ELSEVIER

Human Collective Aggression: A Behavioral Ecology Perspective

Christian G. Mesquida and Neil I. Wiener
York University, North York, Ontario

Moiler (1967/68) proposes that the presence of a large number of adolescents and young adults in a population is a precursor of violent conflicts. But acts of collective aggression are typically perpetrated by males, particularly young males between 15 and 30 years of age. This marked sex difference in the degree of participation is found in all human societies, and it has persisted since the beginning of recorded history. Sexually dimorphic behaviors are invariably found in the context of reproduction, and we discuss male coalitional aggression as a reproductive fitness-enhancing social behavior. This type of social behavior may not increase the welfare of an entire population but it is likely to promote the fitness of the coalition participants. This study argues that the age composition of the male population should be regarded as the critical ecological/demographic factor affecting a population's tendency toward peace or violent conflicts. Our analyses of interstate and intrastate episodes of collective aggression since the 1960s indicate the existence of a consistent correlation between the ratio of males 15 to 29 years of age per 100 males 30 years of age and older, and the level of coalitional aggression as measured by the number of reported conflict related deaths.
KEY WORDS: Collective aggression; Warfare; Mate Choice; Reproductive effort; Age com-

position; Human behavioral ecology.

iots, civil wars, interstate wars and other episodes of collective aggression are unexceptional and pandemic. They have occurred so frequently, in so may different forms of human societies, and over such long periods , o f historical and prehistoric time that many researchers have come to

believe that such events are the product of evolved adaptive behaviors that occur as a consequence of fairly frequent ecological circumstances. It has been argued many times that the roots of warfare are to be found in rapid Received February24, 1995; revisedNovember24, 1995. Address reprintrequests and correspondenceto: Neil 1. Wiener,Departmentof Psychology,York University, North York, Ontario M3J IP3, Canada. Ethology and Sociobiology 17:247-262 (1996) ElsevierScienceInc., 1996 655 Avenueof the Americas,New York, NY 10010

0162-3095/96/$15.00 PII SO162-3095(96)00035-0

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C.G. Mesquida and N. I. Wiener

population growth which, when combined with fixed or slow growing resource availability, increases the disposition to collective aggression (Choucri and North 1975; Vayda 1976; Ehrlich, Ehrlich, and Holden 1977; Conlinvaus 1980; McNeil 1983; Homer-Dixon, Boutwell, and Rathjens 1993). McNeil (1963) for instance, proposes that Latin populations of ancient Italy were much like the Germanic tribes of 100 A.D. " . . . When they, too had but recently settled down to mixed agriculture and solved the problem of rapidly growing numbers by embarking upon a career of conquest that ultimately eventuated in the Roman Empire." These and other scholars vary in their degree of demographic determinism, as well as in their emphases on resource scarcity and distribution as the cause of violent conflicts. But these approaches to the analysis of collective aggression, which might be termed socioecological, agree that group conflicts develop over material goods. Determining precisely what aspect of the relationship between population growth and resources is responsible for group aggression has remained a vexing problem. It is well known, for instance, that there is little correlation between population density and violent conflicts and that resources have to be available to sustain certain military campaigns. A possible solution to the problem would be to consider a human population not as a unified whole but as being composed of different interest groups involved in solving problems at different stages of the life history. The historian Herbert Moller (1967/68), for example, has argued that the presence of a large number of adolescents and young adults is likely to influence political affairs so as to generate violent conflicts. Sustained high rates of population growth can shift the age distribution in such a way that the proportion of young adults in a population increases dramatically, and Moiler suggests that there is a cause and effect relationship between such an increase and political instability. Moiler attempts to explain the observed relationship by suggesting that young adults participate in collective violence, because they have more to gain and less to lose than do older members of the population. His demographic explanation, however, fails to explain why it is young men in particular who engage in collective risk taking, and he does not deal with the potential costs and benefits to individual participants. Clearly, all humans whatever their age and sex have the same basic material needs. It is reasonable to ask, then, why the vast majority of coalitional aggressive acts are committed by young men. Since the invention of gunpowder, and now in the era of high tech warfare, it is not an obvious physical necessity that young males should remain so decisively the main perpetrators of most forms of civil disturbance and warfare. Sexually dimorphic behaviors are invariably found in the context of reproduction, so perhaps intergroup conflict might be better understood by examining competition among groups that differ in their reproductive status and access to reproductively relevant resources. Buss (1989) and others (Essock-Vitale 1984; Hill and Kaplan 1985; Turke and Betzig 1985; Borgerhoff Mulder 1987; Townsend 1989; Prrusse 1994) have argued that women's sexual strategy is to enhance their reproductive success by acquiring mates who have sufficient economic resources for child rearing. This may emerge as the ultimate cause of the propensity of young males to take part in coalitional aggression. The high correlation between warfare and polygyny supports the notion that reproductive competition is a basis for intergroup conflicts (Borgia 1980).

Human Collective Aggression 249

Chagnon (1977, 1988, 1990), for example, has presented evidence that the desire to capture women for wives is often causal in tribal warfare. Aggressive male alliances can be considered adaptive, because they increase men's access to women (Manson and Wrangham 1991) or to material resources needed to have access to women (Low 1993). Aggressors are considered to be collectively seeking inclusive fitness increments (Durham 1976; Boone 1983). We propose that such behavioral ecological argument can satisfactorily explain Moller's observation. The relative number of young males in a given population, we hypothesize, is likely to influence political affairs and to lead to collective violence. It is noteworthy, from a resource competition perspective, that crime-specific homicide offenders are also younger than their social conflict counterparts. Wilson and Daly (1985) have coined the term "young male syndrome" for the fact that homicidal violence and property crimes, in a wide range of societies, are committed largely by young males. They propose that the dangerous, confrontational, competitive interactions of young males, who are typically unmarried and unemployed, are a manifestation of competition for reproductive status (Daly and Wilson 1988, 1990, 1994). Interestingly, homicide rates have been observed to vary according to the relative number of young males in the population; they are elevated when the group aged 15 to 29 composes a large portion of that population (Cohen and Land, 1987). The time/space ubiquity and episodic nature of collective aggression suggest that such behaviors are evolved adaptive phenomena that occur as a consequence of fairly frequent ecological circumstances. Coalitions of resource poor young males, when assessing their chances of success, discount potential risk more than do older males. Laboratory studies of risk taking where both potential cost and benefits are high indicate that males in group tend to make riskier decisions than do individuals or females (Kogan and Wallach 1964; Zaleska 1973/74; Johnson, Stemler, and Hunter 1977). Participation in coalitional aggression can be viewed as an adaptation for the acquisition of reproductive resources, and it is likely to be reflected in the evolved psychology of our species. But knowing what ultimately leads young men to participate does not tell us why warfare is not a continuous process, and why peace endures for long periods of time. Some particular conditions must be present to precipitate episodes of collective aggression. This article further develops and revises Moller's proposition that a particular demographic circumstance--a population profile that is disproportionately young-makes the occurrence of collective aggression extremely probable. We argue, from an evolutionary perspective, that it is simply the relative number of young males in a given population that is likely to be correlated with the presence or absence of collective violence. As a preliminary test of this hypothesis we analyze three different sets of population and conflict data with the intention of bringing to light any relationship that might exist between the two variables.

MALE AGE STRUCTURE

AND COLLECTIVE

AGGRESSION

The term population designates members of the same species that live within a defined area at the same time. But a population can also be regarded as an isolated

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Mesquida and N. I. Wiener

reproductive community. We can therefore realistically conceptualize human populations, organized as independent nation-states, as somewhat isolated reproductive communities. Human populations are in a continual dynamic process of change, and the most important change results from difference between the rates at which people are born and die. Migration is also an important element of population change, which can sometimes alleviate population growth. Fertility, mortality, and migration not only affect the size of a population, they also affect its composition. The most important aspect of population composition is the age structure, also known as the age composition. The age structure reflects the history of changes in births, deaths, and migration over a previous period of time. High rates of population increase can shift the age distribution dramatically, and the resulting large proportion of children has a significant long-term consequence for population growth. Change in the shape of a population pyramid can therefore be relatively slow or nonexistent for decades. Considerable variation in population age structure exists among contemporary independent political states. Figure 1 presents three types of population profiles: type a is an expansive broad-based pyramidal configuration characterized by relatively large numbers of people in the younger age groups and is now typical of high birth rate countries of the developing world. Type b has a smaller percentage of the population in the younger age groups, resulting from a relatively recent drop in prevailing birth rate and occurs in a number of the more economically successful of the developing countries. Type c, characteristic of almost all the developed countries and a consequence of replacement level fertility for two or more generations, occurs when age cohort sizes are nearly equal until the increased death rate of those in their seventh decade and beyond takes effect. There is also considerable variation in political stability among contemporary independent states. It is becoming increasingly apparent that whenever young people come to represent a relatively small portion of the population, as is the case in type c countries, times are relatively tranquil. But when a large portion of a coun-

FIGURE 1. Threetypes of population age structure.Type a is a representative of countries undergoing rapid population growth typified by a number of states in Africa including Rwanda and Algeria. Type b represents a country that has moved from high fertility to essentially replacement fertility for a couple of decades, which seems to be the case for some Pacific rim countries. Finally, type c represents the configuration where replacement level fertility has occurred for 60 or more years and which is today exemplified by most of the European states.
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Human Collective Aggression

251

try's population is young there is likely to be turmoil and political violence. We propose that, because it is primarily males who participate in acts of collective violence, it is the age composition of the male population that represents the important ecological/demographic factor underlying a population's tendency toward peace or violent conflicts. Consequently, whenever it is possible, we will be dealing with male generational cohorts. By considering only the male side of population pyramids, we refine Moller's main thesis, although we suspect that our particular approach does not generate ratios that are significantly different from the ones Moller would have calculated, as in most cases the two sides of a population pyramid are likely to be almost identical. The quantitave analysis that follows will exploit the wide variation in male population age structures to clarify the relationship between the proportion of males who are aged 15 to 29 and the occurrence of collective aggression within and between nation-states.

MALE POPULATION OF CONFLICT

RATIO

AND SEVERITY

The size and age composition of a population are two conspicuous aspects of a species' ecology that are relatively accessible. The quality of demographic data for human populations can however vary considerable from one country to another. Even today, and for a variety of different reasons, census undercounts of 10% are not uncommon (Ericksen and DeFonso 1993). Since the end of World War II, the United Nations (U.N.) has made a considerable effort to improve the accuracy of national population statistics. Periodically, all census data available to the U.N. are evaluated and adjusted if deemed necessary. If retrospective estimates are not available, new estimates are prepared by the U.N. Pol~ulation Division, utilizing available information on fertility, mortality, and international migration. Even U.N. demographic figures can sometimes be unreliable but, in the absence of more accurate figures, and because census data of moderate accuracy are adequate for our analyses, we use as our independent variable mostly ratios derived from demographic data published by that organization. Only in the last of our three analyses, the one dealing with the republics of the former Soviet Union, do we use data produced by the World Bank. We intend to retain Moller's idea of a ratio of males aged 15 to 29 for every 100 males 30 years and over. This ratio provides convenient numerical values that can be used to represent the proportion of young men among the adult males in a given population. The male population is therefore broken into two age cohorts of 15 to 29 and 30+, because age 15 is the approximate age of puberty, when males in significant numbers begin to commit aggressive acts. Age 30 often marks a rapid decline in male participation in individual as well as collective violence. Our dependent variable consists of a measure of the intensity of collective aggression and it will be labeled severity. A good number of possible definitions of the term aggression have been proposed, and this has make the task of quantifying the behavior difficult. However, there is a simple way to overcome this difficulty. Daly

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C.G. Mesquida and N. I. Wiener

and Wilson (1988), for example, have suggest that fatal homicides are a uniquely valuable kind of data for measuring the intensity of interpersonal aggression. Homicide figures, they argue, are readily available and they are less likely to suffer from reporting biases than any other type of measurements. Similarly, we measure the intensity of collective male aggression by considering the number of fatalities sustained by groups directly involved in violent conflicts. In order to avoid having to make value judgements as to which combating army is likely to have been on the defensive, we make not attempt to distinguish between offensive and defensive warfare. Therefore, we treat every country's war deaths as their own "collective aggression" deaths. We assume that official figures for the category "killed in combat" are more reliable than for the category "wounded in combat," and we have obtained data on war-related deaths from such unofficial sources as the World Military and Social Expenditure (Sivard 1988), Measuring Global Values (Sullivan 1991), Conflict Quarterly (Papp 1988), and the International Review of Victimology (Harff and Gurr 1989). Whenever the information had not been previously tabulated, we had to rely on the Keesing's Record of World Events (1960-1970) and Facts on File: World News Digest (1980 to present) for our estimates. Of course, official and press reports of military casualties may not accurately represent the true number of fatalities suffered in each conflict. Certain governments may wish to conceal, for military and political reasons, the actual number of military and civilian casualties. But here again, rough estimates are quite sufficient for the rather crude exploratory analysis we propose to perform. Civilian deaths due to military conflicts can be considerable and even at times exceed those suffered by the military, but reliable data on civilian deaths are quite rare. However, it is young males who most often comprise the bulk of war-related fatalities. In the recent interethnic conflict in Croatia, 91% of those killed were male and of those more than half were between 21 and 35; a significant number were between 16 and 20; and only 22% were between age 36 and 50 (Kuzman et al., 1993). The largest group of female fatalities was among those past reproductive age. It is therefore evident that, even in a conflict where media coverage emphasizes deaths of women and children, it is primarily young males who are the most at risk of being killed by other young males. The shape of a population pyramid tends to change relatively slowly over time, and the ratios of young males to older males conveniently can be calculated from population estimates obtained for a given year. But episodes of collective aggression can be both severe and brief. Consequently, the duration of the time sample for the severity figures must be sufficiently long to account for brief episodes of violence that may take place before or after the year of the population estimate. For most of our analyses, the severity figures reflect a period of at least 10 years (usually 5 years before and 5 years after the estimated population ratio). The severity figures are subsequently divided by 10 to obtain per year estimates. In order to compensate for wide variations in population sizes we also divide the total number of war casualties by the size of the population in millions, to provide severity figures that are per million/per year.

Human Collective Aggression DESCRIPTION OF THE DATA AND THE ANALYSES

253

Our first set of population data is composed of Moller's original sample of 25 countries, which was presented as a table in his 1967/68 article. Moiler made no attempt to quantify the severity of violent conflicts. He simply displayed population ratios in ascending order to illustrate the progressive deterioration in political stability. He offered no method for quantifying political instability, seeming to have assumed that the reader could categorize nations as politically stable or unstable. His ratios were calculated using the 1965 edition of the United Nations Demographic Yearbook, but because Moiler did not differentiate between males and females, his calculations include both sides of the population pyramid. The severity figures, which we collected ourselves from different sources, represent the total death toll per million per year over a period of 10 years extending from 1960 to 1969. Figure 2 is a scatterplot of Moller's ratios displayed on the X-axis, with conflict severity figures plotted logarithmically on the Y-axis. the relationship between the two variables is statistically significant (r = 0.47, p = .017) but the scatterplot shows the presence of three distinct clusters of countries. Most of the countries that show an absence of collective aggression during the 1960s seem to be positioned at the bottom left of the graph, with the exception of the United States (ratio 45) and France (ratio 38), which appear as outliers above that cluster. On the right part of the graph is found a cluster of countries for which episodes of collective aggression during the 1960s were more frequent and lethal.

FIGURE 2. Scatter diagram of M oller's population ratios from the 1960s, in relation to severity of conflict (deaths/per million/per year), plotted logarithmically. Source: for Ratio (U.N. 1965), and for Severity (Sivard 1988; Papp 1988; Harff and Gurr 1989; world Almanac 1995; Facts on file 1960-1969).

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per million/per year) is displayed logarithmically. Source: for population (U.N. 1986), and for Severity -(Sivard 1988; Papp 1988; Sullivan 1991; Facts on File 1980-1990).

Figure 3 is the scatterplot of a new data set composed of all countries of more than 3 million inhabitants listed in the 1986 United Nations Demographic Yearbook. Eighty-eight nations have been included, and ratios have been calculated using only males. The severity figures are given as death toll per million population per year, and they cover the period from 1980 to 1993. A regression analysis using the SAS statistical software indicates a significant influence of the ratio on the severity of collective aggression (F = 79.25, p = .0001, r 2 = 0.47). Again, there appear to be three clusters: one on the bottom left composed of type c developed countries; another on the right-hand side composed exclusively of less developed countries; and a small cluster of four countries that appear as outliers above the developed countries. The four outliers are the former Yugoslavia (ratio 53.7), Argentina (54.1), Ireland (56.6), and Israel (60.7), and we will deal with those special cases later in our discussion. The last sample that we propose to analyze includes the 15 republics of the former Soviet Union. The population figures for this particular data set are based on a 1989 census and were eventually published in World Population Projections 1994-95 (World Bank 1994) a document produced by the World Bank. The severity figures were compiled using different estimates from Facts on File: WorldNews Digest and other press releases. They are presented as the total death toll per million for a period of five years, extending from 1989 to 1993. During and after the dissolution of the Soviet Union some republics began to manifest intrastate and interstate conflicts. A scatter diagram of the population and severity data is presented in Figure 4. It shows that those republics with a male population ratio of >50, with the ex-

Human Collective Aggression

255

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FIGURE 4. Scatterplot of populationratios and deaths per million resultingfrom violent conflictsin the republicsof the former Soviet Union.Severityof conflicts (deaths/per million) is displayed logarithmically. Source: for population(World Bank 1994), and for Severity(Facts on File 1989-1993).

ception of Turkmenistan which appears as an outlier at the bottom right of the graph, suffered a considerable number of collective aggression casualties. The correlation between the two variables is again statistically significant (r = 0.63, p =
.0112). There can never be a definitive evaluation of a hypothesis, and caution is especially appropriate when evaluating a new one. Variables may be selected in more than one way, and what appears to be evidence can always be disputed. The above analyses may not represent a definitive test of our hypothesis. Nevertheless, the results suggest the existence of a consistent positive correlation between the presence in a population of a relatively large number of young males and the incidence and severity of collective aggression. The most important test will be whether this idea can achieve a measure of success in predicting future conflicts.

ALTERNATIVE

EXPLANATIONS

There are several possible alternative explanations to account for the correlations found in our analyses. One of the most pertinent is that countries showing the most

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C.G. Mesquida and N. I. Wiener

intense manifestation of collective violence also happen to be the resource-poor countries of the world, which at the present time have broad-based population pyramids. Our correlations could simply reflect the relationship between the poverty level and severity and not, as we suggest, the relationship between the male population ratio and severity. We are certainly aware of the importance of resource shortage as an aggravating factor in our model. We do claim that, ultimately, young males participate in collective aggression to acquire the resources needed to attract a mate, and we should expect a great majority of the militants to come from that section of the population with the fewest resources. Consequently, the pool of potential participants is likely to increase as a function of the impoverishment of the population, and more specifically as a function of the impoverishment of the population of young males. Resource scarcity may be a necessary condition for the emergence of collective aggression, but it is not a sufficient condition. Impoverished human groups, such as the !kung of the Kalahari desert, seem to live in relative peace for long periods of time, and we feel that such observations can better be explained by the near zero population growth of the !kung (Howell 1979). There is no satisfying definition of impoverishment except as it represents a deficiency in what is desired or in what may constitute adequacy. GNP per capita is an approximate measure of resource availability within a country and a rank order can be taken as a scale of national wealth. In our sample of 88 countries (see Figure 3) we found a moderate correlation (r = 0.69) between GNP per capita and the log of severity. But in order to unequivocally establish the relative importance of this factor in our statistical model, we performed a multiple regression with GNP per capita as the second independent variable. The result of the multiple regression indicates that ratio and GNP per capita, together, explain 54% of the variation in severity (ANOVA, F = 51.41, p = .0001). The presence of the resource scarcity factor does improve our model, given that population ratio, when used as the sole independent variable, produces a squared correlation of 0.47. Of course, many circumstances can result in resource scarcity, but it is, we argue, when this scarcity impacts on young males, usually as a consequence of population growth, that collective aggression is likely to occur. Therefore, it is fair to conclude that in spite of the importance of the resource scarcity factor our model retains its predictive value even in its absence. What remains unclear is whether low per capita GNP is a consequence of population growth outstripping economic growth. We find that the correlation between GNP per capita and our population ratio is r = 0.76, which is not negligible. Another alternative explanation, which is somewhat related to the previous one, is that our results may simply reflect the relationship between the development status of a country and the presence of lethal conflicts. The notion that violent conflicts are an integral part of the development process has often been suggested (Schrijvers 1993), but the idea has never been systematically investigated. It is true that, at the present time, population age structure is correlated with development status at the national level, but this phenomenon is relatively recent. So-called developed countries have not always had population pyramids with narrow bases, and developing countries have only recently experienced a population explosion which is due, at least in part, to the introduction of western-style medicine and to the subsequent drop in infant mortality.

Human Collective Aggression

257

It is pertinent to emphasize that even well-developed countries have, at times, initiated armed conflicts. In the 19th century, Germany replaced Britain as the most rapidly industrializing state, and it has remained an example of what is usually considered to represent industrial and economic development. This did not prevent Germany from initiating two major conflicts in 1914 and 1939. It is interesting to note that during the early years of this century, Germany's population was affected by high rates of natural increase, and in 1910 the ratio of young males 15-29 for every 100 males 30+ was 75 (Mitchell 1992). This was a much different ratio than that prevailing in the other major European states. World War I, although it reduced the number of young German males, arguably delayed entry of survivors into the competitive mating system, The war's economic aftermath, compounded by the global economic slump of 1931-33, aggravated and extended the duration of resource limitation. By 1933, the year the National Socialist Party was democratically elected, the ratio of young males to older males was down to 55. But when we take into consideration the large number of males 30 to 34 whose intry into the employment/marriage stream had been delayed by the war and the depression the figure for the ratio of dispossessed young men to older men is closer to 70. We reiterate that male collective aggression existed well before what we understand as industrial and economic development was initiated, and it has continued to manifest itself even in fully developed societies, as is presently the case in Ireland and the former Yugoslavia. Finally, one might conceivably argue that it is the difference in political organization, and therefore the difference in resource distribution within countries, that is likely to account for the correlations that were found between the age structure and collective aggression. This argument is not supported by the analysis done on the 15 republics of the former Soviet Union. Most of the 15 republics have recently made a radical transition from one form of politico-economic system (communism) to another (free-market capitalism). We would suspect that during such a period of rapid resource redistribution friction would develop between individuals, as well as between competing interest groups. But because of the similarity in the political transition, we would expect the severity of conflicts (death toll/million) to be about the same in each one of the 15 republics. This was not the case and it is illustrated by our scatterplot (Figure 4). The wide difference in ratios found in Figure 4 reflects the fact that some of the republics have had high annual population growth rates for some time, and they now display broad-based population pyramids. Turkmenistan, which appears as an outlier in the scatterplot, is no exception. What is interesting to note about Turkmenistan however is that it has remained firmly under control of a hard-line communist government. Under such a governmental system, political opposition is virtually unheard of. This observation may attest to a better distribution of available resources, but it may also indicate that in such a system political opposition is quickly extinguished. At this point, we may want to consider the possibility that many forms of state repression are in fact manifestations of government-sanctioned collective aggression. We may also wonder whether the choice of political system made by the members of a population is somewhat restricted by the age composition of its male population. It is possible that the shape of the population pyramid ultimately deter-

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mines what type of political organization is practicable. As for democracy, it may very well be a luxury that only countries with narrow-based population pyramids can afford.

DISCUSSION Currently we have a tendency, at least in the Western world, to assume that young men are essentially unwilling participants in armed conflicts. We tend to see such participation as a consequence of manipulation and coercion, and we often make the assumption that a special interest group or class is promoting or financing, in its own interest, the young male coalition. It is likely that our thinking has been conditioned by our recent history of warfare waged at the outer limits of the West's sphere of influence (Korea, Algeria, Vietnam, etc.). We have not had the time to forget that these conflicts were accompanied by resistance on the part of the conscripts. But it is probably because we have no recent experience with territorial expansion that we fail to appreciate the fact that whenever a population has an over-abundance of well armed and organized young men, pure exploitation and coercion are extremely difficult to implement. The confusion is caused by the observation that, at least from a distance, decisions to participate in expansionist warfare also appear to be made by members of elite groups who keep well back from the front line. When we realize, however, that in most of the wars fought over the last 160 years the aggressor prevailed (Bueno De Mesquita 1981), it is not difficult to imagine that in many cases the young men reaped some benefit. Even survivors of unsuccessful coalitions may benefit from their participation. A random thinning of the ranks may enhance the prospects of the survivors by reducing the local competition for reproductive resources. It is likely then that controlling elites astutely underwrite such risky undertakings as territorial expansion or colonization, especially when the alternative is having the aggressive tendencies of the male citizens directed at themselves (Boone 1983, 1988). The mechanisms that would explain how our independent variable causes the emergence of collective aggression are not entirely clear. But we suspect that the social dynamics that engender the emergence of violent uprisings or expansionist warfare might be caused by the presence of resource competition between older and rising generations. Older generations frequently control the political and economic resources of the community, and this may restrict the availability of resources for upcoming generations. The existence of recurring conflicts between generations can be traced back to antiquity, and a body of theories concerning generational patterns of youth movements has been developed (for a comprehensive treatment of the subject see Braungart, 1984). In their struggle for an adequate share of the available resources the success of the rising generation's members may be crucially affected by how numerous they happen to be. Tentatively, we would like to propose that this intergenerational competition for reproductive resources, when exacerbated by the presence of a relatively large number of resourceless young males, might result in the emergence of male collective aggression, which occasionally expresses itself as expansionist warfare.

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The notion that there exist resource transfer problems from one generation to another is elegantly developed in a sociobiological study of the 200-year period of Portuguese expansion (Boone 1983). Boone suggests that the presence of a large number of second and third sons among the nobility, who had little or no access to inherited land, corresponds to the period of Portuguese history when that country was involved in expansionist warfare and colonization. The dispersion effect, which resulted from the land shortage created by the surplus of young males, took the form of direct aggression against both neighboring and distant groups, and it was characterized by a higher war-related mortality rate among landless sons. Boone argues, convincingly, that the behavior of the aggressive coalitions of young Portuguese males is best understood as the aggregate consequence of individual adaptive strategies. Under certain ecological circumstances, participating in coalitional aggression has a better cost/benefit ratio in terms of inclusive fitness than alternatives, and we have suggested that the presence in the population of a relatively large number of young males represents such a circumstance. This idea may be further tested by examining the more distant past for evidence concerning correlations between population profiles and armed conflicts. Historical evidence does present problems regarding the existence and reliability of suitable demographic and casualty data, but those problems are not at all insurmountable. A good example of the kind of demographic information that can be obtained from historical sources is Boone's utilization of the manuscript Peditura Lusitana, which contains the genealogies of several hundred medieval and early modern Portuguese noble families. It is likely that the type of population profile found to have existed in Portugal during its overseas expansion was also characteristic of many European countries during similar episodes of colonial expansion. It is well known, for instance, that the French revolution and the subsequent Napoleonic wars came after France experienced a sustained period of population growth. Valuable information can certainly be obtained from this wide open research area, and the acquired knowledge could be used to either substantiate or disprove our claim. We realize that within a single polity, endogamous groups distinguished by race, ethnicity, religion, class, caste, etc., may not always coexist peacefully. Violent conflicts sometimes develop among them, particularly, we would like to suggest, when one of the groups, majority or minority, presents an age composition skewed to the young. The endogamous group with the fastest growing population is most likely to become the source of collective aggression manifest in several forms of communal strife including civil insurrection, rioting, civil war, ethnic cleansing, genocide, etc. Contemporary events such as the recent civil wars in Lebanon, Sri Lanka, and the former Yugoslavia and the secession conflict in Chechnya may be examples of this phenomenon. Nation-states experiencing a population crisis with one of their endogamous groups present a particular predicament for our statistical analyses, especially when the group in question happens to be a minority. The use of aggregate population data, such as provided by the United Nations, may sometimes produce results that are difficult to interpret and which could not have been anticipated. We suspect that at least some of the outliers found in our analysis of 88 countries have a population problem that is not detectable when using aggregate data. Of particular interest is

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the present demographic situation in the former Yugoslavia, because at least two sources of data seem to indicate the presence of uneven population growth rates (Decroly and Vanlaer 1991, p. 121; Council of Europe 1994). The recent conflict in Chechnya could have a similar origin, and we may be able to learn much about this violent incident by paying close attention to the demographic conditions that exist in that region (Decroly and Vanlaer 1991, p. 121 ). On the basis of a small number of ecological variables, it may become possible to predict certain aspects of our social behavior. The human behavioral ecology argument developed in the previous pages postulates that under certain specific ecological conditions, episodes of collective aggression become more likely. Populations with ratios of young males exceeding 60 per I00 males 30+ are predicted to move toward a state of internal or external conflict, unless there exist particular mitigating circumstances such as an extremely rapid increase in resource availability or new possibilities to migrate to more productive environments. The causal relationship between our population factor and the emergence of violent conflicts may be complicated and may involve many intervening variables, but the presence of a consistent correlation can already allow predictions to be made as to where fatal intrastate and interstate conflicts are likely to occur. CONCLUSION Under what particular ecological conditions are episodes of collective aggression more likely to occur, and what are the potential benefits to members of the coalition? Some insights into the adaptive nature of the behavior can be gained by approaching the subject from a behavioral ecology perspective. Survival and reproductive strategies of individuals, not systemic structures, are the basis of behavioral ecology. We and others (Boone 1988; Chagnon 1990; Low 1993) have argued that collective aggression may be viewed within the behavioral ecology context as an attempt by young males to acquire otherwise unobtainable resources and thereby gain inclusive fitness. We have examined the influence that different population age structures have on the development of violent conflicts and concluded that the presence of a relatively large number of young men makes coalitional aggression more probable, particularly when resources needed to attract a mate are insufficiently available or poorly distributed. Such explanation has the advantage of being susceptible to generalization, and it may account for the existence of tribal as well as modern warfare. Indeed, it has not escaped our attention that if the hypothesis proposed in the present article was to be rigorously tested and proved to have genuine predictive power, it would bring us one step closer to the development of a much needed general theory of warfare and other episodic manifestations of collective aggression.

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