Ecology and Population in the Eastern Mediterranean

Author(s): J. Lawrence Angel

Source: World Archaeology , Jun., 1972, Vol. 4, No. 1, Population (Jun., 1972), pp. 88-
105
Published by: Taylor & Francis, Ltd.

Stable URL: https://www.jstor.org/stable/124144

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Ecology and population in the Eastern
Mediterranean

J. Lawrence Angel

Ecology

The Eastern Mediterranean stretches from the mountain zone of the Balkans, Greece,
Crete, and Anatolia, with the sunken Aegean stretched between, around Cyprus and south
via the hills and plains of the Levant to the Nile delta and North Africa. Easily traversed
steppe grasslands lie to the north and the semi-desert zone of plains and plateaus to the
south and south-west. Ancient crystalline geologic blocks support these eroded areas
around the Mediterranean, and the very recent (Miocene to Pleistocene) Alpine-Balkan-
Taurus-Zagros-Pamir limestone mountain-building, with its accompanying vulcanism
and subsidences, and contrasts of altitudes, water-flow, soils, and coastlines, is what
marks the area as less simply xerophytic than the Near East.
There are three ecologic systems: (a) that of still-forested mountain and bare rock
plus either karst plateaus, or, in Anatolia, high steppe country; (b) the coastal and island
system of scrub-covered rocky and deforested hills, either with rich but stony volcanic
soils from igneous (Gneiss, granite) or metamorphic (schist) rocks, or less fertile sands,
marls, conglomerates or Flysch formations covered with grey, stony forest-built soils,
or more acid Pleistocene red earths washed down from mountains; (c) both large and
small river valley systems again filled with useful silt from Pleistocene soils as sea level
rose between 9000 and 4000 B.c. (Raikes i969; Butzer i964, i965). Results are a steep
diversity of ecologic niches quite closely spaced, a rather small arable area (20-30%)
often both stony and requiring some irrigation and terracing to farm, and a divisive
land topography, so that the sea is the medium of unity (Maull 1922; Angel i946, 1971).
There is another reason for irrigation, terracing, and cisterns (or today, artesian wells
and occasionally river dams). This is the scantiness, year to year variability and sharp
seasonal limits of rain. Rainfall is the key challenge. Water for farming normally here
sets the limit on population density, largely because the typical rainless Mediterranean
summer (May through October) is a time when bulbs, evergreens, and other plants
aestivate. 75% to 95% of the rain and snow falls in winter (October to March). This
winter is like a New England autumn with occasional colder periods (9-12?C), because
of the moderating effect of the sea, while the Balkans and Anatolian plateau are much
more continental (o?C) and snowy (Butzer I965; fig. 3). In the last months of winter
flowers bloom across the hillsides and the spring growing season (March through May)
sees a temperature rise to zo?C and then 25 ?C. Because the prevailing winter winds are
from the west, rainfall varies sharply by region, from 80 to I30 cm. along the steep

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 Ecology and population in the Eastern Mediterranean 89

western coastal mountains down to 35 to 50 cm. in the Aegean basin (Maull, I922) and
inland Anatolia, but below 30 cm. only in the semi-desert areas to the south. The best
Mediterranean climatic zones, adapted to wild and then domesticated olive, grapevine,
and citrus trees, include a oo00-200 km. deep strip of coast and mountains (Butzer I965:
fig. 3), plus an eastward extension to the Zagros slopes. Helbaek (1960) and Harlan and
Zohary (1966) show wild wheats with a highland distribution along the southern edge
of the mountain belt, with Triticum aegilopoides (ancestral einkorn) distributed as far
north and west as Thrace; wild barley also has a highland steppe distribution as far east
as Turkestan. Western plant domestication began on the eastern edge of the Eastern
Mediterranean (Butzer 1965) with wheat and barley, among latest Paleolithic hunters
some of whom were starting to winter-feed and domesticate sheep and goats (Reed I960)
at the end of the Wiirm glacial retreat and contemporarily with the shift to Mesolithic
hunting and fishing further west. Such trees as pistachio, almond, flame-oak, and apple
were used for fruit early, but the olive (requiring grafting) and vine were domesticated
only by the Early Bronze Age (Brothwell and Brothwell 1969; Hopf 1962).
During the last major phase of the Wiirm glaciation (26,ooo-8000 B.C. with a maximum
advance c. I8,000), sea level was oo00 m. lower and mountain glaciation in the Alps,
Caucasus, Armenia, Anatolia went with snowlines at I,800-2,500 m. - 600-900 m. lower
than today - thus implying temperature 2-3?C or 7-8?F lower (Butzer 1964, 1965)
and a harsh temperate but not wet woodland ecology for the Mediterranean. The warmer,
temperate, phase in the eleventh and tenth millennia B.C. (before the brief Dryas-period
last glacial advance) was wetter and favored woodlands, and during the warmer Boreal
Phase and warmer and slightly wetter Atlantic (5500-3000 B.C.) climatic phase there was
no climatic check on woodlands, and the rising sea level reached (Raikes 1969) and by the
fourth millennium surpassed by 2 m. (Butzer I964: 448) its modern level. The succeeding
drier and more continental Sub-Boreal phase down to 6o00 or 400 B.C. began and ended
with slight glacial re-advances (Denton and Porter 1970) and sea levels dropped 3 or 4 m.
so that some Mycenaean and Classical sites are I-2 m. below modern sea level. The wetter
Sub-Atlantic phase had warm and drier interludes (Denton and Porter 1970) about
A.D. I00-400, A.D. 800-I300, and 1900-I950S; and cooler phases about A.D. 700 and
especially from A.D. 1350-I850, with slight sea level changes. One major point is that
since the post-Pleistocene warm up of Boreal times (eighth and seventh millennia B.C.),
climatic fluctuations have been very slight indeed in effect, except where sea level changes
have changed the area of coastal or riverine marshes. Since earthquakes and vulcanism
are frequent, tectonic changes of land level are important as well as the eustatic sea level
changes mentioned. The second major point is the corollary that the drastic ecological
changes in the Mediterranean since Early Farming times have been man-made, the
results of cultural use and abuse of the land.
The Upper Paleolithic period of intense cold, with hunters occupying caves (Butzer
I965; Angel I971) in the Levant, southern Anatolia (cf. Karain), and Greece (cf. Petra-
lona, Peneios valley, Copaic basin, Kaki Skala, Franchthi Cave etc.), but probably
avoiding the high plateaus, saw a cool temperate woodland with deer (fallow, red and roe),
wild cattle and pig, Equus hydruntinus, brown bear, wolf, fox, hyena, leopard, and various
smaller mammals (Butzer I965; Jacobsen I969). The warm up before and after I0,000 B.C.
(Van Zeist and Wright I962) increased the spread of woodland and, after the

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90 J. Lawrence Angel

interruption of the last Pleistocene cold fluctuation in the ninth millennium, led to the
basic post-Pleistocene subtropical drought-resistant evergreen and coniferous woodland,
with warm-temperate forest of black pine, juniper, evergreen oak and some other hard
woods in the extensive mountain areas, and the Pleistocene mountain coniferous forest
now only up in the Caucasus, Cappodocia and north Balkans. In the basic dry Medi-
terranean open woodland at altitudes less than 900 m. live or evergreen oaks were the
chief element, with umbrella and maritime pines, wild olives, in a few rockier areas
evergreen scrub or maquis including heather, myrtle, pistachio, rhododendron, bulbs
(asphodel), anemones, poppies, wild grasses browned in the dry season, and occasional
meadows and marshes (Butzer I964: 66-442; 1965: 9; Maull 1922: 32-42). In this
environment, Mesolithic hunters, probably adapting to the post-glacial spread of
falciparum and other malarias (Angel 1966; I97I: 77-9), lived on red deer, pig, cattle,
and other forest fauna and then by the eighth millennium started sea fishing and also
trading by boat like their western contemporaries.
Latest Paleolithic and Mesolithic development of composite tools and microliths
stimulated trade in obsidian (from Armenia, from Hasan mountains east of (atal Hiyiik,
from Melos in the Aegean) and Jacobs' (1969) stress on trade as a major motive for
establishing fortifiable towns with a secure food supply suggests that the reason for the
rapid post-glacial spread of newly invented agriculture was economic. Three results of
the establishment of farming towns and villages around the Mediterranean in the eighth
and seventh millennia B.C. (from Jericho and Tell Mureybat west to ?atal Hiiyiik,
Ha9ilar, Argissa, Nea Nikomedeia, Lerna, Franchthi, and south to Egypt) were: im-
mensely rapid increase in population (cf. Deevey i960; Barringer I966); consequent
overhunting, start of deforestation and overgrazing; consequent rise and spread of
diseases such as malaria, and dysenteries and hookworm.
Population pressures during the next six millennia not only caused complete lowland
deforestation and replacement by maquis and finally by thorny scrub (Butzer i965) but
also destroyed parts of the already limited fraction of land (about one-fifth to one-third)
which could be cultivated. Fallowing began quite early and after the introduction of stone
and then bronze axes for tree-clearing, and use of the ox-drawn ard or scratch-plow
by the third millennium B.C., scientific farming with adequate manure, ash from burning
over to correct the excess of lime in the soil, irrigation, and control of water, developed
by the Late Bronze Age (Angel 197I). Yet soil minerals were lost, especially potassium;
on the other hand, salinity was spoiling some inland basins; and erosion after loss of
trees was beginning. Plato describes Attica as being skeletally bare, as it was after World
War II, although in the Mycenaean siege-fountain deep down in the north side of the
Acropolis rock at Athens Broneer found beams of full ship-mast size set there 850o years
before Plato. In the intermediate time, the local forest timber had gone into vital ships
for the navy, house-beams, wagons and baulks for the silver and lead mines, and charcoal
for metallurgy (rather than for heating houses or for cremations). Cattle and other
domestic animals also got smaller during Bronze Age to Ancient times (Gejvall 1970).
During Roman times, with the population massing more in cities and still not below
modern density (over 30 per km.2), in part because of grain imports from the Ukraine,
Crimea, and North Africa, land conditions became still worse as slave-farms or lati-
fundia spread, and greater use of wood and continued overgrazing by goats and sheep

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 Ecology and population in the Eastern Mediterranean 91

(now tending to replace cattle) led to still greater erosion represented for example by
the massive silting at Olympia and in the Anatolian and other Mediterranean valleys
(Vita-Finzi I969). The slightly higher sea levels made marsh-control difficult, and in-
creasing malarias as well as more massive epidemics, ending with true Plague (A.D.
531-590) under Justinian, led to loss of population (Ackerknecht I965). Slaves were
probably the hardest hit and there was desertion of villages and areas of countryside.
Weakened by continuous wars since Classical times, and by the collapse of Hellenistic
science after the burning of the library and museum at Alexandria, the culture lacked
vitality to rebuild temples, cities, or sometimes even aqueducts destroyed by earthquakes
(e.g. Olympia, Antioch). The ecological background for the rise and spread of Chris-
tianity and then of Islam was as bleak as the political and military one.
It is amazing that in the chaos of invasions and wars (Goths, Huns, Avars, Slavs,
Albanians, Moslem Arabs, and then Seljuk Turks and Crusaders) the new Christian
Byzantine culture was able to build domed churches over the same range of towns as
the Hellenistic theaters and Roman baths, including the original and miraculously
domed basilica of Santa Sophia in Constantinople. But there was no recovery in land
use, though silting valleys were slowly restoring soil. The rich Byzantine familes tied
down the farmers with taxation and laws, and in a way it was the survival of Bronze
Age village farming which saved them and allowed the ninth- to eleventh-century
Byzantine cultural revival during the drier interval. Cities and monasteries became small
(except Constantinople) and defensible.
In the fourteenth century, during and after the great Plague pandemic, the Osmanli
Turks conquered Anatolia and the Balkans and, after a delay caused by Mongol conquests
under Timur (Ankara fell in I402), took Constantinople in I453, then Greece and then
the Crusader-Arab-held Levant and North Africa (by I517). This sixteenth-century
Golden Age of the Ottoman Empire under Suleiman temporarily helped farming,
without the Byzantine tax-load. Rice, introduced in Roman times, now supplemented
bread and eggs; fish and rare goat or mutton were available, while the new sugar-cane
from SE Asia added food energy. But on the whole the Ottomans did not innovate in
land use, or quickly take over the New World crops (maize, potatoes) except tobacco;
and a new slavery spread. Farming remained on a Bronze Age level through the Moslem
period and Romantic revival, right down to modern times. The climate was cooler and
not dry. But malaria increased and was thoroughly endemic up to the time of DDT;
and plague was endemic from A.D. 53I until I845 (Ackerknecht I965).
With this ecological background, how did population respond to specific cultural
changes and pressures, and what was the biological background for such responses?
Since ecological deterioration and our archaically militaristic and impoverished,
though creative, culture now set problems for us quickly to solve, it should be useful
at least to see how human biological data from the past can serve as an experiment. What
I shall try to do is to derive population densities for the Eastern Mediterranean for
successive periods from Upper Paleolithic to modern and to relate these, as shown in
table 28 and fig. I6, to the number of infant and child deaths in ratio with those of
adults, to the average length of life of adult females, the number of births per female, or
average parity or fecundity, and the consequent number of survivors per female, or
fertility (by subtraction of those dying as infants and children).

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92 J. Lawrence Angel

Population
To set up estimates of population density I shall begin with modern times and shall use
Greece (Steinberg I967; Michalopoulos I932; Stephanos I884) as the key for the Eastern
Mediterranean, since some Classical (Gomme 1933) and some prehistoric (McDonald
and Simpson r969) population data are available for sections of Greece. Based on the
I96I census, Greece has an area, including Crete, the Cyclades and other Islands, of
almost 132,000 km.2 and a population density of 63 -3 per km.2, while the other surround-
ing countries of the Eastern Mediterranean (Turkey, Syria, Lebanon, Israel, Egypt, and
Cyprus) have a combined area of just over 2 million km.2 and a population density of
32-4 per km.2. Since, however, the coastal Levant today has densities over twice as great
as Greece, it seems likely that in past times the balance between Greece and the rest of
the Eastern Mediterranean may have been reversed or at some times about even. In
1940 the density in Greece was 55 per km.2 and in I928, 48 per km.2 (Epstein I944;
Michalopoulos 1932). In the nineteenth century, density was 32-5 in i880, I8-6 in i828
(Peloponnese only), and about 12 in I8oo (Stephanos I884). Stephanos gives for the
Peloponnese a series of estimates based on Turkish census data for the latter part of their
occupation of Greece: in 1777, 7 per km.2; in 1701, 9-3; in I692, 5'4; and 1650, I1'7
per km.2. These may not be completely reliable estimates, but they accord with the
impressions of travelers in Greece and the Levant during the Baroque period, and lead
me to estimate io per km.2 for the phase of Moslem influence as a whole.
For Classical Attica, Gomme (I933), on the basis of censuses, grain imports, and other
data, estimates a population of 315,500 (172,ooo citizens, 28,500 metics, 15,000 slaves)
in 43I B.C. at the start of the Peloponnesian War and before the plague of smallpox (?),
when Athens was dominant and certainly more populous than the rest of Greece; and
258,ooo a century later, at the Classic-Hellenistic period transition, when Athens was
simply the leader of many allied city states; the respective densities are i o km.2 in
431 B.C. and 90 per km.2 in 323 B.C. In I96I, Attica had eleven times the density of the
country as a whole, but the density of the Attic countryside without the city was equiva-
lent to the whole country. In 431 B.C. the Attic countryside had a density of 64 per
km.2 and in 330 B.C. 36 per km.2 (on the bases of urban Athens plus Piraeus populations
of 155,ooo and I68,ooo for the fifth and fourth century dates respectively, as urbaniza-
tion increased). The latter seems to me the conservative estimate to use for Classical
Greece as a whole (including Macedonia and Thrace and Epirus). Another approach,
to argue from a statement by Demosthenes (Sargent I924) that Athenians in the fourth
century imported two-thirds of their grain supply and that in the fifth century this ratio
was at least as high, suggests a general estimate of one-third the i io per km.2 for Attica
in 431 B.C. or just over 36 per km.2. Gomme estimates the number of citizens in 480 B.C.
as I40,000; if there were io,ooo metics (resident foreign merchants and skilled artisans),
and a slave population 50% of the 150,000 free (cf. Sargent 1924) this would total
225,000 or 78 per km.2 at the time when the Athenians had just thrown back two Persian
invasions and the density would more likely be only double rather than triple that of
Greece as a whole. On the whole, therefore, I use 36 per km.2 as the Classical period
density, realizing that fluctuations above the figure are likely.
For the south-western Peloponnese, McDonald and Simpson (I969) over two decades

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 Ecology and population in the Eastern Mediterranean 93

have explored and recorded the number of sites of different periods apparent from surface
observation (sherds and architecture and roads) or from actual excavation; I shall assume
that the area covered is 6,806 km.2, or Elis plus Messenia plus half of Lakonia. For the
Neolithic period they record between seven and seventeen sites - say twelve as a likely
estimate. Weinberg (I965) lists four Middle and Late Neolithic sites and three Early
Neolithic sites in this area, and accordingly I use a factor of 3 to multiply his mapped
sites for the Greek mainland and islands excluding Crete. I take this as an area of 80,832
km.2, using Steinberg's (I967) areas. For the Early Neolithic, if the average population
was I,ooo per site plus environs, Weinberg's 38 sites times 3 means I 14 sites and a popula-
tion density of -4 per km.2, which seems reasonable ecologically. For the Middle and
Late Neolithic periods, I89 (63 times 3) and 375 (I25 times 3) sites at an estimated I,500
people per site give density guesses of 3'5 and 7-0 per km.2 respectively. The twelve
Neolithic sites in SW Peloponnese give a density of 2-6 per km2. For the Early Bronze
Age, McDonald and Simpson list thirty-four sites, and with a population of 2,000 apiece
this gives a density of io per km.2, seemingly reasonable for the whole country. For the
critical Middle Bronze Age, after invasions by Indo-European-speakers, McDonald and
Simpson identify between 80 and Ioo certain sites (o05 maximum, 68 minimum),
at 2,500 per site, suggesting a density of about 36 per km.2, efficiently supported by
plow farming in quite rich and well-watered country, where more sites are located
inland than in the more maritime Early Bronze Age. Caskey (I966) uses Buck's (I964)
count of I60 major sites in mainland Greece and the islands, of which twenty-eight are
in SW Peloponnese. In comparison with about ninety-eight from McDonald and
Simpson's search, this gives a factor of 3-5 (98/28) and hence 560 sites for Greece as a
whole; with average population of 2,500 per site, the mean density is 17.3 per km.2.
For the Late Bronze Age McDonald and Simpson locate I99 sites, and at 3,000 people
per site the density for SW Peloponnese is 88 per km.2, perhaps the maximum which a
proto-urban population in the Eastern Mediterranean can reach without importation
of basic foodstuffs. If the number of sites increased proportionately as much from Early
to Middle Bronze Age for the country as a whole (45%), there would have been 800 or
more Late Bronze Age sites and a general density of 30 per km.2.
For the Early Iron Age's three successive phases, McDonald and Simpson list I3, i6
and 17 sites, at 2,500 per site - a density of 6 per km.2. But this seems too steep a drop in
areas less affected by Dorian and West Greek invasions; the general decline might be
one-third. In Classical times, fifty-three sites in the SW Peloponnese with a population
of 4,ooo each give a density of 31 per km.2, in accord with the area's position under
Spartan rule but perhaps a rather low estimate. In Hellenistic times, forty-seven sites
give a density estimate of 28 and in Roman times, thirty-four sites give one of 20 per
km.2 for the SW Peloponnese, suggesting densities 5 per km.2 higher for Greece as a
whole, at this time including Crete, Macedonia, Thrace. There are several estimates by
historians for population density in parts of the ancient Eastern Mediterranean, and
for the period A.D. IIo, Westfall (I968 and personal communication), a geographer,
gives 49 per km.2 for the Levant as a whole (Roman Greater Syria). This is about
double my estimate for Greece and probably for Anatolia (cf. Beloch i886), as in
modern times. For the Byzantine period density was certainly lower in general and
fluctuating, since it begins with the after-effects of Plague (Justinian's), includes a number

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 TABLE 28

Deaths, longevity, reproductive capacity, population density and ecology

in the Eastern Mediterranean

Population
Longevity of adults estimates
Death ratios to (I5 + years) Per female People
Period adults as Io: Sample Av. age Sur- No. of per en
Dates Infant Child size S ? Births vivors si

I928-29 3 4- 59,000 56'1 54'3 3'9 2'5 48 Rice, bre

A.D. 1900 meat r
Romantic
(4) 4 237 402 37'3 (3-8) (2-2) 25 Rice, her
A.D. I800
meat v
Moslem
(45) 4? 53 33-9 27-8 (3-5) (io8) 1o Rice, her
A.D. I400 sugar
Byzantine (5 +) 5-? 93 37'7 31'I (4.0) (2zo) (20) Bread,
A.D. 600 fish

Imperial Roman (5 +) 3'5 127 40'2 34'3 3'3 I6 25? Bread, he

A.D. I20
(6)
Hellenistic
7? 3 + ? 26 42'6 36'6 3'6 i6 33? Bread, m
300 B.C. (5) balan
Classic
5 3 133 44-5 34-6 4'6 30' (750) 36 Eggs, mea
650 B.c. (5) impor

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Early Iron Age 7'5 3'5 I64 38-8 30'4 4-I I'9 (600) (I9) Bread, cheese
II50 B.C. (7) meat rarely defor
Late Bronze Age 7 4'5 286 39'3
1500 B.C. (IO) (erosi
Middle Bronze Age 8- 5-
2000 B.C. (20) mal
Early Bronze Age
3000 B.C. (34) less meat
Late Neolithic (3??) 3? 4
(Kea) 4000 B.C. Fish le
Middle Neolithic -63 x3 4- Bread, some
5000 B.C. meat
Early Neolithic
6800 B.C. (2I) meat wild and m
tame

Mesolithic (6) 5+ 71 32'0 24-9 (4- 2-) o-o8 Fish,

9000 B.C. grasses mal
Upper Paleolithic 6- 6+
30,000 B.C. vegetable

Note: data for Upper Paleolithic and M

Catal Hiuyutk as well as Nea Nikomed
Karata? as well as from Corinth, Hagio
under births per female are the numbe

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96 J. Lawrence Angel

of dislocating invasions (Avars, Slavs, Albanians, Swedes, crusading Franks, Aragonese,

and Venetians), and ends with Plague becoming endemic, not to mention malaria. In
Moslem times, density was plausibly about half the Byzantine and, as we have already
seen, Stephanos' data suggest about Io per km.2 for the seventeenth to eighteenth
century transition.
The entire fabric of the foregoing series of estimates is shaky, partly because of un-
certainties in guessing the number of undiscovered sites for each period and even more
from the impossibility of guessing the average population of a settlement. Mellaart's
(I967) estimate at (atal Hiiyiik of about 31-45 rooms excavated in one-thirtieth of the
site suggests over I,ooo rooms and, if the average family size was a little over five, the
population in the seventh to sixth millennium B.C. would have been 5-6,ooo. Small
villages of Early Neolithic and later dates would have had only a few hundred. If my
guess of I,ooo for the average early farming trading town is too high, then my serial
increases up to 3,000 in the Late Bronze Age and 4,000 in historic times may be some-
what too flat. Lerna in the Middle Bronze Age plausibly had at least 570-800 people
living within the town walls (Angel I971: 75, based on family size and house number)
and I would assume that outlying farms might have housed an equal number; my
estimate of 2,500 for the average settlement may be high, therefore, rather than low.
What I am trying to offer is a sketch of guessed densities as a demographic skeleton, to
be tested against future archeological and anthropological data, and I do not follow
my own previous estimates (I946, 195I, I969a).
70- -

60 , 55

50- 0

I 50
3~~~~~~~~~~~~~~0~~~ ~50
40-

30- >A I\
~-F~~~~ m
~Z
LU ~~~~~A
j\ j 0
,.
20 - 40
!/
I O
1' 0 o

O 35

10 - 0
~O5
'0 ~
^ 0 ? 30

8000 6000 4000 2000 0 2000

----- BC AD -->-

Figure r6 Relation between population density per km.2 and average lon
of adult females (I 5 + ) in Greece and the Eastern Mediterranean from Mesol
Population density is on a logarithmic scale (left) and age at death on an a

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 Ecology and population in the Eastern Mediterranean 97

Demography
Vital statistics of successive periods shown in table 28 and fig. I6 are based on all Europe
plus the Mediterranean for Upper Paleolithic and Mesolithic, ?atal Hiiyiik and Nea
Nikomedeia (Angel I972a, b) for the Early Neolithic, and Karata? (Angel 1970) as well
as Lerna, Hagios Kosmas, Corinth, and other Greek sites for the Early Bronze Age
(including the scanty Middle and Late Neolithic sample). From then I use only Greek
data. I express infant and child deaths in the first column as ratios to ten dead adults
in the cemetery, because the actual number of infant deaths is often hard to determine.
Also, from Hellenistic times onward, the cemeteries in which we have a secure sampling
of juveniles give small samples. I determine age at death by all available criteria, taking
age 15 as the beginning of adulthood, because age at marriage for women in the ancient
Mediterranean was at or shortly after this time (MacDonnell I913) and menarche
apparently was between 13 and 14 (Amundsen and Diers I969); I am sure that this applied
also in prehistoric times. The number of births for a given female pelvis I determine as
shown in plate 2 from the degree of pitting, grooving, or distortion on the posterior
surface of the pubic symphysis, where stretching of ligaments and hematomas (bruises)
from repeated births change the bone (Angel I969, in part quoting Putschar and Stewart;
Stewart 1971; Gilbert, personal communication); the external pubic surface shows
exostoses and fossae from pregnancy stresses of the rectus abdominis and other wall
muscles. The number of pelves available decreases in later periods, because archeo-
logists in the past, when many Classic to Moslem period cemeteries were excavated, did
not realize that skeletons were worth preserving.
To begin with the earliest changes, the striking thing about the Mesolithic depression
is the short life-span of females, which must have restricted fecundity, though we have
no pelves available to measure this. The Franchthi Cave skeletons (Angel i969b) show
porotic hyperostosis and this would fit the hypothesis offalciparum malaria as a depressant
(cf. Angel J966), as earlier proposed by Zaino and implied by Bruce-Chwatt (i965).
In the Early Neolithic the 2'5 (or 2-6) survivors, shown by the balance of mean parity
and juvenile deaths, imply an increase of 20-30% per generation or i% per year,
astronomically rapid, in fact almost as rapid as present temporary rates for areas like
Ceylon (Wrigley i969). The increase from o.o8 (Deevey I960; for the world estimates,
0-04) in Mesolithic times to I-5 per km.2 involves just over four doublings in about
2,500 years, while in the next 3,000 years three more Neolithic doublings of population
bring us to the Early Bronze level of about io per km.2, a continuation of the rate of
increase which is logarithmic, although plotted arithmetically the curve is concave
upward. Clearly, the rate of increase indicated by the present fertility data from Qatal
Hfiyik and Nea Nikomedeia would be a local and temporary thing apparently resulting
from the striking lengthening of female life-span. This surge of vitality in early farming
times is important to expose for understanding. In the Early Neolithic sample, the adult
incidence of porotic hyperostosis is 43%, of which io% is in the category of marked
diploic thickening, i.e. double the normal blood-forming bone marrow; I assume that
much of this anemia is thalassemia, developed as an evolutionary adaptive response to
newly spreading falciparum malaria (cf. Bruce-Chwatt I965) facilitated by the early
GA

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98 J. Lawrence Angel

farmers placing their trading towns and villages near open water or marsh, where the
ground would be not too difficult to clear (Angel 1966, i967, I971). Sites such as Khiro-
kitia and Kephala (on Kea), in rocky locations, show much less porotic hyperostosis. This
same factor of site location must also have favored the dysenteries and hookworm. Also,
the new diet, with much more cereal carbohydrate, and meat tending to decrease pro-
portionately as population became more dense (Angel i972a, b), promoted growth less
well than in the Upper Paleolithic (stature 7 or 8 cm. less, though 4 or 5 cm. more than in
the Mesolithic). This load of disease and decrease in meat protein show in the lack of
increase in male longevity seen in table 28. Why then does female longevity increase?
Apparently because truly settled life in a fixed town or village, a relatively secure winter
food supply from stored grain, and the high social status of women seen clearly at 9atal
Hiiyiik (Mellaart i967), made pregnancy and family life easier than it was under hunting
conditions.

During the Early Bronze Age, in the third millennium B.C., population growth reached
the top of the logistic curve and ceased or in some cases temporarily reversed. There is no
rise in longevity for either sex. The continued stress from malaria as well as other disease,
and the cereal diet, are factors in lowering fecundity and stature (by 2-4 cm.) although by
this time the addition of newly domesticated olive and grape, and use of the plow, were
improving the diet and malaria was probably decreasing (porotic hyperostosis now 13%),
both from improved farming methods and from the beginning climatic shift from warm
Atlantic with high sea levels to Sub-Boreal. The average of two surviving offspring per
woman fits the apparent stop in population expansion, at the time when population
pressures in steppe country had facilitated massive migrations into the Eastern Mediter-
ranean (and into Europe) of people speaking Indo-European languages; these caused
destructions and resettlings during the last third of the third millennium and beginning
of the Middle Bronze Age. Although the child death rate improves, it is clear that there is
a lag in biological response to ecologic improvement and that maintaining a balanced
polymorphism for abnormal hemoglobins is expensive biologically in pre-adult deaths.
This biological expense is equally true of the Middle Bronze Age, where infant deaths
at Lerna increase (Angel 1971), but by the Late Bronze Age there is improvement in this
respect and porotic hyperostosis decreases from 12% in Middle Bronze Age to 9% toward
the end of the second millennium B.C. The slight dietary improvement certainly lasted
for a good part of the second millennium (cf. Hopf i962), with a large increase in the
inland areas farmed (cf. McDonald and Simpson 1969), as well as irrigation and terracing,
although the size of domestic cattle was starting to decrease (Gejvall i969). The
improved farming plus deliberate draining of marshes, even in areas like the Copaic basin,
added to the effect of the drier Sub-Boreal climate and the lowering of sea level up to
4 m. to reduce breeding areas for anopheline mosquitoes and to reduce malaria. All of the
beneficial factors favored population increase amounting to about i 4 doublings in about
750 years, from after 2000 B.C. to just before the Trojan War, which is only a slight
increase over the previous rate of gain, although in arithmetic or absolute terms the
increase is sudden and enormous, almost comparable to the nineteenth century A.D.
Table 28 makes clear that increasing female longevity (by 21 years) is the direct facili-
tator of the greater fertility needed. Although the sharp rise in average fecundity of
about one more birth per woman does not continue after the Middle Bronze Age, a

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 Ecology and population in the Eastern Mediterranean 99

decrease in infant and child deaths increases fertility. Detailed demographic analysis of
the skeletons from Lerna (Angel I969, 1971) shows a rate of natural increase of 0-3 or
0o4% annually, very rapid, and this may even have increased in the Late Bronze Age,
although our data from family chamber tombs are intrinsically harder to analyze, and
perhaps less reliable, than those from the family burials next to and under houses in the
Middle Bronze Age (Caskey I966), or in pithoi in family plots as at Early Bronze Age
Karata? (Mellink I970; Angel I970). Male longevity rises considerably faster than female
longevity, in response to less malaria and slightly improved diet, but there is less than
i cm. increase in stature, except among the aristocrats (Angel I97I). Heterozygosis from
the really massive biological mixture of the Bronze Age is also a likely factor in the increase
in longevity and fertility and should also have favored increasing body size, unless the
crowding in towns and incipient cities worked selectively against it. Certainly an overall
density of 30 per km.2, which seems to me the maximum possible in early literate and
early urban times in the Eastern Mediterranean, would have produced occasional crowd-
ing stress even though the bulk of the population was rural. It was certainly population
pressure, as well as such factors as the economic and political-military struggles over
land and the new iron metallurgy, which initiated the fall of the Hittite empire, the Trojan
War, the sea raids on the Egyptian delta, and the Phrygian and later the Dorian intrusions
from the north.
Although the middle (or, in the Aegean, Protogeometric) phase of the Early Iron Age,
with its frequent cremation burial, piracy, Homeric warlike adventure, major migrations,
and local separatism, represents something of a return to insular self-sufficiency, until the
spread of the alphabet from the Levant and the revival of regular trade and then inven-
tion of coinage in the Archaic period, there is still considerable unity of culture and it is
likely that some depopulation occurred in the whole area, though perhaps less than in
Greece as shown in table 28. Porotic hyperostosis continued to decrease (4% now) and I
cannot point to any environmental cause for poorer health, except some possible exhaus-
tion of soil. Epidemic disease may have been a factor at the transition from Late Bronze
to Early Iron Age, but in the latter period child, though not infant, deaths lessen.
Nevertheless, longevity decreases, especially for females, where the drop is enough to
cause at least half a birth less per female and accounts for a greater loss in population than
in the Early Bronze Age.
The sudden rise of civilization during late Geometric, Archaic and early Classical
times is as dramatic, striking, and irresistible as any in history and is certainly an outcome
of psychological and biological reaction to the strong mixtures of ideas, of manufactures,
and of people, in the Eastern Mediterranean: Aeolians, lonians, Dorians, Mysians,
Phrygians, Lydians, Lycians, Phoenicians, Assyrians, Egyptians and Etruscans were
directly involved, with Medes, Hebrews, Philistines, and several Italic peoples in the
background. Diet improved, partly from development of better wheats, introduction of
eggs as an added protein (Angel 1946, quoting Michell, Vickery etc.), and probably
larger flocks of sheep, and then largely from increasing trade in food, including oil and
wine in pottery amphorae and especially grain from the Ukraine and North Africa.
Although better diet was a major factor in improving health, there was also a climatic
factor from the added glaciation at the end of Sub-Boreal times; Denton and Porter
(1970) date this from 800 to 400 B.c. and it must have been a factor in lower sea levels.

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1oo J. Lawrence Angel

and lessened marshiness. Porotic hyperostosis drops to % and malaria may have actually
disappeared. Longevity rises very sharply, by over five years for males and somewhat less
for females and there is a significant 2-3 cm. increase in body size. The consequent rise
in fecundity, together with a 30% drop in deaths of infants plus children, results in very
high fertility, as much as a 50% increase per generation, or I 5-2% annually. Since the
actual increase appears to have been almost one doubling between about 700 and 450
B.C., including some immigration into ports and other cities, the fertility data seem to
overestimate, as they did in the Early Neolithic sample; my data on parity, or births per
female pelvis, rest on only five available pelves, although the infant and child death data
depend on the large cemetery at provincial Olynthus (Angel 1971, quoting Robinson).
The actual rate of increase is very little faster than in the Bronze Age.
Slavery is a factor here, since the proportion of slaves increased greatly with the
development of large-scale quarrying and then, in the fifth century, mining as well as
industry, on which economic success depended; but the slaves were relatively short-
lived, especially mine slaves, and their fertility was very low. I have no demographic data
on them.

The Hellenistic period sees another levelling off of population growth, before a decrease
starting in the middle of the period (Tarn 1930), and a steadily increasing movement into
the now very large cities, continuing the trend seen in Gomme's data on the population
of fourth-century Athens versus Attica (I933). Although the populations of Alexandria,
Antioch, Ephesus, Pergamon, and later Rome, were probably two to four times larger
than the I50,000 to 200,000 in Athens, I doubt that the general density in the Eastern
Mediterranean greatly exceeded modern density. Greece and the Aegean may have
begun their population decline sooner than the rest of the area and a return of malaria
(Jones, Stephanos etc., cited in Angel I966, I967, I968) is a likely factor (Io% porotic
hyperostosis) in the decreased fecundity, despite the two-year increase in female longevity,
and perhaps in the apparent increase in infant deaths. Contraception and infanticide are
certainly factors too. Male longevity decreases, though stature does not. Living conditions
and diet for the upper and middle classes (largely sampled by our skeletons) were
apparently as good as in Classical times. In greatly reducing fertility and in shortening
male life span, social factors may have been more important than malaria or other disease
including epidemics; among these factors are the effects of crowded living and the
physical and economic stress of continuous warfare.
Lack of fertility and a declining population are hallmarks of the full Romanization of
the Eastern Mediterranean. By now, the Roman system of slave-farms was breaking
down and the slaves themselves were becoming reservoirs for epidemic as well as endemic
diseases, which had a major effect in reducing population, as well as making people flee
to cities and leaving some farms unworked. Erosion and exhaustion of soil, as well as
deforestation, added to the decline in available food, though bread and a little meat were
enough to maintain the level of body size. The Sub-Atlantic's brief warm and drier
interval, from A.D. o100-400 (Denton and Porter I970), probably raised sea level and
eventually increased marshiness and malaria increased again (porotic hyperostosis rises
to 24%). Probably this climatic shift facilitated both epidemics and the series of invasions
which ended the united Roman Empire. Living and demographic conditions in the city
of Rome (MacDonnell I913) were very much worse than in Greece or in North Africa.

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 Ecology and population in the Eastern Mediterranean IOI

Lead poisoning (Gilfillan I965) as well as malaria probably reduced fecundity and life-
span. MacDonnell's large sample of tombstones shows average adult longevities of 35 8
for males and 30'5 for females, with infant: child: adult death ratio of (8-): 7-: Io. We
have no pelves to estimate fecundity, but if the interval between births averages 4-4 years
(less than in Hellenistic Greece), with marriage for females at I7-8 (MacDonnell I913),
there would have been only 2-7 births per female, actually less than the 2z9 probable
juvenile deaths per female! Thus the population of Rome each generation required over
Ioo% immigration from the rural areas and other cities in order to stay constant. Cities in
the Eastern Mediterranean were probably better off and rural fertilities must have been
higher. But the general picture is of demographic catastrophe, matching the ecologic
destruction which had developed earlier.
The Byzantine period, therefore, starts with a declining population; the decrease in
average longevity for both sexes fits this. Although the demographic picture is at about
the level of the Middle Bronze Age, we lack adequate data on number of infant deaths
and on average fecundity and can only guess that the fertility potential did not go beyond
replacement in general. There was probably a phase of levelling off in the ninth to
eleventh centuries after the first invasion and coincident with the remarkable Byzantine
cultural revival mentioned earlier. Porotic hyperostosis declines to I2%; perhaps there
was less malaria. Plague, however, returned and halved the population in the latter half
of the fourteenth century. The silting of valleys by what Vita-Finzi (I969) calls the
younger fill continued throughout Byzantine and most of Moslem times, probably rather
as a response to deforestation and post-Roman lack of flood control than for reasons of
climate, which was in general slightly wetter than today with cooler winters, except
during the mid-Byzantine warmer spell of the tenth to twelfth centuries.
The Moslem period fits entirely within a minor glacial advance from the fifteenth to
the nineteenth centuries (Denton and Porter 1970), which has been called the little ice
age. This may have restricted farming somewhat, though the new crops, rice and sugar,
seem to have maintained stature despite little meat, while lower taxes and a smaller city
population helped. But there was no restriction on malaria and plague; porotic hypero-
stosis returns to the Early Neolithic frequency, but with fewer marked cases of anemia.
As we saw earlier, population densities are at the Early Bronze Age level partly because of
much smaller cities. Female longevity is even lower than in prehistoric times and fertility
in general sinks below the level of replacement.
The regrowth of population seems to have begun in Moslem times, a little before the
military, political and religious revolution which begins the Romantic period. Between
A.D. I700 and I900 there are two doublings of population, so that the increase is even
somewhat steeper than in early Classical times. Apparently this is a direct result of stimu-
lation from Baroque and then Romantic Europe, plus the new soils brought down into
most valleys, although the Eastern Mediterranean remains largely a rural area until
almost the twentieth century. There is an absolutely amazing increase in longevity,
especially of females, and my guess at excess fertility, without data from pelves or infant
deaths, may be too low. Malaria continues, with adult porotic hyperostosis, of trace to
slight degree, at 4%. Michalopoulos' (I932) data show a huge increase in longevity
expected in the twentieth century and suggest that the same rapid growth rate had
adequate support in fertility, though here the number of births (birth rate 29.o per 1,000

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Io2 J. Lawrence Angel

population) is related to living women in the age range 15-50 over a reproductive span of
32 years; this may underestimate births per female very slightly. The increase is over i%
annually. It is interesting that Michalopoulos' data on adult average age at death still
show a slightly shorter female, as compared with male, life span.

Conclusions

Data from ancient sites and from human skeletal material agree reasonably well regar
changes in population size in the Eastern Mediterranean. Especially in a challen
ecologic setting, population density change depends on fertility, which in turn
on health from diet, from settlement and from cultural balance with ecology.
whole time span from Upper Paleolithic times to the nineteenth century A.D., the av
woman died before menopause and therefore adult female longevity is the majo
tive factor relating health to fertility, although obviously the deaths of infants and
are also important.
Feedback relations are subtle, whether positive or negative, and there is oft
before a given factor (a new crop or food, the plow, endemic malaria, deforestati
clear demographic effect. Thus human destruction of natural resources, for exam
forests and soils of the Eastern Mediterranean, was gradual and produced full eff
about a millennium after its Late Bronze Age beginning.
The positive associations between population density, female longevity, and fe
as here estimated, are extremely strong. The associations between population den
ecologic factors are less strong and more indirect.
Much that I have written is tentative. We need more ecologic and paleoclima
information, more data on food plant and animal remains, and more skeletons ex
like those at Lerna and Karata?, with full demographic sampling in mind.

Acknowledgements
For diverse help in my gathering data for this study (on field trips supported
American Philosophical Society, J. S. Guggenheim Foundation, Wenner-Gren F
tion, N. I. H. Grant A-224, and the Ales Hrdlicka Fund of the Smithsonian), I th
late C. W. Blegen, J. L. Caskey, W. B. Dinsmoor, D. French, M. Gough, M. J. M
the late D. M. Robinson, H. Robinson, H. A. Thompson, and other archeo
and D. Johnson, K. Holland, and my wife.

5.xii. 971 Smithsonian Institution

Washington,

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 Ecology and population in the Eastern Mediterranean I03

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Abstract

Angel, J. L.

Ecology and population in the Eastern Mediterranean

In the mountain, coastal, and river-valley ecologic niches of the Eastern Mediterranean, united
by sea trade and with productivity of grain and olives and animals set by winter rains and
summer dryness, population density depends on health and food as determining, first, adult
female longevity (i.e. time for fecundity) and, second, survival rates of infants and children.
Rapid density increases in Neolithic, Middle-Late Bronze Age, Classic (up over 35 per km.2)
and Modern periods occur apparently with balance or decline of malaria and slight connection
to change of sea level and climate. The major drop in population to io per km.2, in Roman to
Moslem times, lags behind the causal man-produced ecologic destruction (deforestation, ero-
sion, soil-exhaustion). Site-counts and census allow density estimates and study of skeletons
and pubic symphyses allow estimates of longevity, fecundity and survival.

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