Part 15

Biskra lies immediately south of the Atlas Mountains, in the Department of Constantine, 220 kilometers from the Mediterranean and about 400 kilometers from Ouargla. To the northeast of the oasis lie the Aurés and to the west the beginning of the Saharan Atlas, which run south of west across Algeria into Morocco. Just north of the place are small hills and low, jagged mountains—detached spurs from the main ranges. These are the Djebel Bou Rhezal, running nearly northeast and southwest. The highest of them, directly west of Biskra and about 8 kilometers distant, has an altitude of 463 meters. The Bou Rhezal Mountains have a precipitous southern face, but fall away more gradually to the north, where the slope joins a wide and undulating plain. The latter extends to the base of the main Atlas ranges. Southwest of the oasis, and about 2 kilometers distant, a range of rocky hills extends for a distance of about 6 kilometers, or until they join the mountain range of the Saharan Atlas. These hills are called Ed Delouatt. To the south of Biskra, as has already been stated, there extends a vast plain, the reg, which dips gently to the south and drains into the Chott Melrirh, 50 kilometers or more distant. The situation of Biskra relative to the mountains on the one side and the desert on the other, together with its altitude, governs the climate of the place. Except Laghouat no vicinity in the desert proper was seen with so great an amount of precipitation (about 200 mm.) as Biskra, which is indicated by the relatively rich flora. One who has seen only Biskra can not draw conclusions regarding the vegetation or the conditions of plant life of those portions of the Sahara that lie farther to the south, where much more intense conditions of aridity obtain.

The soils of the vicinity of Biskra are various. That of the low hills between the town and the Djebel Bou Rhezal is only a few centimeters in thickness, but in the washes from the hills it is a meter or more. Here the soil is a sandy loam with an admixture of small stones and pebbles. On the flat ground to the north and to the south of these hills it is of a finer texture, approaching the adobe of the southwestern United States. On the reg to the south of the oasis the soil is also fine, and in some places, if not underlying the plain as a whole, there are strata of gravel at varying depths beneath the surface. This soil in places carries considerable salts. It dries to a powder during the long dry seasons and is easily blown by the winds. Owing to outcropping rock, the south face of the Bou Rhezal Mountains has but scant soil, but that of the northern side resembles the soil of the low hills to the south, which has already been characterized. Near the town are dunes of good size. Especially to the southwest the sand banks against Ed Delouatt hills is in large amount. In the opposite direction, but farther from the oasis, the dunes are fairly extensive.

The Oued Biskra is of great importance to the oasis, since it carries water for several weeks of the year and furnishes water for irrigation. Its channel lies about 3 meters, or possibly more, below the general level of the oasis, and possibly in earlier times may not have been so well defined as at present, spreading its waters over its flood-plain during high water. The oued is made up of several tributary oueds which cross the plain north of Djebel Bou Rhezal, unite where there is a pass in these mountains, and finally debouch on the reg to the south of the town, where the channel becomes continually less well defined. Another oued takes its origin in the Bou Rhezal Mountains in several independent branches which unite at a pass in Ed Delouatt hills and extend for a distance of 15 kilometers or more into the reg. One of the feeders of this oued is from hot springs, Hamman es Salahine, about 8 kilometers northwest of Biskra.

PLANT HABITATS OF THE BISKRA REGION.

From the preceding sketch of the leading topographical conditions of the vicinity of Biskra it will be seen that the plant habitats are more diverse than at any other place visited. For the present purpose the habitats may be distinguished as follows: The alluvial desert (reg), which lies on every side of the oasis save the north; low hills adjoining the oasis on the north; Ed Delouatt hills, southwest; the oueds and their flood-plains; the Dj. Bou Rhezal; the hamada (?) lying both to the north and to the south of the latter.

VEGETATION OF THE BISKRA REGION.

A glance over the list of plants which grow naturally in the vicinity of Biskra shows that many are the same as occur farther south, with many unlike these, having affinities outside of the desert proper; also, the number of plants as well as species is greater at Biskra than farther south. This would be expected from the greater rainfall and more diverse topography.

The flora of the Biskra oasis, according to the authors referred to above, consists of 175 or more species. Of cultivated plants there are 25 or more species, the most conspicuous being the date. The other species are mostly the same as have already been noted at other Algerian oases, except that both the peach and the apricot are wanting at Biskra, although cultivated at Laghouat, Ghardaia, etc. On the outskirts of the town the fairly extensive flood-plain is given over mainly to the cultivation of grain, barley predominating.

The hills and mountains, and the bajada at their base, as well as certain oueds with water relations, exposure, and soils different from those of the oasis, have also a very different flora, which, for the most part, is desertic in character. For purposes of comparison, some of the leading characteristics of the plants growing in a half-dozen localities will be given.

As stated above, to the southwest of Biskra there runs a range of hills, Ed Delouatt, to the south and the north of which may be found interesting plants and plant conditions. On the south side the slope (bajada) descends gradually to the great reg, and near the base of the range, in the vicinity of the place where tradition says a Roman town formerly existed, there is a wide, sandy plain, reaching from the Oued Melah, which pierces Ed Delouatt hills, nearly or quite to the western extension of the oasis. There are no large dunes here, but sand billows about a meter in height and sand hillocks diminutive in size. Between these the plain is fairly level. Over this whole tract there seemed to be only one species (Euphorbia guyoniana), but this was fairly abundant. (Fig. 65.) As at Ghardaia and elsewhere, this species grows in small colonies because of its suckering habit, and acts to a small degree as a sand-binder, each group being situated on a sandy hillock. It will be remembered that this species at Ghardaia, as well as at a certain bordj east of that place, had roots which were somewhat fleshy as well as roots which were fibrous, on one and the same plant. I was interested to learn whether similar conditions should obtain at Biskra, since it had been learned, in the case of two species of Opuntia in the Tucson region, that a different habit followed certain differences in habitat. Several specimens of Euphorbia were removed with care from the soil and in no case was it found that the roots were fleshy, but in every instance they were entirely fibrous. It is supposed that the reason for the variation in behavior may possibly be traced to differences in the water relation, as was found to be the case of the variation in cactus roots cited, but no experiments have been made on Euphorbia to prove this.

There is a variety of habitats on the opposite (north) side of the hills, ranging from the large dune (which reaches the summit to the west of the pass through which the oued passes and which must be 100 to 150 meters above the oued) to the flood-plain, with water very near the surface, during rainy season at least. There are rocky slopes, also, and sandy slopes apart from the dunes referred to, as well as a fine clay with sand admixture on the flood-plain. In all of these habitats, except the large dune, the vegetation is actually or relatively abundant. (See fig. 67.) Among the rocks are small shrubs or half-shrubs, and also on the plain below. Here one may find, among other species, Echiochilon fruticosum, Helianthemum sp., Atractylis serratuloides, Gymnocarpos fruticosum, Thymelæa microphylla, Nitraria tridentata, and Acanthyllis tragacanthoides. On the sandy slope was growing a very numerous population of liliaceous forms, mainly, perhaps wholly, Asphodelus fistulosus (fig. 72).

It has been noted that the oued which pierces Ed Delouatt is made up of the confluence of all of the small oueds lying between the Bou Rhezal Mountains to the north of Biskra and the hills lying directly north as well as those (Ed Delouatt) to the southwest. The united oueds reach the base of the hills nearly 1 kilometer east of the pass, and turning abruptly follow the base the remainder of the distance. At the place where the oued touches the range the soil is moist nearly to the surface of the plain (reg) and for several meters back from the oued. Here, then, the water relations of the plants are such as to favor, probably most of the year, the growth of mesophytes (?) or even of hydrophytes, but for some reason there is not much vegetation there; whether large species have been removed as for fuel, have been destroyed by animals, or never existed, was not learned. The most interesting plant found was the well-known parasite Phelypæa violacea, which grows, according to Mobius, on a salsolaceous host. Only a few specimens were seen at the time of the second visit to Biskra, in March, and they were just appearing above the ground. (See figs. 69, 70, and 71.) When removed from the soil the longest specimen was found to have penetrated over 59 cm.; it was 6.5 cm. in diameter at a point 50 cm. from the tip. The plants were exceedingly heavy, being gorged with sap, and appeared to be able to absorb moisture from the wet soil through the white and delicate epidermis of the entire portion submerged. If this is the case the parasitical relation is of especial interest, as the species is wholly dependent on its host for organized foods, but not for water—a condition opposite that found in such semi-parasites as the mistletoes, which obtain water and unorganized foods in solution from the host, but which, save for the fact of attachment, are otherwise independent of it.

To the northeast of Ed Delouatt hills and north of Biskra, but immediately adjoining the town, is an irregular group of low, rounded hills, mostly very arid, which support a scant vegetation of typical desert plants. These hills, without a name, are eroded to a degree and have shallow washes leading from them in every direction. The soil appears to be thin, except in the washes, where it has accumulated to the depth of a meter or more. On the rounded summits it is rather fine, but in the washes there is much gravel and larger stones. Here one finds the greatest range in exposure, and probably also an accompanying difference in the temperature of the air and soil.

In considering the vegetation of the hills it must be remembered that no plants suitable for forage or large enough for fuel would be left untouched, as the biotic factor is quite as much in evidence in modifying the Biskra flora as that of the other regions visited. As the plants are at present, however, and for whatever definitive causes, there is a considerable variation in numbers and apparently also in kinds. An examination of different parts of the hills by which various exposures as well as other conditions are seen bears out the hypothesis. On the southern slopes, particularly the upper portions, the plant covering is especially poor. The population of an area 16 by 16 meters in the upper south slope included 134 perennials and numerous annuals. Of the species, Haloxylon scoparium was the most numerous; there were also Dæmia cordata, Thymelæa microphylla, and Fagonia sinaica (figs. 78 and 80). All of these species were small, so that from superficial examination the area appeared fairly barren (fig. 73).

When one crosses the hills to the north, so that northern and northwestern exposures are seen, a striking difference is immediately apparent. The northern slopes are much richer in perennials than the opposite facing. The population of an area of the size before taken shows this observation to be correct. On a square 16 by 16 meters, with north exposure, 536 living perennials and numerous annuals were found, with Haloxylon scoparium again the dominant species. The species second in number was Ferula vesceritensis, which is to be found chiefly on the north exposure (figs. 74, 75, and 77). There were also a few specimens of Acanthyllis numidica, and many annuals, which, unlike those on the south facing, were not in flower.

On the lower slopes, in the small and open gulches and the lower portions of the washes and their flood-plains, the plants were most numerous and include a variety of forms, with Peganum harmala the most conspicuous and most numerous.

The roots of several species growing on the hills were studied with the following as perhaps the leading results: The root-system of Haloxylon scoparium, as has already been seen to be the case with an allied species in the country of the M’Zab, may be said to be a modification of the generalized type. One specimen, partly exposed by erosion in a wash, had a tap-root over 113 cm. in length (fig. 76). As this root was 4 mm. in diameter where left, and was 8 mm. in diameter at the crown, it may have penetrated much beyond the point where it was left, provided the soil conditions continued favorable. The main root gave off two large laterals, of which one left the parent root 8 cm. and the other 18 cm. beneath the surface of the ground, but there were also numerous filamentous roots about 2 cm. in length, which were borne in tufts. (See fig. 79.) These resembled the deciduous rootlets found on many perennials in the Tucson region and doubtless function quite as they, namely, they are organized at the beginning of the rainy season, operate to increase the water-absorbing area of the plant quickly and greatly, and die as soon as unbearably arid conditions set in.

In the same wash, where the soil was deep with an admixture of small stones and pebbles, the roots of other species were also examined. One of these was Peganum harmala, whose roots were examined at Ghardaia. The root-systems of the plant in the two regions were similar in being generalized. A main root was found extending downward over 61 cm. and it gave off three good-sized laterals, arising from 15 to 27 cm. beneath the surface. The uppermost lateral took a horizontal course. Figure 82 gives a fairly good idea of the general character of the root-system of the species.

There were many specimens of Plantago albicans growing in the same habitat as Haloxylon and Peganum. Its root-system was also studied with the following results: The tap-root is strongly developed. In one instance the slender main root was found to go straight down over 71 cm. Numerous laterals were borne between 8 and 20 cm. beneath the surface of the ground. The species has an interesting habit of propagating vegetatively by means of fleshy stolons. An examination of the stolons showed them to be of very unequal age, some having been lately formed, while others had been organized in previous years and were no longer living. Many specimens were examined to learn the probable service of the fleshiness of its stolons, and the conclusion was that this factor probably enables the species to pass over periods of excessive drought.

Running across country are the Bou Rhezal Mountains, of which the north and south faces were examined in one or two places, on each side somewhat in detail. The south face is precipitous and has little vegetation, but there are many plants on the opposite face, especially near the base, where the soil conditions are relatively favorable (fig. 84). Among the perennials were found a few specimens of Rhus oxyacantha and Zizyphus lotus, which, although dwarfed, were the largest plants seen away from the oasis, being about 1.5 meters high. In another gulch were found the following species: Acanthyllis tragacanthoides, Dæmia cordata, Ferula vesceritensis, Haloxylon scoparium, and other plants unknown. Toward the upper portions of the gulches Haloxylon was seen to be especially abundant, but in the bed of the washes and on their flood-plains Peganum grows in large numbers (fig. 83). In the most favorable places on the northern slope in March the annuals were the most abundant observed in the vicinity of Biskra, although even there they failed to completely conceal the ground. Where the situation was less favorable as regards soil and water conditions (for example, on the side of the gulches, on the summit of ridges, and the like) there were almost no annuals and those present were relatively small.

In this brief account of some of the most striking features of the Biskra flora it should be noted that several habitats are omitted as not being pertinent to the points in view. Especially, nothing has been said regarding the flora of the salt flats along the Oued el Hamman or that of the dunes, since the influences here are largely edaphic, while the present interest lies mainly in the relation between plants and climate; and the flora of the oasis has been largely neglected for similar reasons.

Not only is there in certain regards a larger number of plants in the Biskra region than had been previously seen in southern Algeria, but there are certain types, mostly new, which point to more favorable conditions of plant life. These are such forms as have a water-storage habit, like Asphodelus, Ferula, Plantago, and Phelypæa, which, although not wholly absent farther south, appear to be much more numerous near Biskra. The presence of bulbous plants is well known as being one of the floral characteristics of the High Plateau, and it is also known that similar forms are not to be found where the arid conditions are the most severe, which probably accounts for the facts noted. It may be pointed out also that plants at Biskra exhibit exposure preference where soil conditions appear to be parallel. This condition is not so marked farther in the desert as at Ghardaia, for example, where, provided there is sufficient depth of soil, apparently any species may be found on any exposure. In other words, exposure preference implies a certain amount of water as well as sufficient soil.

So far as shown by observations of the root-systems carried out on similar species growing in the Biskra region and at Ghardaia, the essential root characters of plants growing in the two regions are the same. The single exception to this so far noted is that of the roots of Euphorbia guyoniana, which, contrary to the expectation, are apparently wholly fibrous at Biskra, although they are in part fleshy farther south, where the soil conditions are surely more arid. At Biskra, also, the only plant observed with typical generalized root-systems (Peganum) does not grow where the soil is shallowest, but where it is relatively deep. So far as seen, also, plants growing where the soil is shallow have either a generalized root-system or a root-system approaching this type, even if the branching is only relatively deep.

=GENERAL SUMMARY AND CONCLUSIONS.=

THE ENVIRONMENTAL CONDITIONS OF PLANTS IN ARID REGIONS.

The environmental conditions encountered by plants in the arid regions are widely different from those of moister regions. Precipitation is not only slight, but it shows an enormous range in variation from year to year; the rate of evaporation is high; the temperature of the air and soil varies widely both during the day and with the seasons; the light is of great intensity, and the soil is low in humus content and may contain an excess of salts. These, the most striking physical factors of deserts, are present in different combinations with resultant differences among deserts, and an arid region may be so large as to include such variation within its borders. Also, a desert may be so far from the ocean, or it may include such diversity of topography, as to show great variation in biological features as well as in its surface phenomena.

In the flora of any arid region, the mutual relations of the constituents of the flora and its general and detailed relation to the physical environment are quite different in the main from these features in the flora of the more humid regions. Thus, the leading relation is the relation to water, and on the response of the plants to this relation much of the phenomena associated with plant life in the desert directly depends. For example, in the extreme deserts it is probable that the elements of competition between the perennials, which is an important factor in the survival of a species in the moister regions, is wholly lacking. It should be noted, however, that in the less extreme deserts, as in the vicinity of the Desert Laboratory, competition exists between plants, although this is not at first apparent. In this case the competition is not for room, but for water, and is not manifest by palpable crowding, but by invisible competition of the roots. Thus the reactions are with the physical environment and are exhibited in a variety of ways, some of which concern the plants themselves in an intimate manner, being morphological and physiological, some being concerned with the flora as a whole. The environmental responses are often obscure and complex, but in other instances they are less obscure and apparently direct.

As is well known, a desert flora is in part perennial, lasting with but little outward change from season to season, and in part ephemeral, consisting of short-lived species which appear with the rains and which disappear with the return of the dry season. The ephemeral flora differs in no essential regard from annuals of the moister regions; also, the environment to which they are exposed closely resembles the environment of the annuals of such regions. But the perennial desert flora, on the other hand, offers very striking departures from the corresponding flora of the moister regions, just as the environment to which they are exposed for the most of the year is also different. It will be sufficient, for the purpose of bringing out the point of view, to notice a few of the leading characteristics of desert plants and of their physical environment and, in a few instances, to observe possible relations between the two.