which reflect normal brain lateral- iation (LeMay 1977), Nor does the postulate that brain size increased to conserve body energy make sense. The average human brain consumes 20% of the ody’s metabolic energy. Much more metabolically effective ways of conserving heat would be the evolution of insulating layers of hair, fat, or clothing, In fact, the brain uses so much energy that extensive brain enlargement would be incompatible with survival in food-scarce environments unless it provided cognitive skills enabling increased foraging ef cieney and/or increased cultural adaptation to harsh circumstances. The fact that a correlation between cognition and brain size has not been convincingly demonstrated does not mean it hhas been disproven. Most literature on this subject is either anecdotal or based on questionable brain-size and intelligence data. To answer this question in a scientifically valid fashion will require the development of accurate, culturally unbiased methods of determining both intelligence and brain size in healthy young adults. For now, the most logical explanation of brain expansion remains that the brain expanded because ‘neural functions were selectively advantageous, Further, ample evidence exists that factors other than brain erovth also modify skull form. One of these is cradle-boarding, which occurs primarily in cold climates (Whiting 1981). An- other is masticatory function, Tooth size, masticatory muscle strength, and angle of muscle pull have all been found to correlate with head shape in clinical dental practice Sassouni and Forest 1971). Increased trends toward brachycephalization have also been demonstrated to occur in the archaeological record jn conjunction with changes in both tooth size and muscularity and in the absence of pronounced brain-size or climatic changes (Carlson 1976). Moreover, thorough dental-anthropological analyses have explained Eskimo skull form on the basis of ‘masticatory stress (HHylander 1977). Finally, altered respiratory patterns dramatically affect the form of the face and skull. For Instance, children who habitually breathe through their mouths because of adenoid enlargement develop long heads and long, faces. Removal of the adenoids reverses this growth trend (McNamara and Ribbens 1975). It is probable that patterns of both respiration and mastication vary with elimate. The masticatory stresses experienced by the Eskimo, for instance, would impinge upon any preindustrial Arctic population, Conse- quently, prior to concluding that brachycephalization is a met- bolic adaptation, an investigation of climatic variations in the Vol. 25 + No. 3 + June 1988 Beat, Smith, and Dodd’ CRANIAL CAPACITY AND CLIMATE cultural and biological factors impinging upon skull develop: ‘ment should be initiated by Mactry HenNEneRG tl, Polna 3115, 60-535 Posnas, Poland. 2 x11 8 ‘The authors are profoundly right in raising the problem of the determinants of human cranial capacity. The problem has been for so long a matter of prejudice, speculation, and “intuitively satisfying” explanations based upon the simplistic conjunction “larger head—better thought” (though sometimes veiled by highly sophisticated mathematical theorizing) that it deserves ‘calm, reasonable treatment. The traditional approach to the problem of brain size (not exactly identical with cranial capacity because of the many accompanying tissues, vessels, and fluids) was baed on viewing the brain asa “higher” exceptional organ directing the body. However, the brain is at the same time an ordinary organ demanding proper maintenance from the rest of the bod. ‘There is ever more evidence accumulating, with the paper fof Beals and colleagues being an important contribution, against ‘a direct relationship between cranial capacity and intelectual capacity. Fist, within-group correlations between intelligence test scores of individuals and their head sizes are at best weak, fon the order of 0.1-0.2 (eg., Pearson 1906-7, Wrzosek 1931, Schreider 1968, Susanne and Sporeq 1973), and probably due to differences from family to family in the conditions for individual development. Second, head size diminishes with time ‘over the last 20,000~80,000 years—a period of the most rapid cultural/ntellectual progress (e-g., Tobias 1971, Olivier 1973, Henneberg 19842), Hitherto offered explanations of this fact ‘based on a close link between cranial capacity and intellectual ability (¢g., Tobias 1971) or on autodomestication (Thoma 1969) are unconvincing, Third, the tremendous increase in cranial capacity during hominid evolution seems to be fully explained by increase in body size (Guidotti 1980, Henneberg 1984a) when the dimensionality of measures of the two vari ables is equal. After the scales along which body size and cranial capacity are measured are properly adjusted for di- ‘mensionality (e.g, when stature Is taken as a measure of body size, cranial capacity must be expressed as the cube root—'/s power—of its directly measured size, a simple linear relationship is clearly visible in the data. Body-size increase is en- countered in the evolutionary lines of many taxons of mammals, being the expression of a trend towards optimization of energy expenditure and resistance against environmental stresses. Hence hominid cranial capacity evolution seems to be nothing exceptional or unique. Its not the brain structure that evolves ina particular way, but the patter ofits functions. The change ‘may be not anatomical but biochemical and related to a different structure of sensory input under new environmental and social conditions, ‘There are some minor faults in the paper, of which I will comment upon only a few. Cortelation coefficients are indicative only of coincidences, not of actual causal relationships, land must be interpreted with due care. For instance, the cor relation of head shape with climatic zone may result from different susceptibility of brachy- and dolichocephalics to in- fectious diseases and different distribution of pathogenic factors inclimatic zones. Another problems the possible curvilinearity of some relationships. Discrepancy in scaling may be the cause of the higher correlation between cranial capacity and body ‘weight and surface area than between cranial capacity and stature. By the way, the dimensionality of human body weight seems to be less than 3, though certainly more than I and most possibly not exactly 2, since the human body is a geometric form very different from a sphere. To my knowledge nobody ‘has measured its exact value, but some differences between the 321scaling parameters obtained by various authors on various samples of mammals may reflect differences due to body shape in the dimensionality of its mass, treated as a measure of body size, Different tissues contribute in varying proportions to body weight in different species and populations (e-g., fat accu- ‘mulation, while head size is primarily dependent upon the size land robusticity of bones. In this context one possible explanation for the decline in braincase size during the last 20,000 years is its relation to a process of structural reduction of the human skeleton (graclization) occurring as a result of the re- laxation of selection acting upon mechanical robusticity of the ‘body’ coupled with the Probable Mutation Effect and periodic local selection favouring smaller bodies due to scarcity of re- sourees resulting from overpopulation, natural disasters, ete {Tam referring here not to changes in external dimensions due ‘tw simple thinning of the cranium’ walls but to a true change in its internal dimensions (Henneberg 19840,6). Head-shape changes, being somewhat dependent upon cli matic differences, occurred very rapidly, atleast throughout the temperate zone, in the form of a ecent microevolutionary trend. For instance, in Central Europe during the last 1,000 years alone, and without any important climatic change or historically known mass migeation, the cranial index has increased from about 7S in the Early Middle Ages to almost 85 in modern times (about a 10-unit increase in the mean of a distribution with about 3 units sd. over 30-40 generations) ‘This rapid change is certainly not due to climatic change; rather, itis a result of strong selection favouring brachycephalics in response to cultural change transforming the human environ- ‘ment (dvvellings, food production, diseases, social relations, ete. Se0,€.6,, Blelicki and Welon 1968, Henneberg 1976), This ‘example warns against the acceptance of theories establishing simple relationships between huzsan biological properties and ‘general climatic or ecozones. Human culture adapts to ecological conditions; the human body adapts to conditions ereated by both environment and culture. Tis is the case with the brachycephalization just described, gracilization, and possibly the reduction of jaws and resultant structural changes in cranial architecture (ee Krantz 19806 and my comments upon it). Per- hhaps hominids living in various climatic zones had different conditions for cultural evolution and thus different rates and directions of biological evolution? propose using the computer files (1) to determine the correlation of anthropometric variables with cultural conditions or, stil etter, with ecozone + culture, using ether nonlinear procedures oF simply analysis of variance in its basic form instead of the product-moment procedure, and (2)1o introduce, where possible, body-size estimates into HOMDAT and deter- ‘mine the correlation of body size and cranial eapacity bby RoLaND MEnk Département d'Anthropologie, Université de Genéve, 12, rue Gustave-Revilliod, CH-1227 Carouge-Genéve, Switzerland. 1exm 83 ‘The authors deserve to be congratulated for the realization of their impressive data-processing infrastructure and—as a first slobal-scale application of it—a study that has led to a model of brain(case) morphology as related to climate. Their courage in tackling so delicate and wide-ranging a problem must be ‘warmly welcomed: generalizations like this bioclimatic model are urgently needed, but they imply a high rsk of criticism on ‘minor or major details which may be in contradiction with the (over. simplified vision (Paul Valéry: “Tout ce quiest compliqué est inutilsable, et tout ce qui est simple est faux”). Theit model as suchas a parallel to Bergmann’s law—is of real interest; many explanations are innovative and merit wide discussion but also verification, However, it remains difficult to appre- bend the validity ofthe model: there are important factors uch as duration of undisturbed occupation as well as the complete 32 biological history of a population in a given area) which are ‘wally out of control in this approach, Further, the considerable differences from one area to another in the extent of cranial variation (e.g, eranial index and stature in Europe) mean that sampling could have an unexpectedly strong influence on the strength of the cortelations. The argumentation is staightfor- ward and seems quite convincing at first. The morphometrics— and this is a rare example in which their simplicity is not a dlisadvantage—are quite suitable for this approach, which is basically geometric. It must be borne in mind, however, that there is much redundance among them and therefore some of, the figures presented in the results may be misleading (the increase in cranial capacity of 2.5 em per degree of latitude should be corrected for body weight). The authors propose a functional linkage between encephalization and brachycephalization, In the discussion of “cognitive influence” and brain ‘morphology they restrict their considerations to simple brain sige. Tt would have been profitable to inelude brain surface as ‘8 parameter expressing cortical surface, Indeed, if reduction of relative head surface as a consequence of brachycephal- zation ean be considered an adaptive trait with respect to cold the increase of brain size observed in connection with spheri- zation could be regarded as another adaptive mechanism, coun- teracting the reduction of cortical surface that would oceur if ‘the volume remained constant, by IwataRo Monimoro Department of Anatomy, St. Marianna University School of Medicine, 2095 Sugao, Miyamae, Kawasaki, Kanagawa 21. Tapan. 8 Xi 83 In their very interesting and original paper, Beals, Smith, and Dodd state, on the basis of a large number of materials, that the strongest effects on changes and variation in individual cranial capacity occur with solar radiation, winter temper ture, and vapor pressure and that the increase in capacity is 2.5 em per degree of distance from the equator. I the average cranial capacity on a global scale is taken as 1,353 cm! as they suggest, the inerease in endocranial volume in a racial move ‘ment from the equator to 80° N. can be estimated at 200 cm’, 14.866 of the average capacity. This increase would be too large to disregard. Concerning the progressive increase of endocra: nial volume in the human evolutionary process, however, it ‘must be kept in mind that a basic difference between Nean- dertal and H1. sapiens lies in the surface ratio of the different cerebral lobes. Tagree with the authors that cranial breadth is ‘the most important structural determinant of cranial capacity, for the shift of the maximum breadth to an area high above the cranial base apparently strengthened the tendency of the yhuman skull to assume a globular form in the course of the evolutionary process. Here T would like to know whether the slobularty in human skull form due to a northern, cold environment could more or less be explained by Allen's and Berg. ‘mann’ rules. In recent centuries, brachyeranie skulls show a considerable increase in frequency in Eurasian populations, Including the Japanese, that live in warm climates. It is de- bhatable whether climatic factors have become the principal source of cranial variation. by Ropuar R. SOKAL Department of Ecology and Evolution, State University of New York at Stony Brook, Stony Brook, N.¥.11704, U.S.A. 283085 ‘The authors are to be congratulated upon this very compre: hensive analysis of an important anthropometric variable, An approach that would complement and corroborate these find- ings would be through spatial autocorrelation ofthe cranial as well as the climatic variables, If well-developed clines could bbe demonstrated through spatial correlograms for both the an-‘thropometric and the putative climatic variables, then a study’ fof the regression residuals of cranial capacity or cranial module fon climate might be of interest. Continued clinal structure of these regression residuals with climatic factors kept constant ‘would describe the remaining phyletic component of the phe- ‘nomenon. Lack of further spatial structure of regression resid- uuals would indicate a largely environmentally caused {determination of cranial capacity. Another question that should be looked at in conjunction with the hypothesis put forward Dy the authors is whether currently observed differences in cranial capacity could have arisen under reasonable population genetic models given the amounts of time available. An especially crucial test case would be the differentiation among the Amerindian populations. A final caveat: the statistical sig- nifcance of the correlations and regressions observed is probably not at the conventional level as given in table 7 and elsewhere in the paper. There are two complicating factors: spatial autocorrelation among the variables invalidates the oF- inary distribution assumptions of bivariate analysis, and the spatial distribution ofthe points at which samples are obtained biases the computation of the coveelation coefficient. This prob- Jem has been pointed out by several authors e.g. Mather 1976 and King 1979), by Enix Triwkaus Department of Anthropology, University of New Mexico, Al- buguerque, N.M. $7131, U.S.A. 30 x1 83 ‘The authors have argued, on the basis of a selection model for cerebral thermal stability: and correlations across modern hu- ‘man populations, that variations in neurocranial size and shape among H. sapiens can be explained as a product of climatic adaptation. Even though they do not provide a proximate mechanism, other than general adaptive considerations, they are convincing that atleast part of modern human neurocranial variation is de to climate, However, their statement that “within ‘our own species (including Neandertals) climatic factors have become the principal source of the variation” cannot be substantiated During most of our species's evolution there has been a con= tinuation of the encephalization that characterized the genus Homo. Middle Pleistocene specimens usually included within 4H. sapiens and providing reliable cranial-capacity estimates have a mean of 1,231 em’ (N = 4), in between the means of earlier Middle Pleistocene H. erectus (1,101 em, N= 11) and. Upper Pleistocene archaic H. sapiens (1,459 em’, N = 17). It is only with Neandertals and fosils of a similar grade and with ‘anatomically modern humans that encephalization is no longer ‘a consideration; do they exhibit the postulated climatic pat tering with respect to size and shape? ‘When the available cranial-capacity estimates are tabulated with archaic 7. sapiens in the archaic sample and early anatomically modern humans in their sample correcting some of the values given in the appendix, omitting questionable esti= mates, and adding specimens), the supposed climatic patterning largely disappears. There is litte difference among archaic HL. sapiens between “glacial” (1,482.4 = 173.4 em, 1,200- 681 em!, N= 8) and “temperate” (1,438.9 © 176.5 cm’, 1,200-1,740 em, N= 8) samples. The one “tropical” specimen in this group (Ome-Kibish 2: 1,435 cn) falls in the middle of this range, Among early anatomically modern humans, the “temperate” and “tropical” samples are indistinguishable (TM: 1487.2 © 91.1 om’, 1,300-1,587 om!, NV =9; TR: 1,496.0 = 166,1 em’, 1,230-1,650 em, V = 5), even though the “glacial” sample is higher (1/570.4 129.1 cm’, 1,875-1,880 cm’, 13); only the early modern “glacial” sample supports the supposed pattern. Could the lack of patterning be due to body- size differences? This is possible but unlikely, since eranial- capacity’stature indices (emv/cm) for Neandertals (GL: 9.04, 8,0-9.7, N = S;TM: 8.07, 8,0-9.7, V = 3)and early moderns Vol. 25 + No. 5 + June 1988 ‘Beals, Smith, and Dodd’ CRANIAL CAPACITY AND CLIMATE (GL: 8.89, §.5-10.3, NV = 9; TM: 8.50, 82-88, N = 3) show ‘only slightly lower values forthe “temperate” samples than for the “glacial” ones, ‘A similar pattern is evident in cranial indices ifthe samples are rearranged as above. “Glacial” and “temperate” archaic H. sapiens samples are indistinguishable (GL: 74.3 2.7, 67.8— 16.1, NV = 9; TM: 75.3 2.6, 71.6-78.9, V ~ 8), as are the three climatic samples of early anatomically modem humans (GL: 72.2 © 3.1, 66.5-77.8, N= 17; TM: 72.1 = 2.1, 69.3— 15.1, N'= & TR: 71.9 = 25, 69.0-75.0, V = 4). The data are not available to compare “coefficients of cranial morphol- ‘ogy in the same manner as above, but the cranial indices and published assessments of Upper Pleistocene neurocranial mor- Dhology (e.g., Stringer 1978, Trinkaus 1983, Vandermeersch, 1981) suggest that they are unlikely to show Significant differences between climatic samples of Upper Pleistocene humans. of the same grade One could argue that small sample sizes, possible sex biases, and decisions on sample composition have unduly infuenced these results, If this is the case, the available data should not bbe used to test any hypothesis of climatic patterning in neurocranial morphology and size. Yet, if one uses the maximum data available, if the comparisons are between hominids of similar grades, and if specimens are assigned to samples on the Dass of their total morphological patterns, the results should pot vary markedly from those presented here Although it is worthwhile investigating Pleistocene human ‘morphology from a thermoregulatory point of view (e.., Trin- aus 1981), it appears unlikely that neurocranial size and pro: portions were primarily influenced by thermal stress. Itis more probable that the variation in size is due to a combination of tencephalization and the influence of body mass (not merely stature) (the “meat-hed” hypothesis [Holloway 1981). Neuro: cranial shape is controlled by relative rates of cerebral and. ‘eurocranial growth (Trinkaus and LeMay 1982), which are influenced by a variety of environmental and genetic factors, possibly in conjunction with the apparent shortening of human sestation length in the Upper Pleistocene (Trinkaus 1984, 0.4). Regardless of the relative importances of these and other in- fuences on brain shape and size, the observed patterns are likely to be the result of a complex combination of them, not ‘merely one such as thermal stress. Reply by Kenweri L., BEALS, COURTLAND L, SMITH, and STEPHEN M. Dopp. Corualis, Ove., U.S.A, 6.1 84 Many hypotheses are suggested in the comments. For most, however, no data are available, nor are they presented with sufficient information to enable their evaluation, Our responses to the comments are mostly in alphabetical order, although similar comments are combined and the paleontological portion left for the end ‘As Angel remarks, there is little new about the thesis of thermoregulatory effects upon cranial morphology. It derives primarily from original ideas of Thomson and Coon. The pres tent paper does quantify and document the ecological associa tions involved. The data are more comprehensive than those of other surveys, and the variables are for the fist time geo- sraphically mapped. Conclusions concerning the adaptations within the Americas are not materially affected even ifthe Inuit are eliminated from the sample on the basis of recent movement from Siberia—still, fan arctic environment. All the distribution maps ate conven- tionally depictions of group location at the ethnographic present. a3In writing of neotenous mutations we were referring to a ‘mechanism of the phenomenon rather than the phenotypic re sult, Redundant cortical surface may’ have some relationship to variation in human intelligence, but our conclusions are limited to the morphology of the brain's container. ‘We have no contribution to make toward understanding how the brain functions, Its true that 4,000 generations is theo- retically sufficient me for selection to “iron out kinks and equalize intelligence.” The point is that no matter how many generations are involved, no sufficient evidence has ever been presented that variation'in population brain size, head size, hhead shape, or cranial capacity has a connection to intelligence in the first place. We need to rephrase Armstrong’s initial summary of the hypothesis presented. Shor of qualification, itis that increased cranial capacity is a result of increased cranial size (not brain size) in combination with rounder cranial shape—both of which are partially the result of thermoregulation. We concur that the brain is metabolically expensive. In fact, the cranium as a ‘whole is thus expensive, especially with its lack of vasoconstric- tion, It doesnot, however, fllow that metabolic expense would prevent thermoregulatory adaptation from occurring. Cranial variables tend to be more closely associated with climate than the body as a whole despite the metabolic expense. Moreover, the size of the brain generally increases throughout the paleontological record, which clearly indicates that factors over- riding its metabolic cost do exist. ‘key phrase in Armstrong's comment is “enlarge passively.” 1s the cranium increasing in size because the organ it surrounds {s expanding, or is the organ expanding to fil the size and shape of te container? Versions ofthis query may be also gath- ered from other comments (Blumenberg and Gibson). Our dis- tributional data cannot answer it, The nature of the interactive biology between the brain and its housing can nonetheless be separated from evaluation of the end product (observed endocranial volume), and itis observable that that end product hhas thermoregulatory adaptations, ‘We do follow a passive-enlargement interpretation in regard to cognitive significance. We mentioned, for example, the vir ‘ual identity of mean cranial size in Choctaw and Aleut, whose ndocranial volumes are reported to differ by 226 em’. This ‘surplus” results from the differential geometry and apparently produces no behavioral difference. The additional 226 em’ must indeed be metabolically active, but anything cognitively affected thereby remains obscure ‘Armstrong notes that the difference between winter-frost and lected a trait (the cranial index), a portion of the globe (the ‘Mediterranean area), atime for the map to correspond to (20,000 2.9.) and a segment of the evolutionary rate of change for the Intercept = 80.7 b= -3.67 Standard error of & = 1.830 = 0.58 Stgnifteance of x = 0.08 io. 18. A condensed model of hominid eranlal index to minimize Late Quaternary discovery bas. Datapoints fellow chronologieal gaps. A, 9t specimens less than 130,000 yeas +B, 13 specimens feom 175,000 to 500,000 years; C, 1 specimen s $00,000 years, D, specimens fom 150,000 to 600,000 years; remainder of indvidl specimens x-axis, log of age (> 1,000; std line, regression; dated line, empiriea, with brgin at heterographie composite Vol. 25+ No. 5 + June 1984 327SON * 10W 45E S5N Fic, 16. Time machine projection of cranial index around the Mediterranean at 20,000 a.. Horizontal striping, 7278.9 unis, coss-hatching, 7475.9 units dagonal lines, glacaton; question mark, inadequate data for reconstruction: slid ne, approximate coastline trait rom 110,000 to 10,000 8... Specimens were then sorted by time, sex, and site. A multiple regression predicted values at each site as adjusted to the selected time. Distance was then ‘added as an “experimental” stepwise variable and identified as the space between each site and Cabo da Roca in Portugal ‘The result is, to our knowledge, the first attempt at a clinal reconstruction of a human trait in the Pleistocene. Tt should be taken as an illustration of method rather than finality. ‘This method expands the scope of anthropology. In addition to investigating change in traits over time, it is possible to analyze geographical complexes of traits through space over time. Our attempt here to test the method and improve the files has not been notably successful. Perhaps its failure may stimulate colleagues to resolve the problems th system contains. References Cited Awovte-Coustenanes, A. G., D. W. Vavouns, and J. W. Hips. 197, Rehablitation following early malnutrition inthe rat: Body ‘weit, brain sie and cerebral cortex development. Bravn Research Toorzi-ss." TEAL AsutsrioN, ESTE 1085, Relative brn size and metabolism in mam- tale Sclence 220°1302-4.-— (EA, BB) 1064" *Allomett consideration of the adult mammalian bran ‘with specal emphasis om primates” in Adeances in primatolor Size ad scaling in primate biology Belted by Wilkam LJungers New York: Plesum Press, (E8] Anwouny A. J.-and K. Fnisrav?. 1082, The thory of evolution by Tatura selection: A hierarchical expansion. Puleobioloyy 8:15 23. (BB) ‘Bake, P1960, Climate, culture, and evolution. Humon Biology 32:5- Bamicozzt,C. Raitor. 1982. Mechaniime of speciation. (Progress in {Clinical and Biological Research 96.) New York: Alan R Tis. (BB) BARON, G. 1979. Quantitative changes in the fundamental structural batlern of the diencephalon among primates and insectivors. Folia Primatolgica 374-108. (BB) BiaLS, K. 1972, Head form and climatic stress. American Journal of Physical Anthropology 318892 Biats, K. and AJ. Ke1so. 1975. Genetic variation and cultural ‘volition. American Antiyopolgit 17 366-79. Brats, KC. Suits, and. Dood. 1983. Climate and the evolution ‘ot brachyeephallztion American Journal of Physical Antiyopolo acs 328 BEAUMONT, PB, Hoe Vinttens, and J.C. Vacs. 1978, Modern ‘man in sub-Sahiran Africa prior to 40,000 year bp: A review and ‘valuation with particular reference to Bordet Cave. South ajvican Sournal of Science 1409-19. [BB] BEnNHARD, WA. HANCRE, G, BRAUER, and V. P CHOPRA. 1980, ‘Quantitative genetical snaivis of morphological characters of the human head and face. Yournal of Human Evolution O:621= 20. TSB) Biasursi, R 1989, Le reser © § popolé della tera ‘Tipoeratico Bugulesi, Ty and 2, WELON, 1964. Operation ofthe natural selection ‘on the heal form: Homo 1422-40." [MH] [Bjow Anse. 1980. Some biological aspects of prognathsm and the foclusion of the teeth Acta Odontologica Scandinavica 9:1= 40. 'TFGG, ST) [BivMeNnERG, B. 1978. Hominid ECV vers time: Availale data ‘doce not permit a choice of model. Journal af Human Boats eas". (Bi “1983. The evolution of the advanced hominid brn. CURRENT ‘awtiimorovocy 26889-624." [B) ernd.a. Allometry and evolution of Tertiary hominoids Ms. "(BB) 1nd. Population characteristics of extinct hominid! ECV. ws. i) BRENGELMANN, G., and A. BROWN. 1965. "Temperature regulation In Phystoloy end bophyses, 19h edtion. Philndelpbia: Saunders. ‘Baoca, Ps) Sur ln mensuraion dela capacté du crane. Memes “dels Socié @Amhropolagie de Pars 168-182. BuoWN, A. and G. BRENGELMANN. 1966, Energy metabolism” in Phonology and biophyrier, 1th edition. Philadelphn: Saunders ‘Bavrs, A 1017. People and race. New York: Macmilian (Ciais6s, D_S. 1976" Temporal variation im peeisone Nublan eran “American Journal of Phyrical Anthropology 4S467~3. [KRG] CCitvenub, J"M0 1982. Phenotypic, genetes and environmental mor- ‘phologalinteration in the rami Bvoluion 6.499~516. (BB) CCoxtoy, Guise C- 1980, Cerebral venous hemodynamics and bast ‘rani of Cebus. American Joural of Physical andhropoloy S3'37— a2 (ECG, St] Cook, J, CB. STaINOER, A, P. Cunnant, HL P Scuwance, and "A'G) Wine 1983- A review of the chronology of the Exropenn Mice Pleistocene hominid record. Yearbook Physeal Anthro pology 2810-65. TBE) CCoox, C- 1985" Some problems of human varabilty and natural Selection in climate snd eultre. dmerican Naturalist 89:257-79, 1962. ‘The origin of races New York. Koop.» LA) 1965. The ting races of man. New York. Knopt Caoesten, E1981. Climate and anthropometric variation in Europe land the Mediterranean area yal of Human Biology 899-109 Donmine, J and J SaNDs. 1975. Quantitative growth and devel ‘opment of human brain. Avchives of Diveases i Childhood $8.757— or eal DODGE, PLR AL. PRENSKY, and R. D. FEIGIN. 1978. Nutrition ‘andthe developing nervous ryem. St.Louis Mosby. (EA) rin: UnioneDowna, G. A..and RB. FLaveLt. Baits. 1982. Genome colution The System ‘demic Pres Fit, E1081, “Human bioclimatolog” in World exroey ology: General climatology, val 3. baited by H. Lansberg. Am- erdams Elvis, [Fouttes, D. and C. SMITH. 1978, “Using computer graphic in an Ubvopoloes® Proceeding of the Westere Eaducattonal Computing Conference. North Holivwood: Western Ferlodial. Grxoves, 5. 1970. "Anthropometry of late prehistoric human re: ‘alns" in Hayalbook of Middle merican Indians, vol 9. Ealted Dy R Wauchope, pp. 35-49. Austin: Univenity of Texas Pres. Gopraiy, Ly and KH. Jacons. 1981. Gradual, autocaalyie, and ‘punctustotal model of hominid brain evolution: cautionary tale Journal of Human Evolution 10288-72 [BB) GOULD, 8.1977” Boer since Darin. New York. Nowton, ors. Morton's ranking of races by cranial capacity. Science 200-8089 ‘Gorbort, A. 1980. Considerations sur le rapport entre stature et Tapacté di crane par rapport & la poston ebout de homme Blometris Humaine 1673-56. (Mia) Hawt, W. 1947” Cranial capacities A studyin methods, Piediana “Antivopolgy 36:25-7S. Henn, B. 6. 1349. Organi Wiley. OLA) Henweneno, M1976, The influence of natural selection om brachy fophallstion in Poland” Studies iw Physical Anthropology 2:3 i 19880. Ewolucia méegu a intligencja:Preckonania, up ‘renin fakty (The evolution of brain and intlligene’ Convictions, Prejudices, ets) Posnonstie Studia 2 Pilosans Nouns. I rest (Mi) nS 19846, Redukeje trukturane w mikroewoluci Homo sapiens “Aparatrucia, pracylzaca, brachycclallzacja(Steuctual reductions in Homo sapiens microcvolution: Jaws, geaclization, brachy- ‘cephalizaton) Prectlgd Antvopologicany 49. In press.” [MIE] Hignwavse, [1068 La diverite humoine on Afigue subsaharienne ‘Brasels Eaitons densi de Soiologie. Horniax, M-A. 1983: Encephalizston in hominids: Evidence forthe ‘modal of paactustionaiam. Brain, Behaoton and Repltion 22102 i BB) ion of behavior, New York: 1968. “Cranial eapacty andthe evolution ‘ofthe human brain in Culture: Man's adaptive dimension. Edited by MF Ashley’ Montagu. New York: Oxford University Pres) (FG, ST), host. Volumetric and asymmetry determinations on recent hominid endocass: Spy T and Tl, DicbelIrhoud 1, and the Sale Homes erectus specimens, with some notes on Neandertal rain size, ‘American Journal of Physical Anthropology S688-58. TET] Howaut, FC. 1978. *Hominidar” in Rooution of Aican mammals, Talted'by Vi J. Magio and TI. B.S. Cooke, pp. 154-248, Cam- bie: Harvard University Pest. (D1) Howents, W.W. 1075. Evolution of the genus Home. Reading ‘Addison Welles.” LAI 1905. Homo evectus—who, when, and where: A survey Year boot of Physical Anihrpology 25:23. (BB) HapLices, A. 1024742" Catalogue of huseam cromia iy tke United ‘States National Muscwm collections: Washington: Smithsonian instuion 925. Relation ofthe sizeof the head and skull to capacity in he two sexes. American Journal of Physical Anthvopalogy 249" £0. 1942. Ceania of Siberia. American Joural of Physical Anthvo- ology 29:438"81 198, th dion, Practical andropomety: Pladepbia Wistar Hytanea, W.1917. "The adaptive significance of Eskimo caniola- ‘al morpholog” in Orafcial growth ond development. Edited by ‘Av Dahlberg and TM. Graber, pp. 129-09. The Hague Moston TKRG] Jacos, T1972. The absolute date of the Djetis bes at Modjokerto “imtigity 46:148. (BB) Jenson, H.'1970. Brain evolution: A new ight on ol. princples, Selence130:1224-25 075, Evolution of the brain and intelligence. New York ‘Academie Pree 1077. The theory of encephalzation. Annals ofthe New York “Academy of Sciences 299:1%6-90. (BB) JsiGENSEN, JE. PaRtbon, and F. QUAADE. 1961. The correlation tween external cranal volume and bran volume. American Jour: tal of Physical Anthvopology 19:317-20. JORGENSEN, Ty and F. QUaADE. 1956, External cranial volume as an timate of cranial capacty. American Journal of Physical Anthvo- potors 18:661-66 KELS0, A. 1066, The subdivisions of physical anthropology. CURRENT ‘axriinoroLoey 7315-19, Kino, J, 1979, Problems of spatial enalysisin georraphic epidemiology: ‘Sorts, Science, and Medicine 19Dr249-43. TRS) Mass Vol. 25+ No. 5 + June 1988 Beal, Smith, and Dodd: CRANIAL CAPACITY AND CLIMATE Kotxrvetvrssy, T1972, Relatlonships between the frontal sinus and ‘limatie conditions: skeletal approach to eald adaptation. Amer: ‘con Journal of Pharcal Anthropology 8761-12 Kaan, 6.19804, Climate, aces, ond descent groups. North Quincy, ‘Mase! Christopher. ‘99ab" Sapienzaton and speech. CURRENT ANTHROPOLOGY aiisio2. (MH) Lacace, R1979. The HRAF probability sample. Behavior Science ‘Reseach 211-29. LLEMAv, M1977. ASymmetries ofthe skull and handedness: Phre ‘nology revisited. Journal of Neurological Sciences 32 243-53 TieRG) Leurentcore, W. 1982. “Encephalization and obstetrics In primates "with particular reference to human evolution” in Primate brat ‘rolulon: Methods and plenum concepts. Edited by E. Armstrong, Sind D. Fal pp. 8-95. New York: Plenum Press” (EAT LUMSDEN, C.J 1985. Neuronal group selection and the evolution of hominid cranial capacity. Journal of Human Evolution 125162~ ae EBB) LusspeN, C.J. and E. 0. Witson, 1981. Genes, mind, and etre (Cambridge: Harvard University Bese (BB SANTORS. Promethean fire, Cambridge: Harvard University Pres (BB) Maclin, P1982, The co-evaation ofthe rain and fami (MeNastana, JA. and K. A. RIBDENS. 1979. Nasorespivatory func: ion and erinioficil growth. Ann Arbor Center for Human Growth and Development, Universiy of Michigan. [KRG] MANGOLD, Rey L. SOXOLOFF, E. CONNER, J. KLEINERMAN, FG. ‘Tureen, and’S. 5. KETy. 1085 The effets of sleep and lack of sleep on the cerebral metabolism of normal young men. Sowrtl of Clinical Investigation 34-1092-1100. [EAL ‘Mastin, "1081, Relative brain size nd basal metabolic 2457-00, Manin, Ry and K, SALLER. 1959, Lehrbuch der Anthropologie, Stu fart: Flacher. MATER, P'ML 1976. Computational methods of mutivaiate analysis, ‘in pica geography. London: ohm Wiley" [RRS] MAGA, D. Wy nd R. R-SoKAL” 1980. Properties of Gabriel graphs leva to geographic variation research ad the casera of pots Inthe plane’ Coogrphical Anais 12.208-22. (BB) Munmaeak, Ke Editor 1982. Perspective of evoludon. Sunderland, Mass Sinauer Associates. (BB) Minter, A'K Hy and JA. N. Conse. 1077, Evidence for a Secular Increase in husnan bain weight during the past century. Annals of Human Biology $253-87. (EAL ‘Moss, ML, 1968,"The primacy ofthe functional matrices in orofacial frowth. Dentat Practice 19:68-75. (KRG MURDOCK, GP 1981. Alas of world cultures. Pitsburgh: University of Ptsburgh Pres -Monbock, GP, and D. WHITE. 1969, Standard eros-cltural sam: le. Euknlogy 8329-53 Mens, R'E D. E Hut, A.B. Hou, RE, Scors, B.D. Mezzars, and D. B. Cugek. 1931. Fetal growth retardation produced by ‘xpermenial placental isulicteney inthe shesus monkey. Biology fhe Neonate 18379-94. (EA] NEEL, JV. 1978. The population structure of an Amerindian tribe "The Vanomama. Anna Review of Geneier 268415." (BB) Nects, V.E- 1088. The comparative anatomy and physiology of the tote ond poranatal sinuses, London: Heinemann, [FGG, ST) Newauam, ME 1985. The application of ecolopieal rules tothe rail ‘htheopology ofthe aboriginal New Work dmerican Anthropologist 196. Biological adaptation of man to his enviconment: Heat, old, alttude, and nutrition. Annals of the New York Academy of Sciences o1csh7-33, [Ninkovieny D., and LH, BURCKLE, 1978 Absolute age ofthe base ‘ofthe hominid-bearing beds in eastern Java. Notwre 275'306~>. tar OBFTEKING, BRUNO, 1950, Cramiology ofthe north Pacific coat, New "Yorks stechert Ouiviniy 6.19%, “Hominiation and cranial capacity” in fama ‘evolution: Eated by M. Hl Day, pp. 87-101, London: Taylor and Francs IM) Passinenian, Ru E, 1973. Anatomical diferences between the neo crter of itn aid other primates: Brin, Behevion, and Eveation 1557-30.) 1975. Changes Inthe size and organization ofthe brain in man and isancestor. Bratt, Behavior and Eeoltion 172-00, (BB) PEARSON, K- 1906-7" On the relationship of inligence to size and shape of head and other phisieal and mental characters. Biometrika Si08—46.— (MI PicxERING, S. 1950. Correlation of brain and head measurements nd relation of brain sze and shape to shape and sie of the hea. American Journal of Physical Anthropology 181-82. te, Nature 329Putas, D., and S. GovLD. 1974. Size and sealing in human evo- ation. Seine 186:892-901 Porm, G, G. 192. The antiquity of the Asian Hominidae, Physical “Anthropology News 12h -3. (BB) Ronbirts, Df. 1953. Body welah, rac, and climate, American Jour nal of Physeal Anthropology 11585-38. 1076. Climate and human variability. Menlo Park, Calif Camimines Rocite, AE. 1981, “Recent advances in chil growth and develop Tent” Eleventh intomational Compress of Anatomy: Glial and nee ‘onal cll oly, pp. 321-29. {EAI Ssateras,'S. 1930, Relation of cranial module to capacity. American “Journal of Physical Anthropology 1420$-16. SsAssovst, Vand EJ. Fonear 1971, Orthodontics in dental proctice. ‘Se Louie Mosby. [KRG] SCHLAGINHAUPEN, O77O. 1940, Shull from northwestern Siam. ‘American Journal of Physical Anthropology 26°361-81 scitutd, Cand E- MacCascweut. 1985. Basle problem, technique, ‘and theory of opleth mapping. American Statistical Assocation Journat50:220"35. ‘Souaztons, E1968, Beological cules, body-heat regulation, and hur ‘man evelution, Evolution 1:13 RAEN. Quelaues correlations somatiques des tests mentaux. Homo 19.3843. (ME ‘Scrwroereny, 11952, Slektions Theorie und Rassenbildung beim “Menschen. Bxpevimentia 88808, ‘semen G. 1011, Hominidae: Sistema naturale. Turn: Bocea ‘Shtentian, D_ 1979. "A diacusnon ofthe SVMAP program.” in Map. ‘ping sfsoare and cartographic data bases, vol. 2 Cambridge Har ‘ard University Pree C.D. Fostnin, and R. MeNavowro, 1979, Cross-cultural vigone Corvallis: Oregon State University Priting Office swousty PE, JV. Neg, and W. Liv. 1985. Multiple locus depar tures from panmictic eqlibrium within and between wilage gene pools of Amerindian tesa different stages of agglomeration. Ge Renee touiseess. (BB) Somat, RR. ad P. MeNowst. 1982, Spatial autocorrelation of HLA Treaiencles in Europe suppor demic dlfesion of carly farmers American Natural 1107. (BB) Sonat, Re Ry and N-L- ODEN. 1978e. Spatial eutocortelaton in ology. Methodology: Bsological Foural af the Linwacan Soctety 0199-238," (BB) 107Rb, Spatial autocorrelation in biology 2. Some biological implications ant four applications of evolutionary interest. Biolog. {eal Journal of the Linaaean Society 1029-40.” [BB] SoxoLOPr, L. 1981. “Circulation and ener metabolism ofthe brain,” in Baste newrociemistry. Edited by GT. Sigal, RL-W. Alber, BW. Agrnott, and R, ‘Katsman, pp. 471-95" Boston Lite, Brown. (EAI SOKOLOFY, LR MANGOLD, R. L. Wecustin, C. KENNEDY, and ‘S'S Key. 1985. The elect of mental anthmetic on cerebral ei ulation and. meiabaism, Journal of Clinical Investigation 36 ioe. EA} ‘Sreowais, A. 1970, Cold adaptation and the human face. American “Journal of Physical Anthropology 32:243-0. cots, Human adaptation to cold” im Physiological anthro ology: Eaited by Albert Damon, New York: Oxford University bres SHEPHAN, HL, and O. J. ANDY. Quantitative comparisons of brain "ructufes fom instctivores to primates. American Zoolopst 39" 2 TSB) Srewasr, TD. 1988. Cranial capacity studies. American Journal of "Physical Anthropology 28.337-88 srewant, TD, and MT Newax, 1950, “Anthropometry of South ‘American indian skeletal remains” in Handboot of South American Intans, vl: 6. Rated by) Steward, pp. 19-0, Washington, D.C. USS" Government Printing Office. raivorm, C2 B. 1978, “Some problems in Middle and Upper Pll tocene hominid Felationships,"in Recent advances in primatology Enited by D. J. Chivers and KA. Joysey, vol. 3, pp. 398—418, Tondon: Academie Pres. (ET) 330 ——, 1981, The dating of European Middle Peistocene hominids ‘andthe existence of Homo evecus in Europe. Anthropologie 19:3 ey ‘Sweasovn, C1977, esitabilty of anthropologi "Biology 49:573-80."— [BB] Susanne, C., and J. SeORCO. 1973, Etude de cortéatons eristant ‘nis ds tte paycho-tecniqueset des mensuratonscephaliaues Bulletins de tr Societe Roval Belge anthropalgie et Prokstvve ‘Tavuom, Jats V., and RoweKT DIBENNARDO. 1980 Cranial eapac- ‘ieaval base relationships andl prediction of Vault form A ean ‘nicl correlation anaes mericanJoural of Physical Ant hvopolony Ssisi-ss. | (FGG, ST) ‘Twonta, A. i969. Le caractee aromorphotique de evolution humaine ‘ia lume des nouveau Tosi. Symposia Biologica Hungarica bavae. (ME) ‘Twontson, A. 1903. A consideration of some of the more important ‘actors concerned in the production of man's cranial form. Journal ofthe Royal Anthropological Institute 3813866, 1513, The cortlation of etherms with warations in the nasal Index. Intemational Congress of Medicine 89-0, ‘Trupestay, Mand N. Dara MajoMpen. 1944. Cranial capacity iv data on the techiquer of Maedonell snd Brenget journel of Physical Anthropology 2235-81 ‘Tomas, 1911. The Bain and amas evolution: New Vork: Columbia University Press ‘Toon, TW. 1923. Cranial capacity and linear dimensions in White ‘and Neer. American Journal of Physical Anthropology 697-198. ‘Toon, TW, and W. Kexze1. 1925, The estimation of cranial ax ‘pacly A comparison ofthe dieet water and seed methods meri Journal of Physical mropotagy 8281-59 ‘Toman, P1878. Anthropology. Philadephia: Lippincott. ‘Toxcensen, J. 1981. Hereditary actor in the sutaral pattern ofthe Shull deta Radiologie S6.874-82."" (BB) ‘Taaxnaus, E. 1981. "Neanderthal limb proportions and cold adap Tallon” in aspects of human evolution Rated by CB. Stringer, bp 187-224. London: Tuslor and Francis. (ET) Tks, The Shonidar Neawderals. New York: Academic Press "UT 1994 "Western Asi." in The origins of modern mans. Baited by F Spencer and HL Smith New York" Alan Liss. In press (ET) nnd Neandertal puble morphology and gestation length ten E., and M. LeMay. 1982. Occipital bunning among later Pelstoctne hominids, American Jounal of Physical Anthropology staress. (ET) YVagmensteenscn, B. 198, Les hommes fosiles de Qafch (ora, Paris CNRS. (ET) Weck H. 188s. Die capacitit und de drei Heuptdurchmesser der ‘Schaelkapel bel den verschiedenen Nationen, Ava ar Andhra. polos l1-159 Wesring, A. 198i. A note on how many humans have ever lived BioScience 315823-28, \Warrneo, |, W.M. 1981, “Environmental constraints on infant care Practices” in Handbook oferoneruttural muman developmen! Ea Kee by RH. Munroe, RL. Munroe, and B. B. Whiting. New York! Gaviand STPM Pres.” “TKRG} Wovouist, Rawson J. and Joun A. Bevan. 1980, Temperature Shstivty of tone inthe rabbit facial vein: Myogene mechanism or ‘tania thermoregulation. Setenee 29 @07$1001-2.. [FGG, ST] Warn, J.1054, Nowe shape and climate. American Journal of Phyi- teal Anvopoogy 12:— Worrorr, MH “To68, Climatic tence on skeletal nasal aperture lmevican Journal of Physical Anthropotogy 29408-24 Wazocen, A. 1901, © stonunit niektrych pomiaréw antropologiz- veh | (ypaw rasowyeh morflosienyeh do sprawnosct umyslowe) (Gn the relationship between anthropometric varias and mor phologieal types and mental ability). Pretled Antropologicany 51 ton Tana aracters uma

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