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0007-2745/78/277-293$1.95/0
278
THE BRYOLOGIST
[Volume
81
1000
800
no vegetation
600woody
plants
400
growth
ofopen
perennials
0o
cryptograms
S200-
i~n
no vegetation
vegetation-
and associated areas of extensive upwelling of deep ocean waters off the Atacama
Coast produce a mild, uniform climate in the coastal zone and such stable air mass
conditions that precipitation is rare or even absent. In the coastal Chilean Atacama,
mean annual precipitation ranges from a high of 26 mm at Caldera to an incredible
0.6 mm at Arica over a 44 year record. In central Peru precipitation increases slightly,
reaching 41 mm yr-1 at Lima. Available climatological data for the Atacama desert has
been detailed by Hajek and di Castri (1975) and di Castri and Hajek (1976). Less data
are known for coastal Peru, but available records are summarized by Meigs (1966). A
summary of climatic data for selected coastal stations is shown in Table 1.
Although precipitation is slight, water input from moisture condensation from
coastal fogs allows local development of rich biological communities. Thick low fogs,
termed the garua in Peru and the camanchaca in Chile, form along the coast almost
daily during the winter season from June to October. Although the elevational limits
of the garua and camanchaca vary somewhat with latitude and local topography, fog
is typically present in a well-defined band extending from 300-800 m (Rundel &
Mahu, 1976). Where the coastal topography is low and relatively flat, the effects of
the fog are dissipated over broad areas with little resultant biological influence. Where
steep coastal mountains are present, however, fog condensation is concentrated in a
narrow belt where moisture input allows the development of relatively lush fog zone
vegetation. Conditions for condensation of fog moisture are particularly favorable
along the Atacama coast (Ellenberg, 1959). The temperature of the Humboldt Current
is warmer than that of similar currents off the coasts of Baja California and Namibia,
producing high water vapor contents and consequently high aerosol liquid water content of air masses with subsequent cooling. Also important is the constant light southwest winds, blowing onshore at velocities up to 4 m/s. This wind is important in
providing a continuing source of new aerosol water for condensation against the steep
coastal mountains.
Vegetation development in the coastal lomas of Peru has been described by many
(e.g. Troll, 1932; Weberbauer, 1945; Ellenberg, 1959; Koepcke, 1961; Walter, 1971).
A diagrammatic representation of a typical coastal fog zone is shown in Figure 1. The
coastal plain below the fog zone is barren of vegetation, except for scattered stands
of unrooted terrestrial Tillandsia (Bromeliaceae) which occurs on windblown sand
1978]
279
TABLE1. Climatic data for coastal stations in the AtacamaDesert of Peru and Chile, Sonoran
Desert on the Pacific Coast of Baja Californiaand Nanib Desert of Angola, Namibia and South
Africa. Data from Meigs (1966), di Castri and Hajek (1976) and Hastings and Humphrey (1969).
Mean
Mean
Mean
Trujillo
Lima
Mollendo
Atacama-Chile
Arica
Iquique
Los Condores
Cerro Moreno
Taltal
Chafiaral
Caldera
La Serena
Sonoran-BajaCalifornia
Isla Todos Santos
San Telmo
El Socorro
El Rosario
Vizcaino
Bahia Tortugas
Punta Abreojos
San Juanico
Bahia Magdalena
La Aguja
Todos Santos
Namib
Luanda
Lobito
Mogamedes
Walvis Bay
Port Nolloth
Klaver
6037'
22.2
6041'
22.9
15.2
8
3-4
19
21.1
27.9
2-4
49
18.9
23.9
16.1
45.7
26
17058'
17.8
19.4
16.1
22.1
13
18028'
20015'
23029'
25025'
26020'
27003'
29
515
518
119
39
9
28
18.7
17.9
15.5
17.0
17.4
16.4
16.1
22.2
21.2
19.0
20.1
22.0
19.3
19.7
15.1
14.1
10.2
13.3
14.5
12.2
12.9
0.6
2.1
0.0
2.2
25.1
1.7
25.8
44
49
7
7
21
7
49
29054'
32
14.8
18.9
11.2
127.4
91
8005'
12000'
20012'
31048'
22
255.6
30058'
30020'
30004'
27059'
27042'
100
10
15
10
16.3
16.9
20.9
18.8
172.1
137.7
95.0
79.8
17
11
14
10
26044'
5
15
20.3
21.6
95.8
76.8
11
12
26016'
12
21.1
64.1
24038'
12
21.4
73.7
31
23059'
10
22.4
49.9
23026'
18
22.0
169.6
30
8050'
12013'
15010'
29010'
60
1
3
7
7
24.4
24.4
21.1
17.2
13.9
27.2
27.2
25.0
22.2
18.3
21.7
21.1
16.7
11.7
9.4
322.6
353.1
53.3
22.9
58.4
59
19
21
20
64
31043'
42
19.4
26.7
12.2
22053'
dunes. The lower margin of the fog zone is characterized by cryptogamic communities
of Nostoc and Teloschistes on stable sand surfaces and crustose lichens on rock substrates. Higher up, scattered stands of herbaceous plants and low woody vegetation
merge into a true fog forest of evergreen trees 5-8 m in height at the center of the fog
zone. Epiphytic mosses and lichens are abundant. As the density of the fog zone
decreases at 700 m, open stands of vegetation give way to scattered cacti and tillandsias and, finally, to barren land free of plants. Ellenberg (1959) found that vascular
plant development over the fog moisture gradient was related to the depth of penetration of soil moisture. Virtually no data are available on lichen zonation, however.
Vegetation zonation in the coastal fog belt of northern Chile at Paposo has been
described in detail by Rundel and Mahu (1976), and broadly by Reiche (1911) while
THE BRYOLOGIST
280
[Volume
81
-1000
NO
VEGETATION
.. ..
. ..
....
..
...:.
..
..
CO PIAPOA
..
-800
....
.................
:EULYCHNIA:?.. F-600
..COPIAPOA
..
EUPHORBIA.EULYCHNIA
/
H:::::::::::::
A::::::
EUL
YCH
IA-
PU
-200
SOPEN
COASTAL PLAIN-COPIAPOA
o-10
FIGURE2. Diagrammaticview of vegetation zonation in the coastal fog zone at Paposo,
AtacamaProvince, Chile. From Rundel and Mahu (1976).
the flora of this zone has been treated by Johnston (1929). Here the coastal plain
below the fog belt supports scattered stands of small globular cacti grading to a distinct
belt of terrestrial Puya (Bromeliaceae) at the lower margin of the fog zone (Figure 2).
Within the central fog zone, woody vegetation becomes dominant in forming dense
stands 2-3 m in height with 60% ground cover. At higher elevations in the upper fog
zone, open slopes with increasingly scattered cacti replace woody vegetation (Rundel
& Mahu, 1976), with the area above 1000 m virtually free of any vegetation. While
the development of woody vegetation is not as luxuriant as in the lomas of Peru, the
diversity and biomass of lichen epiphytes are greater. Detailed studies of the lichen
flora of the coastal Chilean zone have been made at Cerro Moreno near Antofagasta
(Follmann, 1960, 1967c), at Paposo (Rundel, unpublished data) and at the desertsclerophyll transition zone of Coquimbo and Aconcagua Provinces (Follmann, 1960;
Follmann & Redon, 1972; Redon, 1972).
Coastal Sonoran Desert of Baja California.-The
coastal Sonoran Desert extends
from El Rosario south along the Baja California peninsula. Coastal fogs and high
humidity, extending as much as 50 km inland, are characteristic of this desert area as
well as semi-arid sclerophyll and thorn forest vegetation transitions on the northwestern coast and southern end of the peninsula. The general vegetation characteristics and vascular plant flora of this desert region were described in detail by Shreve
and Wiggins (1964). Extensive climatological data for Baja California have also been
published (Hastings, 1964; Hastings & Humphrey, 1969). Data for coastal stations are
summarized in Table 1.
Unlike the South American coastal desert region, no distinctive fog zone forms
along the coast of Baja California. While fogs are common along the coast, they lack
the temporal and structural regularity of the garua/camanchaca. Winter fogs are particularly common, occurring down to sea level, and moist marine air close to moisture
saturation blows strongly onshore throughout the year. Contrasting with the gentle
1978]
RUNDEL:
DESERT
281
but steady onshore winds characteristic of the Coastal Atacama, strong irregular winds
blow off of the Pacific Ocean in Baja California, stunting vascular plant vegetation
along the coast and frequently continuing only slightly abated up to 20 km or more
inland. Steep topographic features are rare along the Baja California coast, allowing
maritime influences to reach well across the peninsula in both the central Vizcaino
Region and the Magdalena Plain (Shreve & Wiggins, 1964). Moisture condensation
from humid air masses and irregular fogs allow the development of a moderately
diverse epiphytic lichen community (Nash et al., 1977) and a locally dense growth of
Tillandsia recurvata on a variety of shrubs and succulents.
Although true coastal fog belts do not form along the coast of Baja California,
distinctive vegetation zones dominated by lichens develop where steep cliffs on moderate slopes are topographically developed along the immediate coast (Rundel et al.,
1972). Vascular plant vegetation is commonly restricted severely in these habitats by
soil aridity, wind desiccation and salt spray. Local conditions are optimal, however,
for luxuriant growth of epiphytic, saxicolous and terricolous lichens, and lichen biomass may rival or even surpass that of vascular plants in these coastal areas. Although
published studies of the floristics and ecology of lichens in this coastal zone are few
(Rundel et al., 1972; Nash et al., 1977; Dodge, 1936), the lichen flora is relatively well
known.
Namib Desert.-The
Namib Desert stretches along the southwest coast of Africa,
km
2800
from
Luanda (8?45'S) in Angola to St. Helena Bay (32?45?S) in
extending
South Africa (Meigs, 1966). Like the deserts of Baja California and the coast of Peru
and Chile, the coastal Namib is characterized by cool, moist sea fogs, resulting from
the cold Benguela Current. Observations at Luderitz Bay showed 285 days during the
year with fog or dew (Meigs, 1966). Even when there is no fog, relative humidities
remain at 100 percent during most of the day. Surface condensation of this water is
the major source of moisture for an extensive lichen flora, as well as for several unusual
succulent plants (Walter, 1971; Giess, 1962; Bornman et al., 1973). Annual precipitation is commonly in the range of only 20-50 mm along the coast. Although several
general discussions of the environment and vegetation of the Namib Desert are available (Cannon, 1924; Logan, 1960; Walter, 1936, 1971), detailed descriptions of the
flora are scarce (Giess, 1962, 1968). Vascular plant adaptations have been discussed
by Schulze and Schulze (1976) and Schulze et al. (1976). The lichen flora of this area
has not been described.
Level, gravelly desert plains of the outer Namib support only rare individuals of
vascular plants, but the windward sides of even small pebbles have well-developed
lichen growth. This growth, dominated by Caloplaca, occurs both as upright subfoliose morphotypes which take advantage of direct fog interception and small crustose
growths at soil level which utilize runoff from fog condensation on the rock above. A
variety of strange specialized lichen growth forms from the southern Namib were
described by Vogel (1955). Farther inland, dense colonies of Teloschistes form a distinct stabilizing element on low sand dunes, comparable to the zone of T. peruensis
in southern Peru. The windward slopes of quartzite and marble ridges in nearby areas
provides vertical faces where fog condensation may become concentrated. A variety
of both stem and leaf succulent plants dominate these slopes, together with a rich
saxicolous lichen flora (L. Kappen, pers. comm.).
FLORISTIC
COMPARISONS
Despite the wide geographic separation of the three major regions of desert fog
zones, considerable floristic similarities exist among the lichen floras of these areas.
282
THE BRYOLOGIST
[Volume
81
Chile
Dirina chilensis (Nyl.) Follm.
D. limitata Nyl.
D. lutosa Zahlbr.
Dirinastrum chilense (Dodge) Follm.
Lobodirina cerebriformis (Mont.) Follm.
L. mahuiana Follm.
Fruticose-Foliose
Darbishirella gracillima (Kremph.)Zahlbr.
Dendrographa leucophaea (Tuck.) Darb.
1978]
283
TABLE 3. Occurrence of species of Ramalinaceae in the coastal fog deserts of Baja California and Chile. Data from Rundel et al. (1972), Follmann (1967c), Follmann and Redon
(1972), Redon (1972) and Rundel (unpublished data).
Baja California
Chile
284
THE BRYOLOGIST
[Volume
81
produce large biomasses in the immediate coastal zone where aerosol moisture is
high. Away from the coast, corticolous Ramalina is dominant. In Chile, saxicolous and
terricolous habitats for Ramalinaceae are relatively unimportant although 28% of the
species are typically on such substrates. Large biomasses of saxicolous taxa are totally
absent. Numerous taxa of Ramalinaceae are present in the Namib Desert, but the
majority of species of Ramalina are endemic and remain undescribed. Trichoramalina
melanothrix, with a related species in Baja California, is an unusual endemic (Rundel
& Bowler, 1974).
A third family which forms important elements of desert fog zone lichens is the
Teloschistaceae, most notably the genus Teloschistes. Four species of this genus are
present in the fog deserts of both Baja California and Chile-T. exilis, T. flavicans, T.
chrysophthalmus and T. villosus. The latter is typically restricted to the immediate
coast, while the others occur well inland. As previously described, the endemic T.
peruensis is extremely important ecologically on sand dunes at the lower margin of
the fog zone in Peru (Thomson & Iltis, 1968). The Namib Desert and adjacent parts
of South Africa are likewise characterized by many ecologically important species of
Teloschistes. The unique fruticose Xanthoria flammea is also restricted to this region.
Many other floristic elements are important in all of the desert fog zone regions.
Species of Parmelia with upright growth-forms occur in each region and may dominate
the flora locally as with P. hottentota and similar species on rocky slopes in the Namib.
Chile and Baja California are alike in the frequent occurrence of ciliate species of
Heterodermia.
TRENTEPOHLIA SYMBIOSES
An unusually large percentage of the lichen floras of coastal fog zones of Baja
California and Chile are composed of species with Trentepohlia as an algal symbiont.
Trentepohlioid algae are the symbionts in such ecologically important fog zone genera
as Roccella, Dendrographa, Darbishirella, Dolichocarpus, Hubbsia, Pentagenella, Ingaderia, Reinkella, Dirina, Schizopelte, Opegrapha and Coenogonium, as well as in
a variety of less dominant crustose genera in the Pyrenulaceae, Arthoniaceae, Opegraphaceae and Graphidaceae. Although not well investigated, lichens with trentepohlioid symbionts are also ecologically important in the coastal Namib (L. Kappen,
pers. comm.). Within the macrolichen flora, only Ramalina, Niebla, Usnea and Heterodermia, all with Trebouxia, rival the ecological dominance of the genera with
trentepohlioid symbionts.
There is considerable controversy in the lichenological literature concerning the
biological basis for the establishment of symbioses between spores of lichenized fungi
and Trebouxia. Since Trebouxia is not commonly observed free-living in nature, the
source of the necessary algal cells is unclear, and yet sexual reproduction clearly takes
place with fertile lichen species (Bowler & Rundel, 1975). With Trentepohlia, however, the potential source of symbionts for lichenization can be easily seen in the
field. Free-living colonies of Trentepohlia are common on both rock and plant surfaces
in both the Baja Californian and Chilean coastal deserts.
Little is known of the physiology and ecology of Trentepohlia. The characteristic
orange-red color results from 8-carotene dissolved in fat deposits (Geitler, 1923).
This pigment hypothetically relates to the resistance of Trentepohlia to high light
intensities. In shaded localities the pigment may be almost completely lacking
(Fritsch, 1948). With full pigmentation, Trentepohlia is extremely resistant to high
insolation, much more so than typical green and blue-green algae. Nevertheless, many
1978]
285
RUNDEL:DESERTFOG ZONELICHENS
Interior Desert
Algerian SaharaDesert
(Faurel et al., 1953)
Negev Desert, Israel
(Galun, 1970)
Sonoran Desert
Rock Valley, California
(Nash et al., 1977)
Silverbell, Arizona
(Nash et al., 1977)
MaricopaCo., Arizona
(Nash, 1975)
Chihuahuan Desert
Jornada,New Mexico
(Nash et al., 1977)
Coastal Fog Desert
Baja California, Mexico
(Rundel & Nash, unpublished data)
Cerro Moreno, Atacama
Prov., Chile
(Follmann, 1967b)
Number of
Species
Fruticose
Foliose
Crustose &
squamulose
114
98
44
93
17
12
88
21
19
81
78
28
72
48
40
60
67
54
13
33
146
25
14
60
lichens with Trentepohlia appear to be very tolerant of low light levels, notably spe-
286
THE BRYOLOGIST
[Volume
81
in desert fog zones, however, the structure and morphological characteristics of lichen
floras differ greatly. The most significant environmental factors appear to be moderate
temperatures, high atmospheric moisture content, strong winds and variable solar
insolation. Natural selection should be expected, therefore, to favor adaptations for
optimizing water uptake, reducing desiccation rates and minimizing mechanical damage due to wind.
Contrary to the situation in hot, dry deserts, fruticose lichen growth forms dominate
in coastal fog deserts (Table 4). Crustose lichens in fact, comprise only a small percentage of the total lichen coverage in many areas of unusually high fog frequency in
both Baja California and Chile. It appears, therefore, that the fruticose growth form
may have selective value for optimizing moisture uptake, provided fogs are frequent
and temperatures moderate.
Morphological adaptations to maximize water uptake have been demonstrated in
fog zone lichens. The size and structure of reticulations in Ramalina menziesii Tuck.
(= R. reticulata Kremph.) are related to frequency of fog and relative levels of atmospheric humidity (Rundel, 1974). Morphological variation related to fog occurrence in
Ramalina usnea is present in tropical areas (Rundel, 1978).
Desert lichens may have capabilities of accumulating water rapidly, but only slowly, losing it through desiccation (Zukal, 1896; Galun, 1963; Follmann, 1965). Blum
(1965, 1973), however, rejected the significance of xeromorphic adaptations for protection against evaporation. Experimental studies with arctic and subarctic lichens
have clearly established that morphological characteristics may profoundly affect thallus water relations (Larson & Kershaw, 1976). Thallus surface to volume ratio, branch
shape and degree of clumping of branches all influence the evaporative resistance,
producing significant intraspecific and interspecific population variability. These same
morphological characteristics are clearly important in habitat selection by coastal fog
zone lichen taxa.
Saxicolous and terricolous substrates in coastal fog zones characterized by aerosol
liquid moisture input are dominated by thallus morphologies which expose large areas
of branch surface perpendicularly to the direct moisture-laden winds. This thallus
surface area may be exposed as flattened branches (Niebla josecuervoi) or dense
clumps of terete branches (Niebla "ceruchoides," Teloschistes spp.). Crustose growthforms of lichens on horizontal surfaces are poorly adapted for moisture uptake from
aerosol sources. Fruticose growth forms typically dominate, but some crustose forms
are ecologically successful on such surfaces by utilizing morphological adaptations to
increase thallus surface areas exposed to moist winds. Many taxa in groups typically
crustose become virtually fruticose, forming a clumped to caespitose growth form.
This morphological development can be best seen in Caloplaca coralloides and Lecanora phyganitis in the California-Baja California coastal flora. Raised verrucae in
coastal taxa of Thelomma may represent a similar adaptation.
On corticolous substrates, where moisture condensation on foliage and branches
produces considerable available water, the thallus growth form is less critical. Characteristically, however, thalli are most abundant well inside shrubs rather than on
small branches on the outer surface. On cacti in northern Chile, downward pointing
spines act as major condensation foci and epiphytic lichens are concentrated near the
spine tip rather than on the aereoles. Pendulous or tufted lichens near the spine tips
are ideally suited to maximize water uptake (Figure 3).
In inland areas of fog zones where major moisture input is in a water vapor form,
vertical orientation of thallus branches is less important and foliose and crustose
1978]
RUNDEL:
DESERT
IsUs
ng3~-
i-':?-:
p~nga
~~:~ ..........s
er
aa:i ..........
.......
7.......
..
hL
FIGURE3.
287
288
THE BRYOLOGIST
[Volume
81
growth forms increase in importance. Total biomass of lichens in such areas is relatively low, however (Kappen et al., 1975; Nash et al., 1977).
Many individual morphological characteristics of lichen thalli may also be important in water relations. Cilia frequently occur in taxa of desert fog zone lichens (e.g.
Heterodermia, Parmelia, Trichoramalina, Tornabenia). Although definitive experiments have not been completed, such cilia would be expected to have adaptive value
in straining aerosol water droplets out of humid air. Pseudocyphellae and related
white striations (maculae), present in many taxa, may also relate to water uptake.
These structures are positively correlated with populations of the Ramalina usnea
complex from foggy microenvironments (Rundel, 1978).
Thick cortices are characteristic of many desert fog zone lichens (Rundel & Bowler,
1974). It is doubtful if this adaptation is primarily a response to high light intensities
as in lichens from hot, dry deserts. More likely, cortical thickness is largely influenced
by potential mechanical damage and desiccation by wind. Morphological adaptations
to sand blasting in Antarctica include the formation of a cortex with thick-walled
hyphae closely appressed (Dodge, 1965). Such cortical structure is characteristic of
the Ramalinaceae in coastal deserts (Rundel, unpublished data).
PHYSIOLOGICAL
ADAPTATIONS
289
1978]
DENDROGRAPHALEUCOPHAEA
0.3-
0.1---------
0
CO2 FIXATION
(mg CO2gd-wt.:
h -I)
-0.I
-----
---------
45000
lux
-0.3
9000
lux
-0.5
1900 lux
dark
5
10
T5
20
Temperature (C)
25
30
290
[Volume 81
THE BRYOLOGIST
80-
70
60
Nieblo
50
Thallus
Moisture
Content
(%drywt.)
leucophoeo
ContueDendrogropho
4Nieblo
40
homaleo
30
20
10
10
14
18
Time
22
(hours)
26
30
34
38
42
FIGURE5. Water uptake by salt-encrusted and non-encrusted thalli of Dendrographa leucophaea Tuck. Darb. and Niebla homatea Ach. Rund. & Bowl. Points are mean value for four
thalli.
Comparative data for moisture uptake of air dry thalli previously saturated with
salt water and with deionized water are shown in Figure 5 for two species of lichens
from Baja California. In Niebla homalea (sand form) saturation at 100% relative humidity is reached at a water content of 40% of dry weight in thalli without a salt crust.
1978]
291
Publ. 354.
292
THE BRYOLOGIST
[Volume 81
. 1970. The Lichens of Israel. Israel Acad. Sci. Hum., Jerusalem. 116 p.
GANNUTZ,T. P. 1970. Photosynthesis and respiration of plants in the Antarctic peninsula area.
Antarctic Jour. 5: 49-51.
GEITLER, L. 1923. Studien iiber das Hamatochrom und die Chromatophoren von Trentepohlia.
Osterr. Bot. Zeitschr. 73: 76-83.
GIESS, W. 1962. Some notes on the vegetation of the Namib Desert. Cimbebasia, Windhoek,
No. 2.
-. 1968. A short report on the vegetation of the Namib coastal area from Swakopmund to
Cape Frio. Dinteria 1: 13-29.
HAJEK,E. & F. DI CASTRI. 1975. Bioclimatograffa de Chile. Univ. Catolica de Santiago.
HASTINGS, J. R. 1964. Climatological data for Baja California. Tech. Rep. on Meteorology and
Climatology of Arid Regions. No. 14. Univ. Ariz. Inst. Atm. Physics.
& R. R. HUMPHREY. 1969. Climatological data and statistics for Baja California. No. 18.
Univ. Ariz. Inst. Atm. Physics.
HiRISSET, A. 1946. Demonstration expierimental du r8le du Trentepohlia umbrina (Kg.) Born.
dans la synthese des Graphidees corticoles. C. R. Acad. Sci. 222: 100-102.
HOWLAND, L. J. 1929. Periodic observations of Trentepohlia aurea Martinus. Ann. Bot. 43: 173202.
JOHNSTON,I. M. 1929. Papers on the Flora of Northern Chile. Contr. Gray Herb. 85: 1-171.
KAPPEN, L. 1973. Response to extreme environment. In: V. Ahmadjian and M. E. Hale (eds.),
The Lichens. New York.
, O. L. LANGE, E.-D. SCHULZE,M. EVENARI& U. BUSCHBOM. 1975. Primary production
of lower plants (lichens) in the desert and its physiological basis. In: P. J. Cooper (ed.),
Photosynthesis and Productivity in Different Environments. London.
KOEPCKE,H. W. 1961. Synokologische Studien an der Westseite der peruanischen Anden.
Bonn. Geogr. Abh. No. 29.
LANGE, O. L. 1969. Experimentell-bkologische
Untersuchungen an Flechten der Negev-Wiiste.
I. CO2-Gaswechsel von Ramalina maciformis (Del.) Bory unter kontrollierten Bedingungen
im Laboratorium. Flora 158: 324-359.
& M. EVENARI. 1971. Experimentell-6kologische
Untersuchungen an Flechten der Negev-Wiiste. IV. Wachstumsmessungen an Caloplaca aurantia (Pers.) Hellb. Flora 160: 100104.
& L. KAPPEN. 1972. Photosynthesis of lichens from Antarctica. Antarctic Research Series
(G. A. Llano, ed.). 20: 83-95.
- , E.-D. SCHULZE, L. KAPPEN, U. BUSCHBOM& M. EVENARI. 1975. Adaptations of desert
lichens to drought and extreme temperatures. In: N. F. Hadley (ed.), Environmental Physiology of Desert Organisms. Stroudsberg, Penn.
- , E.-D. SCHULZE& W. KOCH. 1970a. Experimentell-6kologische
Untersuchungen an Flechten der Negev-Wiiste. II. CO2 Gaswechsel und Wasserhaushalt von Ramalina maciformis
(Del.) Bory am natiirlichen Standort wahrend der sommerlichen Trockenperiode. Flora 159:
38-62.
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