B The Peruvian-Chilean Desert

Introduction
In South America there are two very exten-
sive subtropical deserts. One is the Monte
Desert in the Argentine (p. 255); the other is
the Peruvian-Chilean desert which forms a
Fig. 3.37. Map of the Pacific coast of Ecuador and,
from 8 S to 28 S, the coastal desert in Peru and
Chile. Fine hatching shows the foot of the Andean
chain
narrow strip along the Pacific Coast, extend-
ing from about 8 S across the tropic of
Capricorn, to about 28 S (Fig. 3.37). It lies in
the rain shadow of the high Andean chain,
which, rising to more than 6000 m, shelters
this desert from the SE trade wind. Another
important factor determining the character
of this desert is the cold Humboldt or Peruvian
current off the west coast of South America.
The rotation of the earth results in a west-
ward deflection of this current, so that north
of 30 S it is fed by cold water brought from
the depths of the ocean to the surface. The
current leaves the co ast only at ab out 4 S, at
Cabo Blanco, and flows westwards to the
Galapagos Islands. The temperature of the
water on the Chile an and Peruvian coasts is
thus relatively low, 17C in winter and 24C
in summer.
The older literature on this area includes
the geographically-oriented works of Ber-
ninger (1925) and Troll (1930); the climate
was described by Dberitz (1967). the vege-
tation of the Chile an part by Reiche (1907)
and Schmithsen (1956), that of the Peruvian
part by Weberbauer (1911) and Rauh (1985).
A more recent review on Peru has been pre-
pared by Koepcke (1961). The description
which follows is based mainly on the latter,
since we have no personal experience of this
region.
The Peruvian Desert
1 Climate
Under the influence of the cold Humboldt
current the climate of this desert is always
cool. Mean monthly temperature range is
5.T-7.7C. March is the warmest month,
September the coolest.
The Peruvian-Chilean Desert
Monthly means:
Callao (on the coast)
September
17.9C
Lima (30 km inland) 23C 16.6C
The latitude has little influence on the
temperature: the temperature difference be-
tween Callao and the harbour Mollendo, 5
further south, is only 5C.
Rainfall along the coastal strip is almost
too small to be measured. At Casa Grande
(7 45' S) it fluctuates between 8 and 33 mm,
but rain in measurable quantities fell on only
17 of 69 rainy days. In Lima the measurable
rainfall from 1943 to 1947 was between 11
and 51 mm. Mean values calculated over
many years are thus not very useful. This is
especially true of the driest part in northern
Chile.
Schmidthsen (1956) gives the following
data for rainfall:
Arica: In 39 years there were only 4 years
with more than 2 mm of rain (annual mean
O.6mm).
Iquique: In 49 years there were only 17
years with more than 2mm of rain (annual
mean 1.9mm).
TRUJ l lLO(60m)
J
......
.. .. .. .. .. ..
..................
L.. ._._._._._ ... . -.... : .. :.:.: .. : .. : .. : .. : .. : .. : .. : .. : .. : .... ..
.. "." ......................................... .
:::::::::::::::::::::::::::::::::::::::::::: :
............................................
.. .. .. .. .. .. .. .. .. .. .. .. .. .. ..........
"." .................. + .. "
Ll MA(I58m)
115- 18J
263
Tocopilla: In 16 years there were only 6 with
more than 2 mm (annual mean 3.8 mm).
This must be borne in mind when the
climatic diagrams are studied (Fig. 3.38).
The seasonal distribution of the rare bouts
of rain is shown in Table 3.11: in the most
northerly parts of the desert the rain falls in
summer, the central region has rain at any
time of the year, while in northern Chile, if
rain falls at all it is during the winter months.
The rare "nifto" (= Christ Child) years are
an exception: these are years when, from
Ecuador to far into the desert, rain falls at
Christmas. In such years there is stronger air
circulation in the tropics so that the cold
Peruvian current is replaced by a warm
equatorial current trom the Gulf of
Guayaquil and extremely heavy rain falls on
the coast. The water flows off the hard sur-
face crusts of the desert soils, causing vast
floods. The last such catastrophic year was
1982, when the floodwaters in the most
northerly part of the desert even washed
away concrete bridges over gullies that are
normally almost completely dry.
ANTOFAGAS TA m )
[9 - In
.. ............ ..
.. ................ ..

Fig. 3.38. Climatic dia grams for stations in the coastal desert of SW Peru and N Chile (Antofagasta). The
rainfall data are very different from those in Table 3.11; this is commonly the case in arid areas when
data are presented for different years
Table 3.11. Rainfall (in mm) recorded at meteorological stations from north to south of the Peruvian
coast: stations situated somewhat inland are shown in brackets (from Ellenberg 1959)
Station m Summer Winter months Summer Year
NN (total)
Jan Feb Mar Apr May June July Aug Sept Oet Nov Dee
Chicalayo 31 0 0 0 0 0 0 1 0 5
Trujillo 26 0 0 0 0 0 0 0 0 0 0 0 3
Paramonga 6 0 0 0 6 3 2 0 0 0 15
(Lirna) 137 0 0 3 5 6 7 6 2 34
Ganete 36 0 0 9 3 4 3 3 1 12 40
Pis co 6 0 0 0 0 0 0 0 0 0 0 2
Lomas 10 0 0 0 0 0 0 0 0 2 0 0 3
(Tacna) 457 3 2 0 2 4 3 3 7 8 6 3 2 43
264
s.W.
Zonobiome III: Subtropical Deserts (The Arid Zonobiome)
N.E.
Over 20'C
800
Rare
- ... .. _-:.
\000
'100
200
Rare only lasting untll morning)
Aboutt7C .
__
o '=-- TIliandsias
: Pacific -
Fig.3.39. Schematic profile through a Peruvian coastalloma to show the frequency of fog (garua) and
distribution of vegetation during the southern winter (after Ellenberg 1959)
Rain is, however, less important for the
vegetation of this coastal desert than fog,
known as "garua". This forms above the
cold ocean current and is blown inland by
the south-westerly winds that arise as a re-
sult of the greater warming of the land mass.
In some places the cordilleras rise to 1000 m
dose to the coast and here the fog is carried
upwards across the slopes. As it rises, the air
cools and the fog becomes denser and satu-
rated with even larger water droplets. Thus
the wetting effect, small on the flat beach,
increases with altitude up to 700 m NN;
thereafter it decreases as the greater part of
the water content of the air has condensed
(Fig. 3.39).
These wetting fogs occur during the
winter months, May to October, but espe-
cially in July and August, while in the sum-
mer the air over the land warms up so rapid-
ly that there is hardly any condensation.
This garua or winter fog makes possible the
development of the "Ioma" vegetation, so
that this region is less desertlike in character.
Since the fogs rarely pass over the coastal
cordillera, the extreme, barren desert begins
behind this mountain range. Further north,
on the coast of Ecuador, fog also occurs
(Diels 1937), but in this rainy area its effect is
less noticeable.
2 Soils
As in other deserts, these are mostly raw
soils.
3 Sub division of the Vegetation
3.1 Peruvian Desert
The sub division of the vegetation of the
Peruvian part of the desert is shown in Fig.
3.40. The following components can be dis-
tinguished:
1. An area of desert along the co ast, con-
sisting of the flat coastal plain and the west-
ern slopes of the coastal mountains; this re-
gion is affected by the fog.
2. An inland, extreme desert, lying be-
tween the coastal mountains and the lowest,
western slopes of the Andes; this area is not
reached by the garua fog, has almost no
rainfall, and is a barren rock desert. On the
western slopes of the Andes rain fall in-
creases with altitude and the desert is re-
placed by Orobiome ur, which extends to
the nival belt of the highest peak (p. 272).
The islands lying off the coast are the
breeding grounds of sea birds and are cover-
ed with guano; for this reason they have no
vegetation.
3.1.1 Subdivision oi the Fog Desert
This is shown in Hg. 3.39. On the coast itself,
fog is rare, occurring only at night and form-
ing dew.
The sand of the beach is barren except on
dunes, where narrow strips of large species
of Tillandsia (Bromeliaceae) occur. The Til-
landsia rosettes lie loosely on the sand and
The Peruvian-Chilean Desert
'000
lOOO
...
. .. . _. - -. .. .. ..
.- .
265
1 ___ - .... 1 ${JMwIJ'1
It"'y-....v I AJnff
lLi!j FS/IIp.l dtgf'imr"'4S
Sm.nl'p.nIS<t,so(on
"110 IMl'tbnJlollO
[W btep'''rlJUSt/1-'4

I'*" tt I

1
1
;, ,an herl14/ 1ft ...
rn Comvrml,rt,s Uf"t(4'y
d, eler'",s f:D/uml'lltAt
Fig.3.40. Cross-section through the Andes at the latitude of Lima from the coast to the high Andean
region: 1 sea- and freshwater; 2 deserts; 3 guano fields on islands; 4 cover of garua fog; 5 lomas (fog-
green slopes) of the coastal zone; 6 stands of rootless Tillandsias in the desert; 7 riverine forest;
8 glaciers and snowfields; 9 subalpine cushion plant communities; 10 puna grassland; 11 grass steppe
with scattered shrubs; 12 rocky slopes with bromeliads; 13 open, evergreen forest; 14 scrub steppe;
15 valley slopes with summer-green herbaceous communities; 16 stands of Carica and Jatropha;
17 stands of column cacti (enumeration from 1eft to right and vertically, from top to bottam) (after
M. Koepcke, in Troll 1959)
Fig.3.41. Successive rows of Tillandsia 1atifo1ia,
T. purpurea and T. straminea at the southern foot
of the Morro Solar, south of Lima. Sea in the
background (photo H. Ellenberg)
appear to be without raots (Fig.3.41), be-
cause they can be easily lifted. Rauh (1985)
has shown in transverse sections of the axes
that the raots run downwards within the
outer cortex of the shoot and, sheathed by
the leaf bases, seldom appear on the out-
side. They are thus functionless. According
to Rauh the species which occur here are Til-
landsia recurvata, T.latifolia, T. purpurea
and T. werdermanii. Of these, only the latter
is limited to this fog area, the others being
distributed far into the mountains as epi-
phytes and epiliths.
These Tillandsias are the only known true
homoiohydrous fog plants, that can survive
without water uptake through raots. They
obtain sufficient water from fog or dew that
condenses on the leaves, absorbing it through
sucking scales which function like a valve
and are typical of the Bromeliaceae.
Many experiments have been conducted, using
labelled water, to estimate the uptake of water
through leaves or their hairs or even the thorns of
cacti. In most cases the quantities involved are so
small that they are of no ecological significance.
The process is simply a physical one which will
occur wherever there is a sufficient gradient in
water potential.
The water economy of these true fog plants has
been examined in detail by P. de T.Alvim and
Andres Uzeda (personal communication). The
species investigated was Tillandsia pa1eacea. The
potential osmotic pressure of the leaves, deter-
mined by the cryoscopic method, was 7-8 bar.
When the leaves were suspended for 24 h over
solutions in a closed chamber in an atmosphere of
80-100% relative humidity, only a relatively
small increase in weight was detected. This may
have been caused in part by condensation of
266 Zonobiome III: Subtropical Deserts (The Arid Zonobiome)
Tlilandsla straminea
JiXIIJ
8 .'11 .7
July 1958
Fig. 3.42. Water balance of Tillandsias in the Peru-
vian coastal desert during July 1958. A Weight at
09.00h, after dew fall: B weight at 17.00h (after
Alvim and Uze da, personal communication). For
further explanation, see text
water. It follows that the Tillandsias can only
absorb water through their leaves when they are
weUed directly.
Water gained and lost can easily be determined
in the natural habitat. Plants weighing about 3 kg
were placed on a plastic sheet and left to grow in
situ. Successive weighings were done at 09.00 h,
when the dew had already disappeared, and at
17.00 h, after the plant had transpired the whole
day. The investigation was made in July under
the most favourable conditions. Fog and dew are
particularly frequent at this cool season of the
year. The mean temperature was 14C and the
mean vapour pressure was 12 mmHg. The results
are shown in Fig.3.42. The increase in weight
during the night and its loss during the day can be
clearly recognized.
Calculation of the mean fresh weight increase
during July gave a growth rate of 0.17'Yo day'.
Thus the plants grow at an extremely slow rate
even in this favourable season. This investigation
shows that the Tillandsias survive entirely on the
water which they absorb from condensation of fog
or dew with their absorbing scales and they are
thus able to live in a practically rainless fog desert.
The Tillandsias reproduce mainly vegeta-
tively, from parts that are broken off and
scattered by the wind. Their arrangement in
rows, at right angles to the direction of the
wind, suggests that this orientalion is espe-
cially favourable for condensation of fog or
dew, which occurs here at night and in very
small quantities only. In some the growing
tips are oriented to the sea, from which the
fog comes, while the parts of the shoot that
are directed inland die and turn black. They
have colonized a habitat on which other
plants, even cryptogams, cannot grow.
Tillandsias are also found in another
biotope in this fog desert, behind the coastal
cordilleros, in finger-shaped stretches where
the fog penetrates the mountain barrier: this
is indicated in Figs. 3.39 and 3.40.
As can be seen in Fig.3.39, the crypto-
gamic zone begins a few kilometers inland,
where the land begins to rise and condensing
fog wets the ground more heavily. Accord-
ing to Ellenberg (1959), thick, jellylike masses
of blue-green algae, Nostoc cummune and
others, are found here. They cover at most
one-third of the surface. Unlike the Tilland-
sias, the algae cannot stand being covered
with blown sand. South of Lima, on stable
sand, there are large colonies of lichens
(species of Cladonia). while colourful crus-
tose lichens are found on stones. The effect
of fog and the distribution of the vegetation
are closely related to the relief of the terrain.
Rauh has pointed out that amongst the Til-
landsias there may be smaller or larger
stands of cacti; these are low creeping forms
which are spherical or have short columns.
According to Ellenberg (1981). these "cactus
lomas" occupy special, localized biotopes,
where at night dry winds blow from the
mountains and disperse the garua fog. The
tender loma herbs cannot survive this
periodic drying out, but it is actually favour-
able to cacti for it prevents cactus seedlings
from being infected with fungi and also
eliminates competition from the herbs (Fig.
3.43).
South of Trujillo, in the more humid north,
groups of Haageocereus repens occur; its 2-
m-Iong shoots are flat, have roots on the
lower surface and are partially covered with
sand: only the erect tips emerge and point
seawards. The water economy of these cacti
has not yet been investigated. It can, how-
ever, be assumed that they have diurnal
acid metabolism, so that the stomata are
open only during the humid nights, and that
water uptake is through the roots. Near
Chala in southern Peru species of Islaya
form their own community ne ar to the coast,
in an area which otherwise includes only
cyanophytes. In northern Chile the Copiapoa
cinerea cactus community is characterized
The Peruvian-Chilean Desert
267
NE
SW
LOMAS OF MOllENDO
.. MOLLENDO .........---... __ -_
__ _
rare /t'J dry land breeze
800 . ______ - - - .- -- - . : 'opE,n lOg (tlerbs) . : . ' .. : . ' " . (at night)
600 -- -- - .. /:E2>
GaruafrequentweHlng :. :.:::':.<: .:;: .-, : ... . :.; .. _: .. Infrequent wethng logs;
.,' . . . ..... <ifo-'"' ...... -
400 _________ . -.':"': : :: ':-: -:-- -. -- a" olten dry. espec,ally at n,ght
less caclus _ Ihornbush seml-desert
200
xeromorph,C. herbaceou$ log vegetatIOn
o
Pacilic
Fig.3.43. Schematic profile through the lomas of Mollendo, which lie on a slope down to the coast from
a desertlike, dry high plateau. Succulents occur here only where the fog-herb area is not completely
closed, namely, on the floor of the valley through which dry air blows at night (after Ellenberg 1981)
L..L..L.J
A
o 2 .. 6' c.on
Fig.3.44. The root system of the earth cactus,
Neochilenia napina, with beetlike storage root,
and very shallow, but extensive lateral roots. A
Lateral view; B from above (after Weisser 1967)
by the presence of migrating lichens; these
are not anchored to the ground and are rep-
resented in this desert by Rocella cervi-
cornis, Parmelia vagans and Tornabenia
epheba (Follmann 1966) . The "earth cacti"
are a very unusual group. They occur further
south in the region between Antofagasta
and Coquimbo in northern Chile. Since they
are sunken entirely into the soil they are
hard to find and were first discovered by
Weisser (1967). At a depth of 1.5 cm, fine
lateral roots emerge from the thick root
tuber and can take up condensed fog (Fig.
3.44).
Somewhat higher up the slope the true
loma vegetation starts (loma = hill in Span-
ish). The rising moist air cools and there
is an increase in fog droplets. The droplet
content of 1 m
2
air varies greatly (0.01-5 g)
but is usually 0.4-0.8 g. As soon as the fog,
which is driven by the wind, meets with a
fixed structure in its path, it condenses. The
larger the surface of the obstacle, the greater
the quantity of water precipitated. Thus a
rain gauge will catch only a little water. On
the soil surface, condensation is slight; it is
greater on vertical rock faces and very great
on tall plants, while trees "cornb" the water
out of fog that is blown through their
crowns, resulting in a constant dripping
from leaves and twigs. Wind velocity is im-
portant in this regard. If, for example, the
branch system of a tree combs out 0.5 g
water from 1 m
3
passing air, this would re-
sult, with a wind speed of 1 m S-1 (= 3.6km
h -1). in the condensation of 1.81 h - 1; this is
the equivalent of precipitation of 2 mm h -1.
Although the fog lasts for many hours a day
on about 120 winter days in the loma area,
rain gauges set up in the open measured
only 121-219mm (mean 168mm) annual
"rainfall" for the period 1944-1954. Meas-
urements made beneath plantations of
casuarines showed, however, a precipitation
of 262-819 mm (mean 488 mm) and beneath
eucalyptus 252-1240mm (mean 676mm).
Ellenberg (1959) made similar measure-
ments in the lomas of Lachay, near Lima. His
results are shown in Figs.3.45 and 3.46. It
can be seen that during the dry year of 1957,
bare soil was wetted to a depth of about
10 cm, but beneath herbaceous plants to a
depth of 50 cm and under eucalyptus trees
to more than 100 cm. Thus plantations of
casuarines and eucalyptus create for them-
selves the soil humidity they require. On
foggy days, potential evaporation and tran-
spiration are hardly detectable. On moder-
268 Zonobiome III: Subtropical Deserts (The Arid Zonobiome)
""
I." ...
.....,
.......
Mt
"""
s..,
,
,
so
,
"

"
ISO
Fig.3.45. Soil moisture conte nt (%) beneath an
open Eucalyptus stand in the lomas of Lachay in
1957. In this year the fog season began only in
June (after Ellenberg 1959)

, So'.-\gs,. aV8I'age
CUlt1eme"
Oryytar1e57
59
.. rnrn
NO veQe!lIIhon
... ... .. _-
_'\. _ _ _ r a
LJm.lcl
RooI.
Fig.3.46. Quantity of precipitation and soil mois-
ture in sandy soils beneath different vegetation in
the lomas of Lachay (after Ellenberg 1959)
ately cloudy days evaporation, measured
with Piche evaporimeters, reached a maxi-
mum of 0.45 ml h -\ only on dry days, with
an air temperature of 28C, did it exceed
0.7ml h-
1
.
The natural loma vegetation on the lower
slopes consists entirely of annuals. These are
various indigenous Nolana spp. (Nolanaceae,
aff. Solanaceae, Convolvulaceae). At first
there are no grasses, but they become more
frequent with greater soil humidity higher
up the slope; the herbs still dominate, how-
ever, forming a thick carpet, 60 cm high.
This is comprised of the tender-Ieaved Nico-
tiana paniculata, Galinsoga, Palaua malvi-
folia (Malvaceae). Nama dichotoma (Hydro-
phyllaceae). Drymaria weberbaueri and
Spergularia collina (Caryophyllaceae). Bow-
lesia palmata (Apiaceae). Plantago limensis,
Loasa urens, which is covered with stinging
hairs, and many others. Plants regarded as
weeds in Europe are also found here, includ-
ing Sonchus oleraceus, Stachys arvensis,
Erodium cicutarium, Stellaria media. None
oi these species can be regarded as fog
plants, for water uptake is mainly through
the raots from the well-moistened soil. A
possible small uptake of water through the
leaves is of no ecological significance.
Apart from the annuals, a number of geo-
phytes (Iridaceae, Solanum, Oxalis and
others) are also characteristic, as are the
large-Ieaved narcissus Hymenocallis aman-
coes, the Amaryllidaceae Zeyphyranthes
and Stenomesson coccineum; the latter
developed only leaves in the foggy winter,
while the orange-red flowers develop from
the bulb only in summer. Semi-succulent
species of Calandrinia (Portulacaceae) occur
in large numbers in some places.
Woody plants are found in places where
the fog is thickest, at 450-600 m NN; they
are usually isolated specimens, however,
growing on rocks; they include Acacia
macracantha, Carica candicans, Capparis
ptisca, Caesalpinia tinctoria and the poison-
ous Croton shrubs. Near Atiquita which is
close to Chala (Fig. 3.32) there a small patch of
forest with Eugenia ierreyrai, which has hard,
evergreen leaves and a thick crown canopy
above 5-8-m-high trunks. The branches of
these woody plants are thickly covered with
liverworts and mosses, that hang down in
long beards. Even the small semi-succulent
Peperomia crystallina occurs as an epiphyte,
and so do species of Polypodium. In the
forest shade there are several hygromorphic
ferns.
Ellenberg (1959) believes that such forests
were at one time widespread, but have since
been destroyed by man; the reason for this
view is that during the main period of
growth of the loma vegetation at 300-500 m
NN, Indians drive their cattle out of the dry
mountains into the loma region and use the
trees for firewood. Rauh (1985) argues, how-
ever, that the loma trees nowhere form a
closed forest belt, occurring always as is-
lands in places where much fog collects,
separated by wide expanses of desert. Man
has, however, probably caused a certain
amount of degradation.
The Peruvian-Chilean Desert
In northern Chile the coastal cordillera re-
treat from the coastline, so that there are
wide sand flats between the mountains and
the sea, and no fog is formed. A loma vege-
tation is found here only far from the coast,
at 1000-1200m NN.
The last spurs of fog vegetation, consisting
of colourful crustose lichens, are found on
stone heaps at 800-1000m NN. Higher up,
often to 2000 m, the mountain slopes are al-
most completely barren.
4 Loma Fauna
There is, of course, a characteristic fauna as-
sociated with the loma vegetation in Peru
(Koepke 1961). Flies swarm around the
flowers, carabid bettles are very numerous;
the spider Lactrodectus and the scorpion
Hadruroides lunutas also occur here, the
latter being found on the Galapagos Islands
as weH; woodlice further the process of soil
formation with their excrement. Several
varieties of Bulinus snails are found on the
separate loma islands. This is true also of the
bird, Geositta peruviana, which seeks shelter
in holes in the sand. There are also lizards
and a smaH fox, a species of Dusicyon.
5 Oases
The valleys of the 41 rivers of the Andes are
densely populated oases. In the Peruvian
part of the desert they penetrate through the
coastal cordillera and flow into the sea. Salix
humboldtiana and the pepper tree, Schinus
molle, grow on the banks of the rivers and
along their edges the tall grass, Gynerium
sagittatum. On the broad terraces crops are
planted and irrigated; besides grain crops
and vegetables, cotton, sugar cane and
lucern are planted. In southern Peru there
are olives, in central Peru citrus fruits and
higher up the valley, near the rock desert,
apples, pears and plums. Viticulture is
mainly concerned with the production of
spirits.
Along the coast north of Lima as far as
Chidayo (see Fig.3.37) there are large irri-
gated areas with cotton and sugar cane
269
plantations; these stretch beyond Trujillo.
The Tinajones project has made it possible
to lead large quantities of water into the
desert and extend the areas under irrigation.
The North-Chilean Desert
Conditions are far more extreme in the part
of the desert which lies in northern Chile,
because there is no coastal fog here and rain
falls on very rare occasions in the winter.
The region is almost completely barren of
flowering plants. Between Iquique and
Arica there is, at the most, a grey-green
tinge which doser examination shows to be
fructose lichens Anaptychia leucomelaena
and Ramalina cerruchis and air alga Trente-
pohlia polycarpa. Follmann (1965) has de-
scribed window lichens from the Atacama
Desert, between 22 Sand 27 S, members of
the families Buelliaceae, Dermatocarpaceae,
Lecideaceae and Acarosporaceae. Their
only source of water is the nightly dew; they
lie buried in the sand, only the disc-shaped
fruiting bodies projecting just above the
grains of sand. It is possible to cultivate
some blue-green algae from sand taken
near Caldera and Antofagasta - Schizothrix
atacamensis, Calothrix desertica. There are
also two forms of Cyanidium which are
endolithic and occur in cracks in diorite.
This is also an area where the migratory
lichens and earth cacti are found (p. 267).
There is no loma vegetation in this region.
If rain falls in the winter, the desert becomes
carpeted with flowers for abrief period.
Kohler (1967) described such an event in the
north-Chilean Atacama Desert in 1965. By the
18th of July, 22 mm rain had fallen. Since temper-
atures are relatively low in the winter months,
growth was slow. The main flowering of Nolana
baccata and Hippeastrum thus took place only at
the beginning of September, that of Calandrinia
and Cristaria at the beginning of October; the
seeds were ripe by the end of October. Amongst
the ephemerals, a distinction must be made be-
tween annuals and geophytes (ephemeroids). The
former survive the rain-free period as seeds, the
latter as tubers or bulbs underground. The geo-
phytes appear to require more water far their
development, for where the relief is undulating,
only annuals are found on the elevations; here
they have a cover of 10-40'X" and their roots
penetrate to a depth of 10-20 cm. Geophytes are
found only in the hollows, where they occur to-
gether with a dwarf shrub, Skytanthus acutus
270 Zonobiome III: Subtropical Deserts (The Arid Zonobiome)
(Apocyn.). Besides species of Nolana there are
several other annuals, a yellow Viola, for exam-
pIe, and even Cuscuta, which parasitizes these
ephemerals. Calandrinia, which has succulent
leaves, is able to prolong its growth period by
utilizing the stored water.
On the Pampa deI Tamarugal of the Salar
de Pintados, south-east of Iquique, there are
large stands of Prosopis tamarugo, a relative
of P. julif1ora, the mesquite of the Sonoran
Desert. In this extreme desert these trees
seem especially striking.
Went (1975) has reported the astonishing inves-
tigations of F. Sudzuki, which purported to show
that, at night, the leaves of Prosopis bushes take
up large quantities of water from the vapour-
saturated air and conduct this to the roots; these
excrete the water, so that the soil around the roots
becomes thoroughly wetted to a depth of about
1 m, while the soil above and below remains com-
pletely dry. This reserve water, it was suggested,
is then available to the transpiring plant during
the day.
This remarkable idea, untenable on physical
grounds alone, has been shown to be mistaken by
Mooney et al. (1980).
Chile an hydrologists have found that
beneath the Pampa deI Tamarugal there is a
groundwater lake, the level of which had
varied and had dropped in the years prior to
the investigation, owing to the demands
made on it by Prosopis. Formerly there was a
lake here which, owing to the high evapora-
tion in the desert climate, became covered
with a crust of sodium salts which have been
exploited as a source of salpeter.
The underground water lake is fed by
groundwater that flows from the tops of the
mountains through the detritus which lies
above the bedrock on the slopes; at the low-
est point of the Pampa deI Tamarugal the
lake rises almost to the soil surface (Fig.
3.47). As a typical phreatophyte, the mes-
quite is limited to places where there is
groundwater and its tap root can grow to
considerable depths to reach this water.
There can be no doubt, but that the water
requirements of Prosopis tamarugo are met
by the groundwater lake. The leaves of the
plant show absolutely no anatomical fea-
tures that would allow uptake of large quan-
tities of water. It is, however, not easy to ex-
plain the thick mass of roots at 1 m depth or
the accumulation of water in the soil with an
osmotic potential of -15 bar. The mean
Fig. 3.47. Profile through the Pampa deI Tamarugal
(after Klohn, modified from Mooney et al. 1980)

Hour
Fig.3.48. Curves of air temperature (0C), vapour
pressure deficit (VPD in mbar) and water poten-
tial (- bar) of Prosopis tamarugo near Canchones
(Chile) on 5-6, Oct. 1978 (after Mooney et al.
1980)
osmotic pressure of the cell sap of the leaves
was -21.3 bar. In the Sonoran Desert we
measured values of - 20 bar in Prosopis juli-
flora var. velutina, and during the dry sum-
mer period of -30 to -34 bar; these are thus
closely similar to the values for P. tamarugo.
The nightly change in direction of the vascular
flow of water from the shoot to the roots observed
by Sudzuki may indeed occur temporarily during
the spring, when, at night, the water potential of
the leaves rises to -10 bar and that in the roots is
-15 bar (Fig.3.48). Normally, however, the flow
of water is from the roots to the leaves.
Mooney et al. (1980) suggest that the
wetting of the soil in the region of the roots
The Peruvian-Chilean Desert
might arise when, due to the lower transpi-
ration at night, some of the water absorbed
from greater depths by the tap root is ex-
creted by the root felt, and acts as areserve
when transpiration increases during the
day. This is, however, entirely speculative. It
seems to us that the root felt, at 1-1.5 m
depth, might have been formed at a time
when the groundwater was at this level, and
that recent endeavors at reforestation have
led to its rapid fall. The P. tamarugo stands
grow in the lowest part of the terrain, where
the groundwater level is only a little below
the surface.
The question arises as to how the tap root
of a Prosopis seedling reaches the ground-
water if the soil above is dry. This takes
place, as in all phreatophytes, only when
there is an exceptionally rainy year; the soil
is then wet from the surface to the ground-
water and the tap root grows rapidly to the
depth of the groundwater before the soil
dries out.
A rainy year of this sort occurred in this
area at the turn of the century and again in
1977, when the water masses from the
mountains flooded the whole of the Pampa
deI Tamarugo, washing away the salt so that
the salt-sensitive seedlings could take root.
The old P. tamarugo trees should thus be
about 75 years of age, but this has not been
checked. In many extreme habitats the
stands consist of a few generations only.
Detailed investigations in the desert a1-
ways revea1 a simple explanation for obser-
vations which those not acquainted with the
special situation at first find inexplicable,
and are thus often led to unjustifiable con-
c1usions.
In reforestation with P. tamarugo today, seedlings
are planted in a 40-cm-deep hole which goes be-
neath the salt crust; thorough irrigation then en-
sures that the soil is wetted as far down as the
groundwater. The tap roots of these phreato-
phytes grow rapidly in length, without producing
lateral roots, and form a dense felt of lateral roots
only when they have reached the capillary zone
above the groundwater. Too dense afforestation
can result in a fall in the groundwater level, for
the water utilization by phreatophytes in the de-
sert is extremely high.
The extreme desert or "Atacama" extends
in northern Chile over a vast area and is one
of the most desolate and barren areas on
271
earth. Rauh described a transit from Toco-
pilla in the west to Calama in the east as fol-
lows: "Only along small erosion gulleys does
one find isolated specimens of Adesmia ata-
camensis (Leguminosae), Coldenia ataca-
mensis (Ehretiaceae, afL Boraginaceae) and
Cristaria divaricata (Combretaceae). The
lowest hill range of the Sierra Domeyko has
a slightly denser cover of thorn bushes. Be-
hind these hills, in the valley of San Pedro de
Atacama, are oases with irrigated planta-
tions. Here there are the leguminous trees,
Prosopis juliflora and the "Chanar" Gourliea
decorticans. South of this is the 2270 km
2
Salar de Atacama and other salt lakes which
are covered with thick salt crusts. On their
banks are open stands of Atriplex atacamen-
sis, Ephedra andina, Lippia trifida, Teneria
absinthioides and Distichlis meadows,
strewn with the small chenopodiacean Nitro-
phila axillaris".
In Peru, however, the extreme desert is
only a small area of the lower land at the foot
of the western slope of the Andes, where it
occurs as rock or stone desert. The upper
surface of the stone heats up to 70C in the
sun and the humidity of the air is extremely
low. Isolated bushes are found only in places
where, after exceptionally heavy rain, water
flows off the slopes and is stored in the soil.
6 Orobiome III
on the Western Slope of the Andes
As altitude increases on the slopes of the
Andes above the extreme desert, the cactus
desert begins. The floristic composition of
the stands of cactus changes markedly from
north to south (Rauh 1958). It starts in north-
ern Peru with the largest column cactus of
Peru, the 8-10-m-high Neoraimondia gigan-
tea; this cactus is distributed along the
whole length of the almost rain-free western
slopes of the Peruvian Andes. It has a shal-
low, horizontal and very branched root sys-
tem. Growth in height is very slow: in cen-
tral Peru it was estimated to be 20 cm in 50
years.
The most extreme habitats are occupied
by the more spherical genus Melocactus
(see p. 190 and Fig.2.91 on p. 212). Its deli-
272 Zonobiome III: Subtropical Deserts (The Arid Zonobiome)
cate root system penetrates into the finest
cracks in the rocks and is said to absorb
dew, but no detailed investigation has been
made. These cacti can endure high tempera-
tures of about 45C in the central water tis-
sue; they shrivel up completely when it is
dry, refilling their water reservoir immediate-
ly should rain happen to fall. Still higher up
the slope other column cacti replace those
already named; amongst them is the climbing
species Hylocereus venezulensis. Higher
up, above the cactus desert, there may be an
open deciduous forest of Bombax discolor,
the crowns of which are thick with epiphytes
- Tillandsia spp., Vriesea cereicola, Guz-
mania monostachya and others - an indica-
tion that dew probably falls frequently.
The detailed floristic composition of the
cactus desert is, however, very varied, as is
that of the other altitudinal belts. Rauh
(1985) has described this in the Valley of
Chillon, north of Lima. Between 400 and
700 m NN the rock desert is without vegeta-
tion. Above this, from 700 m to 1700 m NN is
the cactus desert. The first species to occur is
Armatocereus procerus. On the basis of
floristic composition, three different cactus
altitudinal belts can be distinguished, al-
though they are not sharply demarcated
from one another. On the upper part of the
slope are also succulent shrubs such as
Jatropha macrantha and Carica candicans,
succulent species of Peperomia and the suc-
culent Pilea serphyllacea (Urticaceae); there
are also poikilohydric ferns - species of
Cheilanthes and Pellaea, and, as well,
Selaginella peruviana. The higher slopes,
from 700 to 2400m NN, are especially dry.
Still higher up on these western slopes of the
Andes, large banks of cloud form from rising
air masses so that rain falls here, the quan-
tity depending on the latitude.
The limits of the altitudinal belts differ
greatly, depending on exposure and on the
orientation of the valleys.
For central Peru the following altitudinal belts
can be distinguished:
300 m Weak effect of fog: barren sand
desert or Cyanophyta lichen com-
munity or non-rooting tillandsia
stands or coastal cacti
700 m Strong effect of fog: loma forma-
tion of annuals and geophytes or
woody plants or cactus stands. At
the upper limit, the effect of fog
is again weak and in places non-
rooting Tillandsias occur
Lower dry belt: only large col-
umn cacti without shrublike
companion flora
m Upper dry belt: low column cacti
with succulent shrubs and bushes
m Summer rain belt: column cacti
with raingreen woody plants and
annuals, and shrubs as com-
panion flora
m Cold summer-rain belt: there are
no cacti, apart from two opun-
tias; evergreen shrubs; grasses
and shrubs increase with altitude
Summer-green grass puna:
above 4500 m only a few detritus
and rock plants
Rauh lists the following fm nmthern Peru in the
Valley of Rio Sana (7 S). that is, within the
reaches of zonoecotone 1II/1I
100m
500m
900m




Semi-desert with scattered woody
groups
Cactus belt
Dry forest with cacti
Dry forest hung with Tillandsia
usneoides
Evergreen montane forest with
Ficus and palms
High montane, evergreen forest
with Podocarpus and Drymis
Ericaceae-Melastomaceae shrub
belt
above 3200 m Paramos, here known as Jalla
This altitudinal belt series is very closely similar
to that in northern Venezuela (Vol. I, p. 206). The
first two altitudinal belts are the equivalent of the
Espinar-Cardonal, the next two of the Selva
decidua belts. Mean annual rainfall measured
over 4 years at 1750 m NN in these northern-
Peruvian altitudinal belts was 1370 mm.
We will not describe here the vegetation
of the deeply incised, often desertlike inner-
Andean valleys. They do not belang to the
Peruvian-Chilean desert, but form part of
the huge meridionally oriented mountain
range of the Andes (Fig. 2.52). Their vegeta-
tion is greatly influenced by the strong di-
urnal winds.
7 Zonoecotone III/ll
Zonoecotone IIIIIV between the Chile an
Desert and the winter rain area of central
The Peruvian-Chilean Desert
Chile will be dealt with in Volume 4, in the
section on zonobiome IV. Here we will con-
sider only zonoecotone III/H, the most north-
erly coastal region of Peru and the most
southerly part of Ecuador. There are no
mangroves along the reach of the Peruvian
coast which is under the influence of the
cold Humboldt current. The most southerly
habitat of this sort on Peruvian soil is the
delta of the Rio Tumbles, on the Gulf of
Guayaquil, a little north of 4 S. Here all the
typical mangrove species are to be found,
such as Rhizophora mangle, Avicennia
tomentosa, Laguncularia racemosa and
Conocarpus erecta. Further south the tidal
zone consists only of a salt marsh, with species
such as Sesuvium portulacastrum, Salicomia
fruticosa, Batis maritima and the halophilic
grasses, Distichlis, Sporobolus.
North of 8
0
S light summer rain falls and the
desert becomes a rain-green grassland with
mainly annual grasses. These include Aristida
adscensionensis, Anthephora hermaphro-
ditica, Bouteloua disticha, Chloris virgata
and Eragrostis spp.
In the Peruvian part of the desert, north of
6 S, there are no long-lasting winter fogs.
Instead, a heavier summer rain falls about
every 5-12 years, following which the sandy
soil becomes covered for a few weeks with
an ephemeral vegetation. Elsewhere, the
plant cover consists of a mosaic of barren
sand dunes, open shrub and tree savanna
with a parkland-like distribution. On the
western slopes of the coastal cordillera on
rocky biotopes is the huge column cactus,
Neoraimondia gigantea; in addition, there
are sm all tree species such as Prosopis juli-
flora, the .. algarrobo" with hard wood and
husks which are rich source of nutrients for
cattle, the evergreen Capparis angulata, the
green-barked Cercidium praecox (Sapo-
tacaea) and Parkinsonia aculeata. There are
also more shrublike forms, Capparis cordata
and C. ovalifolia, Grabowskia boerhavii-
folia (Solanaceae). Galvesia limensis (Scro-
phulariaceae). Monina pterocarpa (Poly-
galaceae). Scypharia spicata (Rhamnaceae).
and others.
273
A striking annual is Luffa operculata (Cu-
curbitaceae) which forms a dense thicket
after good rain.
In the Ecuadorian part of the region there
remains only a small, desertlike area with a
sparse bush vegetation at Cape Puntavilla
near Salinas on the Santa Elena peninsula.
Elsewhere rain-green shrub changes to
savanna and rain-green forests, in which
Ceiba (Bombacaceae) with its mighty barrel
trunk and buttress roots is especially strik-
ing. Cacti are represented by species of sev-
eral genera. In the undergrowth Ipomoea
carnea is often dominant; this is, of course,
the zonal vegetation of zonobiome H.
Galapagos Islands
Before leaving the Peruvian-Chilean desert
areas, we wish to deal briefly with the
Galapagos Islands which, although they lie
on the equator and are 1100 km from the
mainland, are also washed by the cold Hum-
boldt current. As a result, the coastal region
is arid. Annual rainfall is only 100 mm,
falling in the months January to April. Thus,
at low altitudes there is a typical Espinar-
Cardonale vegetation, as on the arid north
coast of Venezuela (Vol. I, p. 206).
The islands are of volcanic origin and rise
to 1000 m. Everywhere above 200 m NN is
shrouded in low stratus clouds even during
the dry season, providing moisture by con-
densation. The flora consists of 463 species
of flowering plants, of which 204 are endemie.
At high altitudes the islands are forested
with woody species such as Psidium galapa-
gicum, Scalesia pedunculata, Pisonia flori-
bunda, Xantoxylum fagara, Piscidia erythrina,
bushes of Miconia robinsoniana with pterido-
phytes and moss undergrowth.
Charles Darwin visited these islands in 1835:
here the large number of finch species later pro-
vided some of the evidence for his theory of evolu-
tion. The large Galapagos tortoises are also well-
known; they have had to be protected from exter-
mination by man. Although the islands are under
nature proteetion, the many tourists are a great
threat.