Le Houerou 1997

Climate, ora and fauna changes in the Sahara over
the past 500 million years

Henry N. Le Hou erou
327, rue de J ussieu, F-34090, Montpellier, France
(Received 22 J uly 1997, accepted 10 August 1997)
The present-day Sahara occupies an area of slightly over 8 million km
2
in
Africa, between latitudes 16 and 32 N, circumscribed within the isohyet of
100 50mm mean annual rainfall. The hyperarid area alternately expanded
and shrank on both sides of a seemingly narrow semi-permanent eremitic
zone along the Tropic of Cancer during the course of the Quaternary epoch
(17Ma). The Cenozoic, Mesozoc and Paleozoc Sahara, in turn, has
undergone drastic climatic changes as the African continent drifted north-
ward from its Antarctic position to reach its present latitudinal situation. But,
seemingly the Sahara was never the large desert it now is, with the exception
perhaps of the Upper Triassic Lower Liassic epochs. The Pleistocene and
Holocene contrasting climate changes induced large variations in ora and
fauna distribution, as well as in geomorphic processes.
The ora shifted from that of typical desert to tropical savanna and
Mediterranean forest or steppe, depending on period and location. Fauna, in
turn, changed more in abundance than in nature, since the same groups have
been in existence since the Upper PlioceneLower Pleistocene. Large
mammals, for instance, were mainly of Afro-tropical kinship throughout the
Pleistocene and Holocene, while small mammals, in contrast, were predom-
inantly of Mediterranean origin over the same periods. And such is still the
case. There were varying large degrees in density of occurrence, but relatively
minor uctuations in nature, in response to such environmental changes as
lake and dune expansion and retreat, and even glacier expansion and melting
at higher elevations. The present-day man-made expansion of desertic
conditions to the north and south actually threaten both ora and fauna alike
in the short- and medium-term. Most African large mammals, still present in
the desert until the second half of the 19th century, have now become extinct,
or are on the very verge of extinction in the Sahara (some may be surviving
further south, and/or in the East Africas parks network). The situation,
however, is far less dramatic for the ora, which still includes almost 3000
species of vascular plants, although some species of economic value or not
are in danger from the man-made destruction of their habitat.
1997 Academic Press Limited
Keywords: Sahara; desert; climate change; biogeography; ora; fauna;
natural resources; biodiversity; desertication
J ournal of Arid Environments(1997) 37: 619647
01401963/97/040619+29 $25.00/0/ae970315 1997 Academic Press Limited
Outline of the present geographical situation and subdivisions of the Sahara
Setting the present upper limit of the Sahara on the 100 50mm isohyet of mean
annual rainfall (MAR), proposed by the present author some 40 years ago, has now
received a broad acceptance among climatologists, geographers and biologists (Le
Hou erou, 1959, 1995a).
Our interpretation of the 1:10 million scale colour map of the Ecologically
Homogenous Territorial Units (EHTU) (Popov et al., 1991) results in an area of 814
million km
2
for the 80 Saharan EHTU recognized in this map of acridian habitats
within the 100mm isohyet bond of MAR (Fig. 1).
Moreover, there are many possible subdivisions in our classications of the Sahara
available in terms of climate, environment, geomorphology, ora and fauna, not to
mention crops, livestock and human populations (Monod, 1936, 1973; Capot-Rey,
1953; Cloudsley-Thompson, 1954, 1984a; Dubief, 1959, 1963; Le Hou erou, 1959,
1986, 1990, 1992, 1995; Qu ezel, 1965, 1978; Schiffers, 1971).
The 80 EHTU mentioned above form six broad divisions.
Northern Sahara lowlands: MAR >25mm, winter rains, elevation <1000m
Southern Sahara lowlands: MAR >25mm, summer rains, elevation <1000m
Sahara Highlands: MAR >25mm, winter and/or summer rains, eleva-
tion >1000m
Oceanic Sahara lowlands: MAR >25mm, elevation <1000m, distance from
the Atlantic <50km
Central Sahara lowlands: MAR <25mm, no marked rain seasonality,
elevation <1000m
Red Sea lowland Sahara: MAR <25mm, winter rains, elevation <1000m
Each, in turn, can be subdivided into at least three to four sub-units, as shown in
Figs 2 and 3 (Dubief, 1963; Qu ezel, 1978; Le Hou erou, 1990).
These divisions and their characteristics should be kept in mind when assessing
climatic, environmental and biological changes during the course of Pleistocene and
Holocene epochs: different geographical areas of the Sahara may contemporaneously
have harboured Mediterranean forest, Mediterranean steppe, tropical savanna and
climatic desert. Conversely, a given geographical division may have alternately
supported forest, steppe, savanna or desert in different times. Things are therefore
globally much more complex than the semantic unit desert might suggest; the natural
temptation for oversimplication should, therefore, be strongly resisted if one is to
avoid deeply erroneous conclusions.
The subject is made the more difcult as our present state of knowledge (and
ignorance) varies from one period to the next. Generally speaking, the higher rainfall
periods are better documented than hyperarid periods as the former permit
conservation of much more datable elements of ora, fauna and sediments. There is,
on the other hand, a decreasing gradient of the amount and accuracy of information on
the changes from moist to dry environments, and from Holocene to Upper Pleistocene
epochs.
Climatic changes before the Quaternary period
The African continent started its slow northward move from the Pangaean South Pole
continent in the Upper Silurian and Lower Devonian periods, some 400 million years
ago (Ma).
Figure 1. (facing page). Ecologically Homogeneous Territorial Units (EHTU) (from Popov et al.,
1991).
H. N. LE HOU
EROU 620
CHANGES IN THE SAHARA 621
Ordovician (500440Ma) continental and shallow channel-sea sandstone deposits
outcrop over large areas in the present Sahara (Furon, 1950, 1957; Fabre et al., 1976),
particularly in the Inner Tassili plateaux of the Ahaggar, around the Precambrian core
(Suggarian, Pharuzian). These show evidence of erosion forms developed below and
around a thick ice cap (Inlandsis) similar to those documented in Canada, Greenland
and Scandinavia for the last Quaternary glaciation (c. 20,000 years ago). The Upper
Ordovician glaciation (Caradoc and Ashgill) is actually one of the best documented
worldwide, apart from the Pleistocenes, with striated bedrocks, U-shaped valleys,
tillites, moraines, hydrolaccoliths and other ring-shaped structures (kettles, pingos),
polygonal soils, etc., all typical of permafrost. All channel and glacial deposits show a
slow northward sedimentation drift (Beuf et al., 1971; Rognon et al., 1972; Rognon,
1989b; Fabre et al., 1976).
But the Ordovician glaciation occurred at a time when there was still no terrestrial
vegetation on earth. The chief biological remains found were Bilobites and Tigillites
imprints of mud-earth worms and arenicolous annelids (Scolites), sometimes very
numerous (Rognon et al., 1972). All this bears wittness to the existence of the future
Sahara in the Antartic Pangaea up to c. 440Ma.
When the Ordovician ice cap melted, huge amounts of water were released into a
transgressing shallow, cold, hypo-saline Silurian ( =Gothlandian) Sea characterized by
graptolithic shale and clay deposits. These presently outcrop over large expansions
across the Sahara from southern Morocco and Mauritania through Algeria and Libya
(Fabre et al., 1976). They, in particular, constitute the Intra-Tassilian trough of the
Ahaggar.
The present-day Sahara was then exundated anew, with the deposition of thick
lagoon and river sandy sediments of Devonian age (395345Ma) that constitute the
Outer Tassili ring of plateaux around the nucleus of the Ahaggar massif. These
Figure 2. Biogeographic limits and subdivisions of the Sahara (Le Hou erou, 1990, 1995).
H. N. LE HOU
EROU 622
1 Mediterranean region (semi-arid to hyper-humid zones)
2 Western arid steppic zone
3 Central arid steppic zone
4 Eastern arid steppic zone
5 Oceanic Northern Sahara transition zone
6 Western Northern Sahara transition zone
7 Central Northern Sahara transition zone
8 Eastern Northern Sahara transition zone
9 Oceanic Central Sahara
10 NW Central Sahara
11 Northern Central Sahara
12 NE Central Sahara
13 Western Central Sahara
14 Central Central Sahara
15 Central Sahara Highlands
16 Eastern Sahara Highlands
17 Oceanic Southern Sahara
18 Western Southern Sahara
19 Central Southern Sahara
20 Eastern Southern Sahara
21 Oceanic Sahel
22 Western Sahel
23 Central Sahel
24 Eastern Sahel
25 Saharo-Arabian region, Eastern Sahara Zone, Red Sea shores
26 Sudano-Angolan region, East African & Erythreo-Sabean domain, E-A Montane Zone
Mediterranean region
Ibero-Maghribian Domain
Saharo-Arabian region
Northern Sahara
Saharo-Arabian region & Mediterranean region
Sahara Highlands
Southern Sahara
Sadano-Angolan region
Sahelian domain
Central Sahara Plains
Figure 3. Sketch of the phytogeographic subdivisions of the Sahara and neighbouring territories
(from Qu ezel, 1978; Le Hou erou, 1990, 1995).
Devonian sandstones are rich in iron oxides, red silts and clays as a result of chemical
weathering on a continent that had become covered with terrestrial vegetation
(Emberger, 1944).
Coral sea deposits as well as palo landforms and rock weathering suggest a rainy
climate with a mild to warm temperature. By the Upper Devonian and Lower
Carboniferous periods (Fammenian, Tournaisian, Visean, Namurian (350320Ma))
arboreal ferns, lycopods and horse-tails were already present in the continental
deposits of the now Djado-Ennedi plateaux, suggesting a wet-tropical climate (Batton
et al., 1965; Busson, 1967, 1970, 1972).
Palo-magnetic data suggest the Saharan area was already then in the vicinity of the
Tropic of Capricorn, in a situation somewhat reminiscent of the present geographic
setting of the Karroo or the Kalahari, but with a different climate (Rognon, 1989a).
Since the late Devonian period (345Ma), throughout the Carboniferous epochs,
most of the Sahara was, to a large extent, under a tropical sea until the Permian
(280Ma).
During the Permian and Lower Triassic periods (225Ma), the sea slowly withdrew
from the continent, from west to east, while Africa continued its northward drift. The
terrestrial arboreal ora included ferns, lycopods, horsetails and primitive gymno-
sperms (Gingkoales, Cycadales, Benettitales, Caytoniales, Araucariaceae). The fauna
included dinosaurs (also present in the Karroo at the same time). Sediments included
red clays and lacustrine limestone. All these characteristics suggest a sub-humid to
hyper-humid tropical Saharan environment. In no way does this suggest a tropical
desert, such as that present at the same time further north in Eurasia from Ireland to
Poland, and particularly well documented by North Sea oil geologists (Rognon,
1989b).
The Permo-Carboniferous series constitutes the oor of the continental Intercalary
(CI) and of its eastern Sahara counterpart the Nubian Sandstone (NS) formations.
The CI and the NS are both most reminiscent of the Karroo formation of southern
Africa, with close faunal kinships (Kilian 1930, 1937; Busson, 1970, 1972; Fabre et al.,
1976; Rognon, 1989b; Lefranc & Guiraud, 1990). Paleomagnetic data suggest the
future Sahara was located between the Tropic of Capricorn and the Equator during
the Permian period.
The rst Triassic marine and continental sediments of Buntsandstein and
Muschelkalk (225210Ma) were then topped with thick saline deposits of evaporites
(gypsum, anhydrite, sodium chloride, dolomite) of the Keuper (Upper Trias,
210190Ma) and Lias (Lower Jurassic, 190175Ma).
All these suggest an arid climate, similar to large parts of the world during the same
periods. Paleomagnetic information infer the future Sahara was between 5 and 10S
by the Lower Jurassic. Such latitudes correspond today with a tropical climate such as
in the Sudano-Zambesian ecozone, while South America had not yet begun its
westward drift off Africa. The CI and NS spread over time from the Permian to the
Upper Cretaceous (Maestritchian) periods. They outcrop over some 12 million km
2
;
about two-thirds of which is located in the eastern Sahara (NS). NS also outcrops over
05 million km
2
in the eastern Sahel of the Sudan Republic. The CI contains one of the
largest aquifer in the world (Guiraud, 1988). It is present over 08 million km
2
of the
northern Sahara of Algeria, Tunisia and Libya, north of the Tropic of Cancer. The CI
aquifer yields some 95m
3
s
1
of good to fair quality water from 110 boreholes and
many foggaras ( = qanats), with a total reserve estimated 6 10
13
m
3
(Gischler, 1979;
Pallas, 1980; Lefranc & Guiraud, 1990; Margat, 1992; Guiraud & Bellion, 1995).
The NS aquifer of Kufra covers an area of 18 million km
2
, producing 4m
3
s
1
of
excellent water (500p.p.m. >total dissolved solids) seemingly dating back to a Palo-
Nile. The reserve is still unknown, but apparently huge. CI and NS waters seem much
older than previously thought (UNESCO, 1972); according to recent isotopic
36
Cl
datation they now appear to be aged over 30,000 years on the margins of the aquifer
H. N. LE HOU
EROU 624
and up to 100,000500,000 in the central trough faults (Guiraud, 1993; Guiraud &
Bellion, 1995).
The ora of the CI and NS was studied by Batton et al. (1965) and Busson (1967,
1970, 1972), and reviewed by Lefranc & Guiraud (1990). This review is summarized
in Table 1.
The recorded fauna includes 12 species of bivalves, 20 species of dinosaurs, shes,
crocodiles etc. Lefranc & Guiraud (1990) conclude their review The Flora of the CI
includes vegetal remains belonging to biotopes typical of high mountain forest, arid,
semi-arid and mainly humid Mediterranean and tropical environments. All of them
evoke warm climatic conditions with alternating dry and wet seasons, sometimes
prolonged dry and short wet seasons and semi-arid conditions. Data obtained from
animal remains have a similar signicance: warm climate, variable wetness, depending
on region. Continental sweet water shes, dominate, associated with estuarine species
including skates and sharks. Reptiles include tortoises, crocodiles, large snakes and
many species of dinosaurs. The fossilized plants and animals of the CI indicate a wide
variety of biotopes that evoke a vast and complex territory, comparable to the present
African continent. One constant fact throughout the region, however, was a tropical
Table 1. Summary of theundifferentiated fossil oras of theContinental
Intercalary and Nubian Sandstoneformations (after Lefranc & Guiraud, 1990)
TriassicLiassic DoggerMlm
(210175 Ma) (175136 Ma)
Order Family Species
Fern Allies
Equisetales 2 2
Lycopodiales 1 1
Ferns
Filicales 6
Gymnosperms
Bennettitales 1 3
Caytoniales 1 1
Coniferales 5 9
Araucariaceae 2 1
Cupressaceae* 1 1
Ephedraceae 1
Pinaceae 1
Podocarpaceae 1 2
Taxaceae 2
Taxodiaceae 2
Cordaitales 1
Cycadales 1 2
Gingkoales 1
Angiosperms 4
Ebenaceae 1
Lauraceae 2
Nymphaeaceae 1
*Subtribe Cupressineae.
Subtribe Abietineae.
Includes Cinnamomaceae.
Table 2. Tentativesketch for correlating landmarks of theUpper Pleistoceneand Holocenein thenorthern southern Sahara
Europe
(Penck & Brckner,
1901, 1905, 1909;
Climatic uctuations Blytt, 18761909;
Years Human activities and Sernander, 1908, 1910; Years
B.P. S Sahara N Sahara industries Mangerud et al., 1974) Fauna Flora and vegetation B.P.
Present Hyperarid Hyperarid Settlement of nomads, Extinction of Increasing scarcity of trees Present
urbanization, man-made large mammals and shrubs; Chenopodiaceae
desertization Amaranthaceae
40 Mediterranean 40
450 Little Ice Age Presence of Forest remnants in the 450
African large north and Highlands
mammals
2200 Historic Period; aridity Cameline Period Sub-Atlantic Mediterranean 2200
3000 Arid to semi-arid Equine Period Sub-Boreal Forest in north and Highlands 3000
Highlands; tropical
3500 Climatic aridization Bovine/pastoralists Rich and Forest and savana in the 3500
diversied south
5500 Tafolian Rharbian Bovine/Round Heads Atlantic Afro-tropical Gramineae and Cyperaceae 5500
semi-arid semi-arid Protomediterranean
Cromagnods
6500 Nouakchottian Rharbian; Buffalo/Hunters Mediterranean forest and 6500
optimum semi-arid to tropical Savanna Gramineae
sub-humid and Cyperaceae
8000 Short dry spell Neolithic Boreal ? Chenopodiaceae 8000
Amaranthaceae,
Artemisia
10,500 Semi-arid to sub-humid Capsian Pre-Boreal, Afro-tropical Mediterranean forest and 10,500
Chadian (Protoneolithic) Younger Dryas tropical savanna
Allerd
12,500 Beginning of Beginning of Iberomaurusian Blling ? Chenopodiaceae, Amaranthaceae 12,500
semi-arid semi-arid (Epipaleolithic, and Artemisia
build-up of build-up of Protomdit. Cromagn.)
Ogolian dune major sand (Homo sapiens)
system seas
H
.

N
.

L
E

H
O
U
E
R
O
U
6
2
6
Table 2. Continued
Europe
(Penck & Brckner,
1901, 1905, 1909;
Climatic uctuations Blytt, 18761909;
Years Human activities and Sernander, 1908, 1910; Years
B.P. S Sahara N Sahara industries Mangerud et al., 1974) Fauna Flora and vegetation B.P.
19,000 Hyperarid; Hyperarid; Aterian (Paleotithic) ? Mediterranean forest and 19,000
Inchirian wet Soltanian (H. neanderthalensis ??) Afro-tropical tropical, savanna
period pluvial-erosion
cycle
40,000 Beginning of Wurm 40,000
wet period
70,000 Hyperarid; Soltanian (H. erectus) ? Chenopodiaceae, Amaranthaceae 70,000
Symm. erosion dry period; and Artemisia
sand ridges sand seas?
125,000 Wet/warm Tensiftian (LevalloisianMousterian) Eemian Inter- Afro-tropical Mediterranean forest and tropical 125,000
pluvial glacial Optimum savanna
150,000 Hyperarid; Hyperarid; Acheulean (H. habilis) Riss ? Chenopodiaceae, Amaranthaceae150,000
leveled-down sand seas? (Paleolithic) and Artemisia
sand sheets
C
H
A
N
G
E
S

I
N

T
H
E

S
A
H
A
R
A
6
2
7
climate. The opening of the South Atlantic had only just begun during the mid-
Cretaceous (120100Ma). The proximity of the American and African continents
produced a continental effect which accentuated the semi-arid character of northern
Africa. Palaeomagnetic data, in good agreement with paleoclimatic and paleogeo-
graphic information, place the centre of the CI deposition directly under the probable
trace of the palaeo-equator during the Jurassic and Lower Cretaceous.
The Lower Cenozoc period (6550Ma) was characterized by a large marine
transgression of a tropical sea that covered most of the Sahara and North Africa
(Guiraud, 1978; Guiraud & Ousmane, 1980). The Continental Terminal (CT) is a
formation of various ages from the mid Eocene (Lutetian) to the Quaternary epochs,
but mostly Mio-Pliocene (2617Ma). It consists of unconsolidated sands, silts,
conglomerates, clays and contains bauxite deposits, iron and lateritic hard pans
(Siderolitic strata) (Guiraud, 1978; Guiraud & Ousmane, 1980; Lang et al., 1990;
Guiraud & Bellion, 1995). CTs bauxite, iron and lateritic hardpans occur up to
2023N (e.g. in the Umm Ruwaba formation of Kordofan). These sediments testify
for a humid equatorial climate in these areas at the time of deposition since bauxite,
iron and lateritic hardpans are nowadays only being laid down in Africa south of Lat.
8 N, i. e. in the Guinean eco-zone (Rognon, 1989a; Le Hou erou et al., 1993). At the
same time, in the present north-western Sahara of Algeria and Morocco, there took
place the deposition of thick lacustrine limestone, the Hammada formation, containing
gasteropods (snails), Characeae and small mammals, all typical of a semi-arid to sub-
humid climate.
The CT is the store formation of very important aquifers in the northern Sahara and
arid zones of Algeria, Tunisia and western Libya. This CT aquifer discharges some
215m
3
s
1
, of which 85 is from 2000 boreholes and 135 from natural evaporation in
the great salt lakes of the AlgerianTunisian border (Gischler, 1979). The recharge is
estimated to be 18m
3
s
1
from runoff on the Saharan Atlas.
The Sahara was at the present latitudes of the Sudanian ecozone (515N) during
the Upper Cretaceous and Eocene epochs (10025Ma). It reached its present
latitudinal situation (1632N) during the Neogene, c. 1020Ma. The northward
drift was then c. 25cmyear
1
; 4cmyear
1
is currently being recorded in the Ethiopian
rift.
The Sahara during the Quaternary Period
During most of the Quaternary Period (16M years), the Sahara had a number of dry
wet cycles, the causes of which are not clearly understood. There is, however, a
consensus that these cycles relate to the strength and weakness of the Saharan
anticyclonic pressure zone and, therefore, to advances and retreats of the Polar Front
(PF) and the Intertropical Convergence Zone (ITCZ). Why the Saharan high pressure
belt is alternately strong and weak is not clear. It may be connected with oceanic
temperatures (Leroux, 1976, 1983; Kutzbach, 1981, 1987) and currents, and with the
Pacic Oceans Southern Oscillation that seems to control El Ni no and drought
phenomena around the world. It is now generally admitted that the forcing of the
earths orbital processes, along with the Milankovitch theory (Milankovitch, 1930,
1941), plays a major role in the wetdry cycles with a periodicity of 100,000, 41,000
and 21,000 years, perhaps in combination with shorter cycles of solar activity
(Bernard, 1962, 1986; Lamb, 1977; Berger, 1978, 1981; Berger et al., 1984; Lorius et
al., 1985; Lorius, 1989; Broecker & Denton, 1990).
There seems to have been at least eight to ten major drywet cycles since the Upper
Pliocene. Thirty years ago, some scientists believed that European glaciations, related
to the Penck & Br uckner (1901, 1905, 1909) chronology for the Alps, were
contemporary with the Saharan pluvial periods and with the Blytt-Sernander
H. N. LE HOU
EROU 628
chronology (Blytt, 1909; Sernender, 1910) for the Upper Pleistocene and Holocene of
Scandinavia. Then came the Dubief-Balout theory (Dubief, 1951; Balout, 1955) of
the climatic swing of the Saharan edges. According to this theory, there are periods
when the Polar Front moves south, alternating with periods of a northward advance of
the ITCZ. Consequently a dry period in the northern Sahara would have corresponded
with a wet period in the south and viceversa. But there is now an increasing tendency
to assume that the weakness of the Sahara anticyclone may produce a southward move
of the Polar Front and, at the same time, a northward march of the ITCZ. The
hyperarid nucleus of the Sahara, along the Tropic of Cancer, would therefore have
alternately expanded and shrunk over a NS distance of 10002000km. The absolute
dating chronology shows an almost perfect matching of the Saharan hyperarid periods
with the higher latitudes glacials and the correspondence between a semi-arid Sahara
and the interglacials (Fig. 5) for the Mid- and Upper Pleistocene and the Holocene.
However, at present we are in a period that is both interglacial in the higher latitudes
and hyperarid in the Sahara, thus contradicting the theory. Furthermore, fossil ergs
expanded 400600km south of the present southernmost limit of drifting sands in the
Sahel, whereas there is no fossil dune system of any substantial size north of the present
limit of the Sahara. It follows that the limits of the Sahara do not seem to have shifted
further north from their present position during the Quaternary, contrary to the
southern limit which moved considerable distances southward (Fig. 4).
ThePliocene, theLower and Mid-Pleistocene
The Sahara seems to have had a semi-arid to sub-humid tropical climate during the
Cenozic Epoch. Geological and palaeontologic data suggest a wet equatorial climate in
the Lower Eocene, shifting in the Oligocene towards a Sudano-Guinean type of
savanna, with a clear-cut dry season (Maley, 1980, 1981). The rst evidence of aridity
is known in the Mid-Upper Pliocene of the southern Sahara from aeolian deposits and
fossils of xerophytic vegetation (Retama cf. raetam, Tamarix spp.). A period of aridity
also occurred in North Africa during the Villafranchian, between the Pliocene and
Pleistocene. This bore witness to deteriorating tropical conditions and the progressive
disappearance of the archaic Neogene fauna (Elephas africanavus, Stylohipparion,
Libytherium and Macharodus). Some large tropical mammals (Rhinoceros simus,
Hyaena striata, Alcelaphus bubalis, Gorgon taurinus prognu, Taurotragus derbyanus)
together with present-day arid and semi-arid zone species still exist. The Villafranchian
fauna of An Brimba, near Kebili, Tunisia, for instance, includes: Gazella dorcas,
Ctenodactylus gundi, Lepus kabilicus, Oryctolagus cuniculus, Canis aureus, Meriones cf
shawi, J aculus sp., Elephantulus rozetti, Hystix cristata, Struthio camelus, Acinonyx
jubatus, Ammotragus lervia) (Joleaud, 1935; Arambourg, 1949, 1952; Arambourg &
Coque, 1958). Most of these were part of the Saharan fauna less than 100 years ago
and a few are still to be found in the desert today. But the presence of a few species
(Ursus arctos, Rhinoceros mercki, Bos primigenius) suggests that conditions were
somewhat colder than now.
The Villafranchian ora of northern Tunisia shows a shift from tropical Pliocene
(21%) to Mediterranean species (52%) and boreal elements (21%). Some 47% of this
ora is still present (Quercus faginea, Q. suber, Q. ilex, Olea europaea, Cetatonia siliqua,
Laurus nobilis) along with some boreal species which no longer exist in North Africa
(Fagus sylvatica, Ulmus scabra).
It is probable that the major sand seas of the Sahara began to be established at that
time.
It also corresponded with a thick generalized limecrust north of the tropics, the
Villafranchian or Moulouyan limecrust, also called the Salmon crust of Helicidae
(because of its colour and the frequent presence of snails within it) (Durand, 1953;
Figure 4. Climatic and biogeographic variation in the Sahara and neighbouring zones over the
past 20,000 years (in part from Duplessy et al., 1989a, b): (a) The Sahara 18,000 years B.P.; (b)
The Sahara 8000 years B.P.; (c) The Sahara today.
H. N. LE HOU
EROU 630
Choubert et al., 1956; Coque, 1962). Limecrust and gypsum crust formations seem to
correspond with transition periods between pluvial and arid conditions (Coque, 1962).
It was at one time believed that the erosion period of each sedimentary cycle
corresponded with arid to semi-arid climatic conditions, whereas the deposition part
Figure 5. Tentative palaeoclimatic chronology of the Pleistocene and Holocene in the Sahara
(after Rognon, 1989b).
corresponded with pluvial periods (Alimen, 1955; Biberson, 1961; Coque, 1962;
Chavaillon, 1964; Rognon, 1967; Conrad, 1969; Camps, 1974). The times of
deposition in these cycles were thought to be contemporaneous with the alpine
glaciations of Penck & Br uckner (1901, 1905, 1909), while the periods of arid erosion
were equated with interglacial periods. These views are being strongly challenged.
Most specialists now consider that glacial is equivalent to dry and interglacial to
humid.
The Lower Pleistocene period is characterized by the Pebble Culture of Aus-
tralopithecinae and of Homo habilis. The long period of Acheulean industries, the
Gunz-Riss interval (1,000,000150,000B.P.), is little known, although Acheulean
industries are widespread, even in the most arid parts of the Libyan and Egyptian
Deserts (McCauley et al., 1982, 1986; Petit-Maire, 1982). At least three cycles of
erosion and sedimentation took place during this period. These were the Moulouyan,
Saletian and Tensiftian, with two or more limecrust formations in the Moulouyan and
Tensiftian, and two or more formations of gypsum crust (Choubert et al., 1956;
Coque, 1962). Homo erectus, known from East Africa, has not so far been reported
from the Sahara. Levalloisian and Mousterian artefacts (80,00050,000B.P.) are very
rare, unlike the Acheulean (500,000100,000B.P.) and Aterian (40,00020,000B.P.)
industries.
The fauna and ora do not seem to have changed much in respect to the Upper
Villafranchian. Among the animals present were: Loxodonta africana, Rhinoceros simus,
Equus mauritanicus, Hippopotamus amphibius, Homoceras antiquus ( = Alcelaphus
bubalis =Bubalis antiquus), Alcelaphus buselaphus, Bos primigenius, B. ibericus, Gazella
dorcas, G. cuvieri, G. rufrons, G. atlantica, Ammotragus lervia, Papio sp., Arvicanthis
sp., Meriones shawi, Gerbillus campestris, Hystrix cristata, Serengetilagussp., Canis aureus,
Crocuta crocuta, Hyaena stritata, Phacochaerus aethiopicus, Sus scrofa, and Camelus
dromedarius. Most of these belong to the Afro-tropical fauna, suggesting a dry tropical
climate.
The ora was predominantly Mediterranean with a number of tropical and
temperate elements which, again, did not change very much after the Upper
Villafranchian and continued throughout the rainy periods of the Pleistocene and
Holocene. Of course, there were differences according to latitude and altitude; the
Sahara mountains showed a more temperate climate ora (Tilia, Alnus, Acer, Betula,
Quercus spp.). Throughout the Pleistocene rainy periods some of the most common
specie were: Ephedra sp., Cedrus atlantica, Pinus halepensis, P. laricio, Cupressus sp.,
J uniperus oxycedrus, Taxussp., Platanussp., Salix spp., Corylussp., Alnusspp., Betula
sp., Quercus afares, Q. ilex, Q. coccifera, Q. mirbecki ( = Q. faginea), Q. suber, J uglans
regia, Tilia spp., Sapindus sp., Argania sp., Pistacia atlantica, P. lentiscus, Rhus sp.,
Tamarix sp., Acacia tortilis subsp. raddiana, Ziziphus lotus, Helianthemumspp., Cassia
spp., Erica arborea, Olea europaea, and Fraxinus spp. (Pons & Qu ezel, 1958; Qu ezel,
1960; Van Campo, 1975; Beucher, 1971; Cour & Duzer, 1976; Brun, 1979, 1983,
1985, 1987, 1989; Maley, 1981; Van Zinderen Bakker & Maley, 1979). This suggests
a vegetation similar to that of the present humid Mediterranean climate of North
Africa, with somewhat cooler temperatures (Alnus, Corylus, Tilia, J uglans, Betula,
Taxus, Cedrus) on the mountains, and remnants of the Pliocene tropical ora
(Sapindus, Argania) in the lowlands. Vegetation seems to have remained semi-arid to
arid in the plains during the pluvials; it was probably comparable to the present-day
arid steppes of North Africa, to the north of the Tropic of Cancer, and to that of the
Sahel to the south of the Tropic (Acacia tortilis subsp. raddiana, Ziziphus lotus,
Helianthemumspp., Combretaceae, Cassia spp.). There is, however, little data on the
dry and hyperarid periods because ecological conditions were not conducive to
conservation of plant and animal remains; but arid and hyperarid periods are
demonstrated by the interstratication of aeolian deposits between the lake and
organic deposits of the wet periods.
H. N. LE HOU
EROU 632
TheUpper Pleistoceneand theHolocene(Table2)
The Mid- and Upper Pleistocene correspond roughly with the two last alpine
glaciations of the Riss, Wurm and post-W urm age (Fig. 5). In North African
chronology, these are referred to as Soltanian and Rharbian erosionsedimentation
cycles (Choubert et al., 1956). In terms of human artefacts they correspond to
Acheulean, Mousterian and Aterian (Mid-to Upper Palaeolithic). The post-W urm
ages (12,500B.P. onwards) correspond with Aterian, Ibero-Maurusian (Epipalaeo-
lithic), Caspian (Protoneolithic) and Neolithic civilisations. These civilisations are
known from many sites throughout the Sahara (Balout, 1955; Camps, 1974; Hugot,
1974), including Neolithic rock paintings and engravings (Fr obenius & Obermeier,
1925; Fr obenius, 1937; Lhote, 1958; Mori, 1965; Hugot, 1974). Palaeontological
data are particularly numerous from 12,500B.P. onwards (Butzer, 1966; Faure, 1966;
Butzer et al., 1972; Van Campo, 1975; Cour & Duzer, 1976; Rognon, 1976a, b; Brun,
1979, 1983, 1985, 1987, 1989; Petit-Maire, 1979, 1986, 1989; Maley, 1981; Dutour,
1984, 1988; Fontes et al., 1985; Faure et al., 1986; Petit-Maire & Dutour, 1987;
Fontes & Gasse, 1989; Wendorf et al., 1990). This situation is in part due to the older
limit of reliable
14
C dating. It is well established that in the southern Sahara and the
Sahel, at least eight to ten arid and wet periods occurred over the past 125,000 years.
The older dune system (150,000B.P.?) is composed of levelled sands. The second
system (125,00070,000B.P.) forms longitudinal, symmetrical (NESW), erosion
sand ridges (Michel, 1973; Mainguet, 1976, 1982, 1983, 1984; Rognon, 1989a, b).
The third Ogolian or Kanemian system (12,00020,000B.P.) is made up of
transverse (NWSE) asymmetrical, depositional dunes, extending some 400600km
further south than the present drifting sands (Hunting Technical Services, 1964;
Chamard, 1973a, b; Michel, 1973; Servant, 1973; Wickens, 1975; Rognon, 1976a, b,
1980; Mainguet, 1976, 1982, 1983, 1984; Servant & Servant-Vildary, 1980; Maley,
1981). The fourth dune system of barchans and barchanoids dates back to 3000 years
B.P. onwards; it is a reshufing of the two former, the third being a reshufing of the
second and so forth.
Pollen analysis from sea sediments has shown there has been a dry period in the
northern Sahara between 18,000 and 24,000B.P. (i.e. slightly older than the Ogolian
transverse sand dunes of the Sahel) (Brun, 1979, 1983, 1985, 1987, 1989; Brun &
Rouvillois-Brigol, 1985). Then an optimum pluvial, during the Neolithic
(10,0004000B.P.) was followed by a return to arid, then hyperarid conditions from
3000B.P. onwards (Faure, 1966; Petit-Maire, 1979, 1982, 1986, 1987, 1988). Semi-
arid to sub-humid climatic conditions prevailed throughout the Holocene in the
lowlands, along the Tropic of Cancer until c. 3000B.P. These facts are established by
sedimentation studies and lake levels. The Mid-Holocene Sahara, including the
present most arid areas, contained a large number of lakes [Lower Saoura Valley
(Touat, Ahnet), Taoudeni, Araouane, the Fezzan (Shati, Murzuk and Wadi al Ajial
Basins)] (Chavaillon, 1964; Faure, 1966; Conrad, 1969; Hebrard, 1972; Elouard,
1973; Servant-Vildary, 1977; Sarnthein, 1978; Rodhenburg & Sabelberg, 1980; Petit-
Maire, 1982; Petit-Maire & Riser, 1983; Callot, 1984, 1987; Stein & Sarnthein, 1984;
Faure et al., 1986; Pachur & Kr opelin, 1987; Fabre & Petit-Maire, 1988).
Other evidence is afforded by ubiquitous rock paintings and engravings (Fr obenius &
Obemeier, 1925; Fr obenius, 1937; Lhote, 1958; Camps, 1961, 1974, 1980; Mori,
1965; Hugot, 1974). Five Neolithic periods have been recognized: (a) the Buffalo
[Homoceras(Bubalus) antiquus] or Hunters period c. 90006000B.P.; (b) the Bovine
period (65002500B.P.), subdivided into three sub-periodsRound Headed
(65004500B.P.), Bovine proper (50004000B.P.), and Horsemen sub-period
(35002000B.P.); and nally (c) the Cameline Period (2200B.P., onwards). The
Hunters period and early Bovine period are characterized by melanodermic people
similar to the Fulani and to East African pastoralists, while the late Bovine (Horsemen
Table 3. Floristic richness of various parts of theSahara and neighbouring areas
Log species
Surface numbers
No. of area Sp. per
Region species (10
3
km
2
) 10
4
km
2
Log area (km
2
) References
Northern Sahara 1100 1000 110 050 Le Hourou (1990)
Southern Sahara 850 1500 57 047 Le Hourou (1990)
Central Saharan Lowlands 500 3000 17 042 Le Hourou (1990)
Saharan Highlands 830 500 166 051 Le Hourou (1990)
Western Sahara 870 3700 24 045 Le Hourou (1990)
Oceanic Sahara 450 70 643 055 Le Hourou (1990)
Eastern Sahara 1700 3600 47 049 Le Hourou (1990)
Sahara (overall) 2800 8000 35 050 Le Hourou (1990)
Fezzan 230 600 38 049 Corti (1942)
Mountains 850 500 170 051 Quzel (1954, 1957, 1958), Kassas (1956),
Bruneau de Mir & Gillet (1956),
Gillet (1968), Hassan (1974)
Djourab (1) 50 150 33 033 Quzel (1965)
Tnr (2) 20 200 11 025 Quzel (1960)
Majabat (3) 7 150 05 016 Monod (1958)
Tunisian Sahara 480 100 480 054 Le Hourou (1959)
NW Sahara of Algeria 380 12 317 063 Guinet (1958)
(Beni-Abbes)
Ahaggar 350 150 233 049 Quzel (1954)
Tibesti 568 200 284 052 Quzel (1958)
Tassili 340 90 384 051 Leredde (1957)
Moroccan Sahara 350 350 100 046 Guinea (1949), Guinet & Sauvage (1954),
Mathez & Sauvage (1974),
Birouk et al. (1991)
Ar 410 85 482 053 Bruneau de Mire & Gillet (1956)
Egyptian Sahara 1000 1000 950 105 050 Tckholm (1974), Boulos (1975, 1995)
(without Sinai)
H
.

N
.

L
E

H
O
U
E
R
O
U
6
3
4
Table 3. Continued
Log species
Surface numbers
No. of area Sp. per
Region species (10
3
km
2
) 10
4
km
2
Log area (km
2
) References
Ennedi 528 70 754 056 Gillet (1968)
N African Steppes 2220 628 354 058 Le Hourou (1990, 1995)
Mauritania 1000 800 130 051 Monod (1940), Lebrun (1977), Barry & Celles (1991)
Libya 1800 1760 102 052 Boulos (1975), Le Hourou (1975, 1988, 1990, 1995)
C
H
A
N
G
E
S

I
N

T
H
E

S
A
H
A
R
A
6
3
5
and Cameleers) represent leucodermic people the People of the Sea, invaders from
LibyaEgypt and probable ancestors of the Kel Tamasheq (Tuareg). Leucodermic
protomediterranean cromagnods, closely related to the makers of the Epi-palaeolithic
Ibero-Maurusian and Capsian industries of the Maghrib, survived until proto-historic
times in the central, western and southern Sahara and in the Nile Valley; physical
anthropology data suggest that these same Cro-Magnods might have been the
ancestors of the Guanches, aboriginal (now extinct) inhabitants of the Canary Islands
(Petit-Maire, 1979; Dutour, 1984, 1988; Petit-Maire & Dutour, 1987; Wendorf et al.,
1990).
In the northern Sahara, wet periods corresponded with the extension of a
Mediterranean-type of vegetation characterized by the dominance of pollen from
Mediterranean trees and shrubs (e.g. Quercus ilex, Q. coccifera, Quercus suber, Pistacia
lentiscus, Pinus halepensis, Cedrus atlantica, Olea europaea, Phillyrea sp., Myrtus
communis, Abiessp., Ceratonia siliqua, Laurus nobilis, Rhus coriaria, J uniperus oxycedrus,
Tetraclinis articulata, Cupressussp., etc.) These are species of the present vegetation in
the semi-arid and sub-humid zones of northern Africa (Libya, Tunisia, Algeria,
Morocco) (Van Campo, 1957; Leroi-Gourhan, 1958; Gruet, 1960; Santa, 1960; Van
Campo & Coque, 1960; Brun, 1979, 1983, 1985, 1987, 1989; Brun & Rouvillois-
Brigol, 1985). The arid and hyperarid periods corresponded with vegetation
dominated by Artemisia and Chenopodiaceae/AmaranthaceaePoaceae steppes,
respectively (Beucher, 1971; Maley, 1973; Van Campo, 1975; Cour & Duzer, 1976;
Van Zinderen Bakker & Maley, 1979; Van Zeist & Bottema, 1982; Brun, 1985, 1987,
1989).
The situation was similar in the highlands of the Central Sahara, but with a stronger
contribution of temperate climate species in the higher altitudes (Abies, Cedrus,
Corylus, Fagus, Ulmus, Erica arborea, Alnus, Daphne, Fraxinus, Quercus mirbecki, Q.
afares, J uglansand Tilia).
During the arid and hyper-arid periods, a ora similar to that presently known in
similar zones developed: Ziziphus lotus, Acacia spp., Retama, Tamarix, Ephedra,
Chenopodiaceae, Artemisiaspp., Compositae, Brassicacae, Cistaceae, Zygophyllaceae,
Cyperaceae, Maerua, Balanites, etc. (Pons & Qu ezel, 1957a, b, 1958; Qu ezel &
Martinez, 1958, 1960; Maley, 1973, 1980, 1981; Van Campo, 1975; Cour & Duzer,
1976; Petit-Maire & Riser, 1983; Ritchie et al., 1985; Neuman, 1987; Neuman &
Schulz, 1987; Schulz, 1987, 1988).
Most of our knowledge relates to the southern Sahara. The humid periods
supported a ora that included many strong Mediterranean inuences (Quercus, Pinus,
Fraxinus, Platanus, Cupressaceae, Alnus, Tilia, etc.), mixed with Sahelian and
Sudanian elements (Combretaceae, Grewia, Acacia spp., Hyphaene, Commiphora,
Maerua, Balanites, Bauhinia, Celtis integrifolia, Hymenocardia, Salvadora, Cadaba,
Capparis, Diospyros, Uapaca, Olea hochstetteri, O. europaea subsp. cuspidata (syn. O.
africana, O. chrysophylla) and subsp. laperrinei (syn. O. laperrinei), Syzygium,
Tamarindus, Lannea, Alchornea cordifolia, etc. (Maley, 1981). Conversely the arid and
hyperarid periods were dominated by Sahelian and Saharo-Arabian elements,
respectively. The latter included Tamarix, Cornulaca, Cleome, Zygophyllum, Artemisia,
Pentzia, Tribulus, Launaea, Plantago, Aerva, Euphorbia, Chrozophora and Polycarpaea
(Maley, 1981).
Present-day ora, vegetation and fauna
Flora and vegetation (Figs 13, 6; Table3)
The ora and vegetation of the Sahara may be subdivided into ve main entities: the
northern Sahara with a ora and vegetation belonging to the Mediterranean region and
H. N. LE HOU
EROU 636
closely correlated with an extreme form of the Mediterranean climate. The southern
Sahara, conversely, belongs to the Sudano-Deccanian region and the Palaeotropical
oristic and ecoclimatic zone, closely tied, in turn, to an extreme form of the tropical
climate. The central Sahara plains are characterized by intense aridity without any
dened rainfall regime. The ora and vegetation are strictly Saharo-Arabian, i.e. a
mixture of Holarctic and Palaeotropical elements showing a high degree of adaptation
to aridity. Perennial plant communities are restricted to runoff or to the presence of
ground-water. The Saharan highlands and mountains bear many similarities with both
the northern and the southern Sahara. Mediterranean elements are, however,
dominant above 10002000m, mixed with tropical species and representatives of the
archaic pan-African Rand ora. The Atlantic Sahara is an attenuated form of coastal
desert including both Mediterranean and tropical species and a remarkable degree of
endemism.
Fauna
The situation is more complex than that for the ora, relationships depending, to a
large degree, on the taxonomic groups considered. Large mammals, ungulates and
carnivores are predominantly of tropical origin; they belong to the so-called Afro-
tropical fauna, formerly called Ethiopian fauna. This has been the situation
throughout the Quaternary; they therefore represent a continuation of the Mid-
Pliocene fauna (see below). Conversely, small mammals, particularly rodents, are
essentially of Mediterranean origin. The same is true for birds: 80% are Palaearctic
(Heim de Balsac, 1936; Dekeyser & Derivot, 1959; Casselton, 1984; Le Berre, 1989).
Reptiles are derived almost equally from Palaearctic and Palaeotropical elements. The
same applies to shes (Lambert, 1984).
Coleoptera are predominantly Mediterranean, whereas Palaeotropical elements
predominate in ants and termites (Bernard, 1954). Arachnids, spiders, Solifugae and
scorpions show a predominantly Mediterranean afnity (Cloudsley-Thompson,
1984a, b).
Tropical elements of both plants and animals reach higher latitudes along the
Atlantic and Red Sea shores. Many species reach 30N and beyond in southern
Morocco and eastern Egypt (and further north-east along the Rift Valley as far as the
Dead Sea). This is probably due to the milder temperatures in these regions as
compared with the central Sahara (Le Hou erou, 1989b, 1990). Winter temperatures
play a role at least as important as seasonal rainfall patterns in the distribution of both
plants and animals. Mediterranean species and communities tend to dominate under
low winter temperatures, even with summer rains, in elevations above 1500m between
the latitudes 18 and 28N. This occurs in the Tibesti, Ar and Gourgueil. Conversely
tropical species tend to dominate in areas with mild winter temperatures, despite a
Mediterranean rainfall regime, as in south-west Morocco, on the shores of the Red Sea
and the Arabian Gulf. In all these areas the mean daily minimum temperature of
January (m) is above 7C: the distribution of Saharan Acacia and tropical grasses
corresponds to the m isotherm of 5C and above; virtually all tropical species
disappear when m drops to 3C or below (Le Hou erou, 1959).
(For further information on the present day ora and fauna, see Le Hou erou, 1992,
pp. 812, and Le Hou erou, 1995a,b).
Conclusions
The biological history of the Sahara appears throughout the Pleistocene and Holocene
to have been a series of wetdry periods corresponding with alternating expansion and
shrinking of a more or less permanent arid to semi-arid nucleus in the lowlands along
the Tropic of Cancer. The wet periods are much better documented than the dry spells
because organic remains are better preserved during sedimentation cycles than during
erosion phases. For the same reasons the later wet periods of the Neolithic are better
documented since previous sediments have often been eroded. The advent of Th/U
isotopic dating techniques in the 1980s showed, however, that many lake deposits were
much older than previously thought (80,000120,000B.P. vs. 30,00040,000B.P.),
on the basis of erroneous
14
C dating (Fontes et al., 1985; Causse et al., 1988; Fontes
& Gasse, 1989; Gasse et al., 1990). There are, however, a few striking facts, some of
H. N. LE HOU
EROU 638
which seem paradoxical. The fauna changed little from the Lower Pleistocene until
about 100 years ago. The Afro-tropical fauna of large mammals, elephant,
hippopotamus, giraffe, addax, oryx, gazelle, lion, cheetah, leopard, hyaena, white
rhino, hartebeest, wildebeest were common until c. 2000 years ago and quite a few
species managed to survive until 80100 years ago. These include the ostrich and the
crocodile. In early historic times, this fauna extended to Mediterranean northern
Africa (lion, elephant, oryx, addax, hartebeest, leopard, etc.). It is probable that these
animals were reduced in numbers during the dry or hyperarid periods, but they
managed to survive in refuges and thrive again when favourable conditions returned.
The situation has now drastically changed since a large number of species have become
extinct, or are at the verge of extinction in the Sahel and East Africa. This is a result
of the impact of an exponentially growing human population and extensive hunting. So
far few protective measures have been enforced. Most protected areas exist only on
paper and wildlife is often persecuted by those who are supposed to protect it (Le
Hou erou & Gillet, 1985). Somewhat paradoxically, the ora and vegetation did not
follow the pattern of the fauna. Mediterranean and temperate species were in existence
in the northern Sahara and the highlands throughout the Pleistocene and Holocene,
with periods of expansion during the wet phases and retreat during the dry periods.
This contrasts again with the situation in the Pliocene, when relicts from the tropical
climate of the Oligocene and Miocene remained dominant. In the southern Saharan
lowlands tropical elements seem to have dominated throughout the Pleistocene and
Holocene. The situation thus seems to have been analogous to the present: a northern
Mediterranean ora and vegetation opposed to a Palaeotropical ora and vegetation to
the south; along the lowlands bordering the Tropic of Cancer, a nucleus of more or less
permanently arid or hyperarid conditions in which both Mediterranean and tropical
inuences played an alternating role, thus developing throughout the Quaternary, the
Saharo-Arabian phytogeographic entity. This explains why the Saharo-Arabian
element lacks taxonomic individuality. It contains elements from both Mediterranean
(65%) and tropical (35%) oras. Perhaps also time was too short (2 million years) for
an original ora, with endemic tribes and families, to develop.
Figure 6. (facing page). Some typical patterns of distribution of plants and animals in the Sahara and
adjacent areas (Le Hou erou, 1990). (1) Mediterranean zone: semi-arid to hyper-humid. Flora: Quercus
ilex, J uniperus oxycedrus, Pistacia lentiscus. Fauna: Macaca sylvana, Cervus elaphus barbarus, Sus scrofa
barbarus, Ciconia ciconia, Erithracus rubecula, Pica pica, Lacerta lepida, Acanthodacylus vulgaris. (2)
Mediterranean and steppic zones. Flora: J uniperus phoenicea. Fauna: Gazella cuvieri, Pterocles orientalis,
Passer hispaniolensis, Passer domesticus, Carduelis carduelis, Prinia gracilis, Parus caeruleus, Tarentola
mauritanica, Chamaeleo chamaeleon, Coluber orentulus, Ophiops elegans, Vipera libetina. (3) Arid steppic
zone. Flora: Stipa tenacissima, Lygeumspartum, Artemisia campestris var. glutinosa, Artemisia campestris var.
cinera, Seriphidiumherba-alba=Artemisia herba-alba, Helianthemumlippii, H. sessiliorum. Fauna: Cursorius
cursor, Emberiza calandra, Naja haje, Echis melanogaster. (4) Saharan and steppic zones. Flora: Stipagrostis
pungens, Retama raetam. Fauna: Zorilla libyca, Gazella dorcas, Hyaena hyaena barbara, Meriones spp. Gerbillus
spp., Psammomys obesus, Uromastix acanthinurus, Varanus griseus, Agama agama, Cerastes cerastes, Chlamidotys
undulata. (5) Saharan zone, sensu stricto. Flora: Calligonumcrinitumsubsp. comosum, C. azel, C. arich.
Fauna: Fennecus zerda, Felis margarita, Vulpes r ueppelli, V. pallida, Passer simplex, Oenanthe leucopyga,
Ammomanes cincturus, Corvus rucollis, Stenodactylus sthenodactylus, S. petrei, Cerastes vipera Psammophis
schockari, Phenops sepioides. (6) Northern Sahara and steppe zones. Flora: Hammada schmittiana,
Anabasis articulata. Fauna: Ctenodactylus gundi, Ammomanes deserti, Oenanthe deserti, Emberiza striolata,
Acanthodactylus pardalis, Lythorhynchus diadema. (7) Northern Sahara. Flora: Anthyllis sericea subsp.
henoniana. Fauna: Ctenodactylus vali, Ramphocorys clot-bey, Scotocera inquieta, Oenanthe lugens, Coluber
choumovitchii. (8) Southern Sahara. Flora: Schouwia thebaica, Balanites aegyptiaca. Fauna: Lycanon pictus,
Gazella rufrons, Hirundo obsoleta, Bitis arietans. (9) Central and southern Sahara. Flora: Maerus
crassifolia. Fauna: Oryx dammah, Addax nasomaculatus, Gazella dama mohor, Procavia capensis, Corvus albus,
Streptopelia senegalensis. (10) Mediterranean, Saharan and Sahelian zones. Flora: Tamarix aphylla.
Fauna: Ammotragus lervia, Capra ibex, Canis aureus, Lanius minor.
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Climate, ora and fauna changes in the Sahara over

the past 500 million years

Etudeg eomorphologique. Paris: A. Colin. 476 pp.

Ecologiques et Floristiques sur la V eg etation de la Tunisie

Ecoclimatiques et Biog eographiques sur les Zones Arides (s.l.)

Epoque Actuelle. Travaux et Documents, no 129. Paris:

Ecologiquedes Biotopes du Criquet P` elerin

Etudedela Floreet dela V eg etation du Hoggar. Monographie

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