THE BIOTIC FACTOR.

It will be well to summarize some of the main facts regarding another

and important environmental feature of the Saharan plants, namely,

the relation to herbivorous animals. In the western Sahara, wherever

there is any forage, animals which subsist on it are to be found. Of

the wild animals the gazelle is probably the most numerous and the

most destructive. All travelers across the desert have noted the

presence of this animal. In the northern Sahara Tristram remarked its

abundance fifty or more years ago, and it may be frequently seen by

the traveler at the present time. In addition to the native animals,

the domestic animals, especially the sheep, goats, and camels, are

very numerous, very destructive of plants, and range great distances

for food. As a result, an area around every well or oasis, extending

as far as 40 kilometers or even much more than this,[38] is repeatedly

grazed over and has been utilized in this manner for centuries. As a

result only the poisonous or the distasteful species, or the plants

especially well armed, are left undisturbed to grow and reproduce,

while the balance are more or less consumed, frequently so much so as

to be quite unrecognizable. From the large number of camels, sheep,

and goats which range the desert pastures it might be concluded

that the leading types of plants to be found would be such as are

not eaten by them, but this is not the case. On the other hand,

possibly the most generally consumed form is _Haloxylon_, which

grows on the oued flood-plain, the reg, and the hamada. Thus,

so far as this type is concerned, there is probably little or no

diminution in numbers because of the attacks of animals.

A similar conclusion would doubtless be drawn after study of other

forms, but there lies at least one notable exception, namely,

the influence of animal grazing on the distribution of the betoum

(_Pistacia atlantica_). The betoum, which is the largest arboreal

species in the Sahara, is confined to the region of the Dayas;

that is, to the country immediately south of Laghouat. The tree

is unarmed and is eagerly sought after by all herbivorous animals

for its foliage and tender twigs. Owing to the presence of such

animals, wild and domesticated, the young tree would have no chance

to survive were it not that, growing in association with it, is

the jujube (_Zizyphus lotus_), which is armed and is not eaten by

any animals. The jujube affords safe protection for the seedling

betoum, and in its capacity as nurse prevents predatory attacks by

animals during the critical period. The survival (and probably the

distribution as well) of the betoum is mainly conditioned on the

presence of its protector.

When I first visited southern Algeria it seemed improbable that any

portion of it, or at least any portion that I should be likely to

see, would exhibit the possibilities of plant growth as unaffected

by herbivorous animals. But finally there were found two classes of

plant formations in which animals either had not intruded at all,

or not to a harmful degree. One of these is the wide flood-plain

of the Oued M’Zab, or one of its tributaries, lying about 50

kilometers east of Ghardaia. Despite the fact that the flocks are

very numerous in the vicinity, and that the flood-plain is on the

regular caravan route between Ghardaia and Ouargla, there are few

signs of grazing. The entire plain is so well covered by shrubs

that the vegetation gives the tone to the landscape—a rare thing

in the desert. Here one finds _Retama retam_, _Genista saharæ_,

and _Ephedra_ sp., as well as other species in fair abundance. Thus

the plants are not only numerous, but are of a good size. It should

be observed that on either side of the flood-plain, on the hamada,

scarcely any vegetation may be found.

The second formation referred to is that of protected areas,

especially at Ghardaia, which have been little disturbed at any

time, and portions of them not at all disturbed for centuries. These

are the cemeteries. Such areas are situated not only in the floor

of the M’Zab Valley, where the plant conditions are relatively

favorable, but also on the hamada, where they are relatively very

unfavorable. Both in the valley and on the hamada, as well as on

the valley wall between the two wherever there chances to be soil,

the plants are relatively numerous and of fair size. This fact has

been detailed under the section on Ghardaia and need not be more

than mentioned here.

From these two general observations (exceptions to the usual

conditions), that on the flood-plain of the Oued M’Zab and the

cemeteries at Ghardaia, it is concluded that the grazing of animals

has had a very marked influence in modifying the flora of southern

Algeria. So far as could be told from the limited opportunity to

observe, the modification has gone along on at least two lines,

which are, of course, closely related. The size of the plants eaten

is much under normal for the particular locality, and at the same

time the capacity of the plant for reproduction has been greatly

lessened. That such species as are not touched by animals have not

spread more rapidly, or even have not become the dominant forms,

is an interesting problem, and one that would have to be worked out

for each species. It can be suggested, however, that the restricted

distribution probably lies in the fact that the struggle of desert

plants is mainly with an adverse physical environment rather than

with one another, and that such conditions would not be affected

by grazing animals. For example, large shrubs do not occur on the

hamada, hence _Tamarix_, _Zizyphus_, and _Rhus_, as well as _Peganum

harmala_, are limited to situations where the soil is fairly deep

and the water relatively favorable. _Euphorbia guyoniana_ also

occurs only in sandy soil, and the number of plants limited in

their distribution to soil characters is necessarily a large one.

COMPARISON OF SOME GENERAL FEATURES OF THE VEGETATION OF SOUTHERN

ALGERIA AND OF SOUTHERN ARIZONA.

When we compare the most striking characteristics of the vegetation

of the Algerian Sahara with that of the Tucson region where

the Desert Laboratory is situated, we find some interesting

differences, which may be summed up in the terms “desert”

and “semi-desert,” as applied to the two widely separated

regions. What is meant by these terms will be apparent from the

following short characterization:

Passing into the Sahara from the Saharan Atlas, over the route which

I followed, one encounters a great variety of topography, of which

the most extensive may for the moment be classed as plains. The

plains are divided into three well-marked regions, that of the

dayas, the Chebka, and the Gantara (hamada). The topography is

further diversified by oueds and their flood-plains and by low,

flat-topped mountains. On the northern portion of the plains one

encounters a sparse population of low perennials, and as Ghardaia

is approached the plains vegetation becomes continuously poorer

until at Ghardaia there appears to be none. On the hamada between

Ghardaia and Ouargla areas are to be crossed, several kilometers

in width, where perennials are wholly lacking. The decreasing plant

population of the plains, until it entirely disappears, is entirely

due to the increase in aridity as one goes from the mountains to

and across the Ghardaia-Ouargla country. The low mountains are

almost entirely barren. The flood-plains of the oueds, however,

support a surprisingly luxuriant population of perennials.

Should we contrast the topography and vegetation of the Algerian

Sahara with that of southern Arizona we would find little that

is similar and much that is different. The wide-stretching plains

(bajada) of southern Arizona are well covered with perennials of

good size. The water-courses are fringed with trees, and often an

open forest is to be found on the flood-plains. The low mountains

have a fairly dense plant population, partly of trees, and the

lower mountain slopes are often covered with a mixed flora of

shrubs and trees. It may be said that there is probably no large

area in southern Arizona, where the soil conditions are favorable

for plants, where the water conditions are too meager to support a

perennial flora of some sort. The greater aridity of the northern

portion of the Sahara is evident, therefore, from the great contrast

in its flora.

In crossing the plains of southern Algeria one is likely to call a

region barren when close inspection will show that this is not the

condition. In fact, it was found that areas on the plain, 16 by

16 meters in extent, carried as many as 330 or more perennials,

although a casual glance did not reveal the presence of any

conspicuous vegetation.[39] Both of these conditions are the immediate

result of the small rainfall. The reason for the large number of

plants in certain areas, as above noted, lies in their small size,

since it would probably be difficult to find an equal number on

this area were the plants as large, for example, as in southern

Arizona. The fact that the perennials are inconspicuous is in part

because they are small and in part because the leaves are either

absent or greatly reduced.

When viewed somewhat more closely, one finds other features

in which the flora of southern Arizona and of southern Algeria

are unlike. Travelers, botanical as well as non-botanical, have

described the armed condition of the Saharan plants until the

impression is general that such plants as persist from season to

season are usually well provided with spines. What may be the

proportion of armed to unarmed plants in the northern Sahara I

do not know, but to a person familiar with the plants of southern

Arizona, where spinose forms are very numerous, the Algerian plants

do not appear especially well protected. As this appears to be a

general condition, it is scarcely an accident that the spines of

the American species of the genus _Zizyphus_, for example, are much

better developed than are those of the Algerian representative of

the same genus. From the circumstance that grazing by wild as well

as by domestic animals is very destructive in Algeria, apparently

more so than in Arizona, where the results of grazing are scarcely

to be noted, the general facts regarding spininess in plants, as

given above, suggest the really small influence such animals play

in shaping such a character in desert plants.

Finally, it need only be remarked that plants with a water-balance

are wanting in southern Algeria, and that they constitute one of

the striking features of the flora of the southwestern United States.

[Plate 2:

Illustration: Fig. 4. Shoot-habit of Acanthyllis tragacanthoides.

Laghouat.

Illustration: Fig. 5. Shoot-habit of Zollikoferia spinosa. Laghouat.

Illustration: Fig. 6. Acanthyllis tragacanthoides on sandy plain.

Laghouat.

Illustration: Fig. 7. Zollikoferia spinosa in habitat, plain

(hamada). Laghouat.]

[Plate 3:

Illustration: Fig. 8. Detail of north slope of Nomad Mountains where

Zollikoferia spinosa is the dominant species. Laghouat.

Illustration: Fig. 9. Vegetation of plain (hamada) at Tilrempt. The

conspicuous shrub is Haloxylon articulatum.

Illustration: Fig. 10. The Daya of Tilrempt from the plain, showing

the character of the depression.]

[Plate 4:

Illustration: Fig. 11. Near view of the Daya of Tilrempt. The

fortified stage station, bordj, and a nomad camp are to be seen. The

flattened, level, lower surface of the trees is the effect of

grazing, mainly by camels.

Illustration: Fig. 12. Jujube (Zizyphus) shrubs and betoum (Pistacia)

at the Daya of Tilrempt.]

[Plate 5:

Illustration: Fig. 13. A young specimen of betoum in the midst of a

protecting jujube, at the Daya of Tilrempt.

Illustration: Fig. 14. Leaf and shoot habit of the jujube (Zizyphus

lotus). Daya of Tilrempt.

Illustration: Fig. 15. Leaves of the betoum (Pistacia atlantica),

from the Daya of Tilrempt.]

[Plate 6:

Illustration: Fig. 16. South wall of the valley of the Oued M’Zab at

Ghardaia.

Illustration: Fig. 17. Detail of an eroded bank of the Oued M’Zab at

Ghardaia. The overhanging stratum is hardpan similar to the “caliche”

of southwestern United States.]

[Plate 7:

Illustration: Fig. 18. Shoot-habit of Haloxylon articulatum, from the

plain near the Daya of Tilrempt.

Illustration: Fig. 19. Agriculture at Ghardaia. The fields are

divided into plots about 3 by 4 feet in size, for the more economical

use of water. Young barley is shown growing (November).]

[Plate 8:

Illustration: Fig. 20. Vegetation in upper part of a “draw” on plain

north of the M’Zab Valley Ghardaia.

Illustration: Fig. 21. Situation of square No. 2. on plain north of

valley at Ghardaia.]

[Plate 9:

Illustration: Fig. 22. Capparis spinosa at base of valley wall at

Ghardaia.

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