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The lichen genus Acarospora in Israel and its vicinity

Article  in  Nova Hedwigia · May 2005

DOI: 10.1127/0029-5035/2005/0080-0433

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 The lichen genus Acarospora in Israel and its vicinity

by

Marina Temina, Eviatar Nevo*, and Solomon P. Wasser

Institute of Evolution, University of Haifa, Mount Carmel, Haifa 31905, Israel

Abstract: Thirteen species of the lichen genus Acarospora, all occurring in Israel and its vicinity, are
described. A key for these species is given. Distribution of these species is discussed.

Introduction
The cosmopolitan genus Acarospora includes ca. 200 species (Nimis 1993). Acarospora
species occur usually on siliceous rocks, sometimes on calcareous rocks, and very rarely - on
wood, soil, and thallus of other lichens. The distribution of some Acarospora species in the
world is poorly known because of taxonomic and diagnostic problems (Wirth 1995).
Species of the genus are subdivided into two groups, which differ in thallus color. There is a
group of yellow Acarospora species and a group of brown, grey or whitish Acarospora
species ("Phaeothallia" H. Magn.). Most yellow Acarospora species occur in areas with arid
climate (Santesson 1944). These species have two distribution centers in the world
(Golubkova 1980, 1983, 1988). One of them is in the subtropical areas of southern and
southwestern Africa: in the Kalahari, Namib, and Karroo Deserts. The second center is in the
arid mountainous areas of North America. High species richness of yellow Acarospora is
known also for the Mediterranean region and South America (Golubkova 1980, 1983, 1988).
The distribution of the "Phaeothallia" species is centered in mountain regions of temperate
Europe (Golubkova 1983, 1988). Most of these species occur in European boreal zone. Many
"Phaeothallia" species are recorded also in southern Europe, Central Asia, North Africa, and
southwestern North America (Golubkova 1981, 1983, 1988).
Contributions to the study of the genus Acarospora have been made by Magnusson (1924,
1929a, b; 1937, 1956), Poelt & Steiner (1971), Poelt & Vezda (1977), Golubkova et al.
(1978), Golubkova (1980, 1988), and Clauzade & Roux (1981, 1985).
In Israel and its vicinity the genus Acarospora is still insufficiently known. In the
monography of M. Galun (1970), “The Lichens of Israel”, four species from this genus
occurring in Israel were presented. They were A. areolata, A. bornmuelleri, A. murorum, and
A. nodulosa var. reagens. A. areolata was found also in the Sinai Peninsula (Galun & Garty
1972). The fifth species from this genus was recorded in the studied region only in 2000. A.
cervina was found in the Mount Hermon area (Temina et al. 2002). All of these lichens
belong to "Phaeothallia". Here we describe 13 species of Acarospora, all from Israel and its
vicinity.
Materials and Methods

The paper is based on the study of the lichenological collection at the herbaria of Tel-Aviv University and also
on the results of some expeditions of the first author to various regions of the studied territory from September
2000 to August 2002. Lichen species were collected in 9 localities from the following natural regions: Mount
Hermon, Golan Heights, Upper Galilee, Central Negev, and Southern Negev. The natural regions of Israel and

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its vicinity are given according to Zohary & Feinbrun-Dothan (1966-1986) (Fig. 1).

Localities:
1) Mount Hermon: Ski resort – upper cable station, 2000 m a.s.l., valley, (3317’N; 3545’E), 18 August 2002.
2) Mount Hermon: Ski resort – lower cable station, 1600 m a.s.l., SE-facing slope, (3317’N; 3544’E), 26
September 2000.
3) Mount Hermon: NE vicinity of the village of Majdal Shams, 1300 m a.s.l., SE-facing slope, old fruit garden,
(3316’N; 3545’E), 27 September 2000.
4) Golan Heights: S vicinity of the village of Mas’ada, 800-900 m a.s.l., oak forest, (3313’N; 3545’E), 20
September 2000.
5) Golan Heights: vicinity of Ziwan, (3305’N; 3547’E), 600-700 m a.s.l., 17 May 2001.
6) Golan Heights: Nahal Meitsar, canyon, (3244’N; 3543’E), 5 June 2002.
7) Upper Galilee: vicinity of Dalton, 800 m a.s.l., (3301’N; 3528’E), 13 January 2001.
8) Central Negev: Negev highland, Makhtesh Ramon Nature Reserve, 900-1000 m alt., (3037’N; 3454’E), 29
September 2000.
9) Southern Negev: Nahal Shaharut - “Evolution Canyon” III, (2955’N; 3458’E), 16 January 2002.

All lichen samples were collected for precise determination in the laboratory. Observations and measurements of
morphological characters of lichen specimens were made under a dissecting microscope. Anatomical details
were obtained by examining hand-cut sections or squash preparations under a light microscope. Microscopic
preparations were mounted in water, 10% potassium hydroxide or Lugol’s solution. Measurements of anatomical
characters were made in water at x 100-1000 magnification. The thickness of paraphyses was measured in the
basal third and apex. The number of spores in ascus was evaluated following an optical section. Only well-
developed spores lying outside the asci were measured. The spot tests on the thallus and medulla were made
using 10% potassium hydroxide (K), calcium hypochlorite (C), and para-phenylendiamine in ethanol (Pd).
Iodine reactions in the hymenium have been tested in Lugol’s solution (I). The following references were used to
identify the specimens: Galun & Reichert (1960), Poelt (1969), Galun (1970), Golubkova et al. (1978),
Golubkova (1979, 1988), Clauzade & Roux (1981, 1985), Wirth (1995), Brodo et al. (2001). To verify
identification of new species, specimens from lichenological collections of the Department of Lichenology &
Bryology of Komarov Botanical Institute (St. Petersburg, Russia) were used as comparative material. Lichen
nomenclature is given according to Kirk et al. (2003). Herbarium specimens are deposited at the herbaria of the
Institute of Evolution, University of Haifa (HAI, Israel).

Fig. 1. Map of natural regions of Israel and its vicinity:

AP – Akko Plain; AV – Arava Valley; BS – Bet Shean Valley; CC – Carmel Coast; CM – Mount Carmel; CN –
Central Negev; DS – Dead Sea Area; EP – Esdraelon Plain; GC – Galilee Coast; GH – Golan Heights; GM –
Gilboa Mountains; HE – Hermon; HP – Hula Plain; JD – Judean Desert; JM – Judean Mountains; LG – Lower
Galilee; LJ – Lower Jordan Valley; NN – Northern Negev; PP – Philistine Plain; SA – Samaria; SH – Shefela;

434
SN – Southern Negev; SP – Sharon Plain; UG – Upper Galilee; UJ – Upper Jordan Valley; WN – Western
Negev.

Results

Key to the species

1. Thallus distinctly yellow, thick, squamulose; squamules 1-3 mm wide, convex or flat,
surface finely fissured. Apothecia 0.1-0.4 mm diam., with prominent thallus-colored margins.
Disc punctiform, reddish brown to very dark brown, often broken up with patches or a
network of sterile thallus-colored tissue; paraphyses 1-1.5 μm wide; ascospores narrowly
ellipsoidal …………..……………….…………………………...….. 4. Acarospora contigua
- Thallus brown, sand-colored, grey or white, without a yellowish tint ……………….…….. 2
2. Thallus chalk-white or greyish white ……………………………………………………... 3
- Thallus brown, grey- to black-brown, reddish brown, fawn, or sand-colored …………..…. 4
3. On soil. Thallus chalk-white, squamulose, continuous, effigurate or not …………………...
…………….……………………………..……….…. 11. Acarospora nodulosa var. reagens
- On rocks. Thallus greyish white, areolated, dispersed or contiguous, indeterminate ………
…….……………………………………………………….……….... 1. Acarospora areolata
4. Thallus verrucose, squamulose or areolated, sometimes inconspicuous .……….………… 5
- Thallus rosetted, rounded or lobated, with verrucose, squamulose or areolate crust in center,
and well-developed margin lobes ……………………………………………...……...…..... 12
5. Ascospores small, 3-5(6) μm long and 1.5-2.5(4) μm wide, sometimes globose, 3-5 μm
diam. Asci 100- to 200-spored ………………………………………...…………...………... 6
- Ascospores larger, 6-13 μm long and 3-6 μm wide, ellipsoidal. Asci to 100-spored ………...
……………….…………………………………………………… 10. Acarospora murorum
6. Ascospores globose ….…………………………….…..….... 3. Acarospora bornmuelleri
- Ascospores ellipsoidal ..…………………………….…..…………………..……...………. 7
7. Thallus C+ red, KC+ red …………………………………..………….………………...… 8
- Thallus C-, KC- ………………………………………………………….…….……..….…. 9
8. Thallus ± densely white-pruinose; areoles markedly convex; apothecia 0.1-0.5 mm diam.;
disc mostly rounded …………….…………….……………........ 13. Acarospora umbilicata
- Thallus without pruina or if pruinose then areoles flat; apothecia 0.2-1 mm diam.; disc
usually angular ………… ………………………………………….… 5. Acarospora fuscata
9. On thalli of other lichens (Aspicilia sp.) …...……………….... 12. Acarospora tominiana
- Not on thalli of other lichens …………………………..………….…...…………….….… 10
10. Thallus poorly developed, usually visible only as a rim surrounding apothecia …………...
……………………………………..…………… 7. Acarospora glaucocarpa var. conspersa
- Thallus well developed, areolated to squamulose ………………………………………… 11
11. Thallus areolated-squamulose; areoles pale brown-grey, with white margins and
undersides; apothecia 0.5-2 mm diam. …………… 6. Acarospora glaucocarpa var. cervina
- Thallus areolated; areoles dark red-brown to black-brown, margins concolorous with upper
surface, underside dark brown; apothecia 0.2-0.5 mm diam. …...... 8. Acarospora impressula
12. Ascospores globose …………………………….………..… 3. Acarospora bornmuelleri

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- Ascospores ellipsoidal …………………………………………………………....……….. 13
13. Thallus C+ red, KC+ red ……………………….…….…....…. 2. Acarospora assimulans
- Thallus C-, KC- ……….………………………………………………….….……………. 14
14. Thallus distinctly lobed, densely white- or blue-grey-pruinose; lobes and areoles
surrounded by thick, raised, white rim; apothecia 0.3-0.5 mm diam., without margins …….…
………………………….. ………………...……………….………. 9. Acarospora laqueata
Thallus indistinctly lobed, not or slightly pruinose; areoles surrounded by narrow, white rim;
apothecia 0.5-2 mm diam., with thin margin, concolorous with disc ……………………….....
…………………………………………………..… 6. Acarospora glaucocarpa var. cervina

1. Acarospora areolata Reichert & Galun, Bull. Res. Counc. Israel, 9: 132, 1960.
Description. Thallus areolated; areoles (0.5)1-2.5 mm diam. and 0.4-0.8 mm thick, dispersed,
in small groups or contiguous, greyish white, densely pruinose, rough and granulous, slightly
convex, the larger areoles occasionally with thin splits; lower side pale, firmly attached to the
substrate. Photobiont chlorococcoid. Algal layer 150-300 μm thick, interrupted by medullary
hyphae perpendicular to the surface. Medulla greyish, made opaque by masses of substrate
particles. Apothecia numerous, mostly solitary, sometimes 2-3 to each areole, with thalline
margin concolorous with the thallus. Disc on a level with the thallus, 0.5-1.5 mm diam.,
black, dull, plane. Hymenium colorless, 100-150 μm tall. Epithecium yellowish brown.
Hypothecium colorless, grumous. Exciple colorless, 15-40 μm thick at the base and expanding
to 150 μm above. Paraphyses septate, 2 μm wide at the base; apices slightly swollen and
yellowish brown, conglutinate. Asci 100-spored. Ascospores cylindrical, 4.5-6 x 1.5-2 μm.
Thallus K-, C-, KC-; hymenium I+ blue.
Habitat. On sun-exposed, dry, calcareous rocks.
Distribution in Israel and vicinity. Judean Desert (Galun 1970, Galun & Mukhtar 1996,
Kondratyuk et al. 1996); Dead Sea area (Galun & Mukhtar 1996); Central Negev (Galun &
Reichert 1960, Galun 1970, Galun & Mukhtar 1996, Kondratyuk et al. 1996), Negev
highland, Makhtesh Ramon Nature Reserve (Temina: HAI, 000613; HAI, 000621); Southern
Negev: Nahal Shaharut - “Evolution Canyon” III (Temina: HAI, 012211); Sinai (Galun &
Garty 1972, Galun & Mukhtar 1996, Kondratyuk et al. 1996).
General distribution. Asia [Egypt (Sinai), Israel].
Note. According to Galun & Reichert (1960), Acarospora areolata resembles two species: A.
bicolor Vain. and A. glaucocarpa var. cervina (A. Massal.) Clauzade & Cl. Roux. It is
distinguished from A. bicolor by shape and width of ascospores, and higher hymenium (A.
bicolor has a 3 μm width broadly ellipsoidal to subglobose ascospores and hymenium of 85-
100 μm high, whereas A. areolata has a 1.5-2 μm width cylindrical ascospores and hymenium

436
of 100-150 μm high). A. areolata is distinguished from A. glaucocarpa var. cervina by a
greyish white thallus, having a distinct, thick exciple, and absence of the lower cortex
whereas A. glaucocarpa var. cervina has a grey-brown thallus, indistinct exciple, and a 15-30
μm thick lower cortex.

2. Acarospora assimulans Vain., Acta Horti Petropol., 10: 559, 1888. (Pl. 1, fig. A).
Description. Thallus rosetted, 1-3 cm diam., with distinct marginal lobes, pale brown to
fawn; center areolated; areoles and lobes surrounded by a narrow, dark rim. Lobes 1-5 mm
long and 0.5-2 mm broad, plane, on the ends expanded, deeply incised. Areoles 0.3-1.8 mm
diam., irregularly rounded or angular, plane, with dark margins and undersides. Photobiont
chlorococcoid. Algal layer 60-120 μm thick, continuous. Medulla thick, well developed.
Apothecia immersed, 2-5 and more per areole, sometimes solitary, at first punctiform, later
concave-flat, without thalline margin. Disc 0.6-1 mm diam., reddish brown, rough and
umbonate. Hymenium colorless, 100-135 μm tall. Epithecium brightly brown. Hypothecium
35-56 μm tall, greyish, granular. Exciple distinct, colorless, 14-17 μm thick at the base and
expanding to 40-60 μm above. Paraphyses septate, 2-2.5 μm wide at the base; apices slightly
swollen, brown. Asci 85-116 x 15-22 μm, elongated-clavated, 200-spored. Ascospores
broadly ellipsoidal, 4-5.5 x 2-3.5 μm. Thallus K-, C+ red, KC+ red; hymenium I+ blue.
Habitat. On basalt rocks in open habitats.
Distribution in Israel and vicinity. Golan Heights: Nahal Meitsar - canyon (Temina: HAI,
021031), Mas’ada, oak forest (Temina: HAI, 001014); Upper Galilee: vicinity of Dalton
(Temina: HAI, 012431).
General distribution. Asia (Afghanistan, Israel, Kazakhstan, Tajikistan, Turkmenistan),
North Africa (Algeria).
Note. According to literature data, this species occurs on calcareous rocks.

3. Acarospora bornmuelleri J. Steiner, Ann. Naturh. Hoffm., 30: 29, 1916.

Description. Thallus areolated or areolated-squamulose, forming patches to 2-5 cm diam.,
thick, dark red-brown; areoles plane and even, angular, 0.4-2 mm diam., circumference
sublobate, with dark brown undersides. Photobiont chlorococcoid. Algal layer 70-120 μm
thick, continuous or in groups separated by strands of parallel hyphae. Medulla thick, contains
many crystals and substrate particles. Apothecia rare to rather numerous, immersed, usually
solitary, sometimes 2-3 to each areole, without thalline margin. Disc on level with the thallus,
plane, granular, 0.5-1 mm diam., concolorous with the thallus or darker. Hymenium colorless,

437
75-120 μm tall. Epithecium brown. Hypothecium 25-35 μm tall, colorless or greyish. Exciple
15-20 μm thick, composed of parallel radiating hypothecial hyphae, colorless except for the
brown-pigmented exterior cell walls. Paraphyses coherent, septate, 2 μm wide at the base,
with brown apical cells to 4 μm wide. Asci 55-75 x 14-17 μm, 100-spored. Ascospores
subglobose or globose, 3-5 μm diam. Thallus K-; cortex C+ red (a very thin line of coloration
is formed at the upper part of the cortex below the brown region); hymenium I+ blue.
Habitat. On calcareous and basalt rocks in open and dry sites.
Distribution in Israel and vicinity. Upper Jordan Valley (Galun 1970, Kondratyuk et al.
1996); Upper Galilee (Galun & Mukhtar 1996, Kondratyuk et al. 1996); Lower Galilee
(Galun 1970, Galun & Mukhtar 1996, Kondratyuk et al. 1996); Sinai (Galun & Garty 1972,
Galun & Mukhtar 1996, Kondratyuk et al. 1996).
General distribution. Asia [Armenia, Egypt (Sinai), Iraq, Israel, Jordan, Kyrgyzstan,
Mongolia, Russia, Syria, Turkey].
Note. According to Magnusson (1929a), Acarospora bornmuelleri occurs on granitic or
volcanic rocks. This species was also recorded on basalt rocks in Mongolia (Golubkova
1981).

4. Acarospora contigua H. Magn., Mycologia 21: 256, 1929. (Pl. 1, fig. C).
Synonyms. Acarospora bella (Nyl.) Jatta, Acarospora xanthophana (Nyl.) Jatta.
Description. Thallus squamulose, scattered or contiguous, bright sulphur-yellow to greenish
yellow; squamules 1-3 mm wide, irregular in shape, convex or flat, surface finely fissured,
with smooth or scalloped margins. Photobiont chlorococcoid. Algal layer 80-150 μm thick,
irregular, discontinuous. Medulla thick, well developed, greyish or white contains many
crystals. Apothecia immersed, 1-4 per squamule, sometimes becoming somewhat prominent
with thallus-colored margins. Disc 0.1-0.4 mm diam., punctiform, reddish brown to very dark
brown, often broken up with patches or a network of sterile thallus-colored tissue. Hymenium
colorless, 130-180 μm tall. Epithecium yellow. Hypothecium 20-35 μm tall. Exciple colorless,
10-35 μm thick at the base and expanding to 50 μm above. Paraphyses 1-1.5 μm wide at the
base, 2-3 μm wide at the apex. Asci 100-135 x 17-25 μm. Ascospores narrowly ellipsoidal, 3-
5 x 2-3 μm diam. Thallus K-, C-, KC-; hymenium I+ blue.
Habitat. On basalt rocks in dry, open habitats.
Distribution in Israel and vicinity. Golan Heights: Nahal Meitsar - canyon (Temina: HAI,
021033).
General distribution. Asia (China, Israel), Africa (Morocco, South Africa), North America

438
(Mexico, USA), Hawaii, South America (Argentina, Bolivia, Chile, Colombia).
Note. According to Golubkova (1983), this species is closely related to Acarospora fulva N.S.
Golubk. It is distinguished from A. fulva by negative reactions of thallus with C and KC. A.
fulva has the C+ pink and KC+ red thallus (Golubkova 1979, 1988).

5. Acarospora fuscata (Nyl.) Arnold, Verh. zool.-bot. Ges. Wien, 20: 528, 1870. -
Lecanora fuscata Nyl., Not. Sällsk. F. Fl. Fenn. Förh., 8: 138, 1882. (Pl. 1, fig. D).
Description. Thallus areolated-squamulose, wide-spreading and often forming extensive
patches, rarely scattered; areoles 0.5-3 mm wide, very variable, mostly contiguous, irregular
in shape, ± flat, at times sublobate or weakly imbricate, surface uneven, irregular, pale to dark
reddish or yellowish brown, often with black margins and undersides. Photobiont
chlorococcoid. Algal layer 40-90 μm thick, continuous. Apothecia solitary or 2-5 per areole,
usually numerous, at first punctiform, later concave-flat, usually angular or irregular,
immersed, never elevated or emergent, without thalline margin. Disc smooth or slightly
rough, level with thallus or ± depressed, 0.2-1 mm diam., reddish brown to blackish, usually ±
darker than the thallus. Hymenium 80-120 μm tall. Epithecium yellowish brown.
Hypothecium 40-60 μm tall. Exciple distinct, 10-30 μm thick. Paraphyses conglutinate, 1.5
μm wide at base, 2-3 μm wide at the apex. Asci 60-85 x 15-17 μm, 100- to 200-spored.
Ascospores 4-6 x 1-2 μm, narrowly ellipsoidal or ± cylindrical. Thallus K-, C+ red, KC+ red;
hymenium I+ blue or red.
Habitat. On sun-exposed basalt rocks.
Distribution in Israel and vicinity. Upper Galilee: vicinity of Dalton (Temina: HAI,
012432).
General distribution. Cosmopolitan.
Note. Acarospora fuscata is a very variable species but characterized by the C+ red reaction
of the thallus. Pruinose form of A. fuscata resembles A. umbilicata Bagl. See further note
under A. umbilicata.

6. Acarospora glaucocarpa var. cervina (A. Massal.) Clauzade & Cl. Roux, Bull. Soc. linn.
Provence, 53: 147, 2002 - Acarospora cervina A. Massal., Ric. Auton. Lich. Crost.: 28, 1852.
(Pl. 1, fig. B).
Description. Thallus areolated-squamulose, often forming extensive patches; areoles or
squamules contiguous or ± overlapping, 0.5-3 mm wide, rather thick, rounded or angular to
slightly lobed, flat or convex, surface smooth or slightly roughened, pale brown-grey, matt,

439
not or slightly pruinose, usually with a white margins, underside white. Photobiont
chlorococcoid. Algal layer 90-200 μm thick, irregular, discontinuous, tooth-like below.
Medulla greyish, made opaque by masses of crystals and substrate particles. Apothecia
immersed, usually solitary, sometimes 2-4 per areole, 0.5-2 mm diam., with a thin, smooth
margin, and dark brown-red, ± naked disc. Hymenium 80-100 μm tall. Epithecium brightly
brown. Hypothecium 30-50 μm tall. Exciple indistinct. Paraphyses free, 2-3 μm wide at the
base; apices swollen to 4 μm, with brown cap. Asci 45-80 x 12-17 μm, 100-spored.
Ascospores 4-6 x 2-4 μm, cylindrical. Thallus K-, C-, KC-; hymenium I+ blue.
Habitat. On sun-exposed calcareous rocks, somewhat nitrophilous.
Distribution in Israel and vicinity. Mount Hermon: Ski resort – upper and lower cable
stations, NE vicinity of Majdal Shams, old fruit garden (Temina et al. 2002).
General distribution. Europe (widespread), Asia (Afghanistan, Armenia, Azerbaijan,
Georgia, Iran, Israel, Jordan, Kazakhstan, Kyrgyzstan, Syria, Tajikistan, Turkey,
Turkmenistan, Uzbekistan), Africa (Algeria, Morocco, Tunisia, South Africa), Madeira
Islands, North America, Australia.
Note. Acarospora glaucocarpa var. cervina is closely related to A. glaucocarpa var.
conspersa, and distinguished from it by well-developed, areolated-squamulose thallus, with
areoles surrounded by narrow, white rim and ± naked apothecial discs. A. glaucocarpa var.
conspersa characterized by a thallus reduced to margins around the apothecia and densely
pruinose apothecial discs.

7. Acarospora glaucocarpa var. conspersa Th. Fr., Lich. Arct.: 88, 1860. (Pl. 1, fig. E).
Synonym. Acarospora cervina var. conspersa (Th. Fr.) Clauzade & Cl. Roux
Description. Thallus poorly developed, usually visible only as a rim surrounding apothecia.
Photobiont chlorococcoid. Apothecia 0.5-2 mm diam., solitary. Thalline margin thick, raised,
reddish brown, ± pruinose. Disc flat or concave, ± concolorous or slightly darker than the
thalline margin, densely pruinose. Hymenium 80-100 μm tall. Epithecium brown.
Hypothecium 30-45 μm tall, granular. Exciple colorless, 15-20 μm thick at the base and
expanding to 35-50 μm above. Paraphyses conglutinate, 2.5-3 μm wide at the base; apices
swollen to 4 μm, with brown cap. Asci 40-70 x 10-17 μm, 100-spored. Ascospores 3.5-5 x
1.5-2 μm, narrowly ellipsoidal. Thallus K-, C-, KC-; hymenium I+ blue.
Habitat. On sun-exposed, dry calcareous rocks.
Distribution in Israel and vicinity. Central Negev: Negev highland, Makhtesh Ramon
Nature Reserve (Temina: HAI, 000614).

440
General distribution. Europe (widespread), Asia (Armenia, Azerbaijan, Georgia, Israel,
Kazakhstan, Kyrgyzstan, Lebanon, Mongolia, Russia, Tajikistan, Turkey, Turkmenistan,
Uzbekistan), North Africa (Morocco), Greenland, North America (Canada, USA), South
America (Argentina).
Note. Acarospora glaucocarpa var. conspersa is closely related to A. glaucocarpa var.
cervina. See further note under A. glaucocarpa var. cervina.

8. Acarospora impressula Th. Fr., Lichenogr. Scand., 1: 214, 1871.

Synonym. Acarospora atrata Hue.
Description. Thallus areolated, contiguous, dark red-brown to black-brown; areoles 0.2-
1.5(2.5) mm wide, flat or slightly concave, variously angular, rounded or elongated; surface
smooth or slightly rough; underside dark brown. Photobiont chlorococcoid. Algal layer
irregular, ± continuous, not tooth-like below. Medulla greyish, contains many crystals.
Apothecia 0.2-0.5 mm diam., 1(2) per areole, immersed, without or with a thin thalline
margin. Disc level with the thallus, angular or round, concave, concolorous with the thallus or
blackish. Hymenium colorless, 90-115 μm tall. Epithecium dark red-brown or yellow-brown.
Hypothecium 50-60 μm tall, colorless. Exciple colorless, 14-30 μm thick. Paraphyses 2-3 μm
wide at the base; apices swollen to 4(5) μm, with brown cap. Asci 60-85 x 12-18 μm, 100-
spored. Ascospores 3-5 x 2-2.5 μm, ellipsoid. Thallus K-, C-, KC-; hymenium I+ blue.
Habitat. On basalt rocks in sunny, open habitats.
Distribution in Israel and vicinity. Golan Heights: vicinity of Ziwan (Temina: HAI,
011021).
General distribution. Europe (Austria, Great Britain, France, Germany, Italy, Norway,
Poland, Russia, Sweden, Switzerland, Ukraine), Asia (Armenia, Azerbaijan, Israel,
Kazakhstan, Kyrgyzstan, Mongolia, Russia, Tajikistan), North Africa (Morocco), Greenland,
North America.
Note. Identification of our specimen of Acarospora impressula was confirmed by Prof. N.S.
Golubkova.
According to literature data, Acarospora impressula occurs on calciferous or siliceous rocks,
frequently on the thalli of other crustose lichens.

9. Acarospora laqueata Stizenb. ex Flagey, Herbor. Lich. Envir. Constantine: 131, 1888. (Pl.
1, fig. F).
Description. Thallus rosetted, often forming extensive patches to 11 cm diam., with distinct

441
marginal lobes, pale or dark brown, densely white- or blue-grey-pruinose; center areolated.
Lobes 0.6-2.2 mm long and 0.2-0.9 mm broad, closely adpressed to each other, plane, smooth,
on the ends expanded and incised, surrounded by a white rim. Areoles 0.2-0.8 mm wide,
plane, angular, with white margins. Photobiont chlorococcoid. Algal layer 50-65 μm thick,
continuous. Medulla thick, greyish, contains many crystals. Apothecia immersed, 1(2) per
areole, without thalline margin. Disc 0.3-0.5 mm diam., rounded or angular, concave reddish
brown, naked. Hymenium colorless, 80-120 μm tall. Epithecium brightly brown.
Hypothecium 30-100 μm tall, greyish, granular. Exciple 14-17 μm thick, inconspicuous.
Paraphyses septate, 2.5-3.5 μm wide at the base; apices slightly swollen, brown. Asci 72-85 x
15-17 μm, 100-spored. Ascospores 4-5.5 x 2-3 μm, ellipsoid. Thallus K-, C-, KC-; hymenium
I+ blue.
Habitat. On basalt rocks in sunny, open habitats.
Distribution in Israel and vicinity. Golan Heights: Nahal Meitsar - canyon (Temina: HAI,
021032).
General distribution. Europe (France, Italy, Spain, Switzerland), Asia (Iraq, Israel, Jordan,
Kyrgyzstan, Tajikistan, Turkey, Turkmenistan), North Africa (Algeria, Morocco, Tunisia).
Note. According to literature data, this species occurs on calcareous rocks.

10. Acarospora murorum A. Massal., Mem. Lichenogr.: 130, 1853.

Synonyms. Acarospora macrospora subsp. murorum (A. Massal.) Clauzade & Cl. Roux,
Acarospora macrospora var. murorum (A. Massal.) Anzi.
Description. Thallus areolated-squamulose, indeterminate, more or less concrescent into an
areolated crust, sand-colored; squamules 0.7-2 mm diam., plane, irregularly rounded or
angular, underside white. Photobiont chlorococcoid. Algal layer 40-70 μm thick, irregularly
interrupted by perpendicular strands of medullary hyphae. Medulla nubilated by granules and
substrate particles; lower part of medulla brown. Apothecia numerous, immersed, solitary.
Thalline margin concolorous with the thallus, slightly prominent. Disc 0.5-1 mm diam., red-
brown or dark brown, naked, concave or plane, on a level with the thallus. Hymenium 110-
160 μm tall. Epithecium yellowish brown. Hypothecium colorless, indistinctly separated from
the colorless hymenium. Exciple colorless, 15-20 μm thick at the base and expanding to 30
μm above, sometimes inconspicuous. Paraphyses coherent, septate, 2-3 μm wide at the base,
3-4 μm wide at the apex. Asci 100-120 x 17-32 μm, 30-to100-spored. Ascospores ellipsoidal,
6-13 x 3-6 μm. Thallus K-, C-, KC-; hymenium I+ blue.
Habitat. On fine sandy soil on shady north-facing slopes.

442
Distribution in Israel and vicinity. Central Negev (Galun 1966, 1970, Galun & Mukhtar
1996, Kondratyuk et al. 1996).
General distribution. Southern Europe, Asia (Israel), North Africa.
Note. This species resembles Acarospora macrospora (Hepp) Bagl., but is distinguished from
it by a sand-colored thallus and solitary per squamule apothecia, whereas A. macrospora has
more dark reddish brown thallus and apothecia numerous per areole. According to Nimis
(1993), these two species have also different ecology and distribution.
According to literature data, A. murorum is most usually saxicolous and only rarely
terricolous.

11. Acarospora nodulosa var. reagens (Zahlbr.) Clauzade & Cl. Roux, Bull. Mus. Hist. Nat.
Marseille, 41: 61, 1981. - Acarospora reagens Zahlbr., Beih. Bot. Centralblatt, 13: 162, 1902.
Synonym. Acarospora reagens f. radicans (Nyl.) H. Magn.
Description. Thallus squamulose, continuous, chalk-white, forming patches to 5 cm diam.;
surface occasionally cracked, covered with a thick whitish pruina. Squamules 3-10 mm diam.
and 0.2-2 mm thick, rounded, slightly lobed. Photobiont chlorococcoid. Algal layer 50-120
μm thick, continuous. Medulla nubilated with crystals and substrate particles, connected to the
substrate by thick root-like medullary hyphae. Apothecia numerous, up to 1.2 mm diam.,
mostly solitary, rarely several per squamule, at first immersed, later sessile, with thalline
margin concolorous with the thallus. Disc plane or slightly convex, more or less level with the
thallus, black, naked. Hymenium 170-180 μm tall, colorless. Epithecium yellowish brown.
Hypothecium colorless, granular, 40-85 μm tall. Exciple 15-20 μm thick. Paraphyses 1.5-2.5
μm wide at the base, simple, septate, with slightly broadened brown apical cell. Asci 72-140 x
10-30 μm, 100-spored. Ascospores globose, 3-5 μm diam. Thallus K+ yellow turning red, C-;
medulla Pd+ orange yellow; hymenium I+ blue.
Habitat. On fine sandy soil and on loess in dry, open habitats.
Distribution in Israel and vicinity. Judean Desert (Galun & Reichert 1960, Galun 1970,
Galun & Mukhtar 1996, Kondratyuk et al. 1996); Central Negev (Galun & Reichert 1960,
Galun 1970, Galun & Mukhtar 1996, Kondratyuk et al. 1996); Western Negev (Galun &
Reichert 1960, Galun 1963, 1970, Galun & Mukhtar 1996, Kondratyuk et al. 1996).
General distribution. Southern Europe, Asia (Armenia, Azerbaijan, Georgia, Iran, Israel,
Kazakhstan, Kyrgyzstan, Tajikistan, Uzbekistan), North Africa (Algeria, Egypt, Morocco,
Tunisia), Canary Islands, North America (Mexico, USA), Australia.

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Note. Acarospora nodulosa var. reagens differs from a similar species A. nodulosa var.
nodulosa (Dufour) Hue in the K+ red reaction of the thallus (Golubkova et al. 1978, Nimis
1993).
According to Magnusson (1929a), A. nodulosa var. reagens is characterized by the absence of
exciple or a very thin (10 μm) exciple. Specimens from Israel have a 15-20 μm thick exciple.

12. Acarospora tominiana H. Magn., Kungl. Svenska Vetensk. Akad. Handl., ser. 3, 7 (4):
216, 1929.
Synonym. Acarospora impressula f. tominiana (H. Magn.) N.S. Golubk.
Description. Thallus dark brown, squamulose, dispersed or contiguous, making small areas
over or between the areoles of other crustose lichens. Squamules 0.5-1.0 mm diam., rounded,
convex, smooth. Photobiont chlorococcoid. Algal layer 85-145 μm thick, continuous. Medulla
greyish, contains many crystals. Apothecia 0.2-0.5 mm diam., usually solitary, rarely several
per squamule, immersed, with a thin, smooth thalline margin. Disc punctiform or angular,
concave, concolorous with the thallus. Hymenium 85-100 μm tall. Epithecium dark brown.
Hypothecium 35-50 μm tall, greyish, granular. Exciple ca. 15 μm thick at the base and
expanding to 30 μm above. Paraphyses 1.5-2.5 μm wide at the base, septate, apices slightly
swollen, brownish. Asci 70-85 x 13-17 μm, 100-spored. Ascospores broadly ellipsoid, 3-6 x
2.5-3.5 μm. Thallus K-, C-, KC-; hymenium I+ blue.
Habitat. On thallus of Aspicilia calcarea on limestone in sunny and dry habitats.
Distribution in Israel and vicinity. Mount Hermon: NE vicinity of Majdal Shams, old fruit
garden (Temina: HAI, 001218).
General distribution. Europe [Italy (Sardinia), Russia], Asia (China, India, Israel, Mongolia,
Russia, Tajikistan, Uzbekistan).
Note. This species is closely related to Acarospora hospitans H. Magn. It is distinguished
from A. hospitans by shape of squamules, larger cortex cells, and higher hymenium. A.
hospitans has a 1.5-2.5 μm diam. cortex cells and hymenium of 60-90 μm high, whereas A.
tominiana has a 5-6 μm diam. cortex cells and hymenium of 85-100 μm high (Golubkova et
al. 1978). These two species have also different ecology and distribution. A. tominiana occurs
on calcicolous Aspicilias of the A. calcarea group and is distributed in areas with a dry
continental climate, whereas A. hospitans is a parasite on siliceous crustose lichens and has
mainly arctic-alpine distribution (Nimis 1993).

13. Acarospora umbilicata Bagl., Mem. Reale Accad. Sci. Torino, ser. 2, 17: 397, 1857.

444
Description. Thallus squamulose, greyish-brown, densely white-pruinose. Squamules 0.5-1.5
mm diam., scattered to contiguous, markedly convex, angular to sublobate. Photobiont
chlorococcoid. Algal layer 35-165 μm thick, continuous. Apothecia 0.1-0.5 diam., 1 or 2-5 per
squamule, immersed, often coalescing, without thalline margin. Disc rounded, rarely angular,
concave to flat, smooth, brown-black. Hymenium 85-110(135) μm tall. Epithecium brown.
Hypothecium 20-35 μm tall. Exciple 8-14 μm thick at the base and expanding to 30 μm above,
yellowish. Paraphyses conglutinate, 1.5 μm wide at base, 2-3 μm wide at tips. Asci 70-100 x
15-17 μm, 100- to 200-spored. Ascospores 4-5 x 1.5-2 μm, cylindrical. Thallus K-, C+ red,
KC+ red; hymenium I+ blue or red.
Habitat. On sun-exposed basalt rocks.
Distribution in Israel and vicinity. Golan Heights: vicinity of Ziwan (Temina: HAI,
011022); Upper Galilee: vicinity of Dalton (Temina: HAI, 012433).
General distribution. Europe (Austria, Great Britain, France, Germany, Italy, Norway,
Poland, Spain, Sweden, Ukraine), Balearic Islands, Asia (Israel), North Africa (Morocco),
Canary Islands, Madeira Islands.
Note. This species resembles Acarospora versicolor Bagl. & Carestia and pruinose form of A.
fuscata. It differs from A. versicolor in the C+ red reaction of the thallus (Golubkova et al.
1978, Golubkova 1988). It is distinguished from pruinose form of A. fuscata by having
markedly convex squamules (A. fuscata has ± flat squamules) and smaller apothecia.

Discussion

Among the lichen species mentioned in this study, eight are recorded for the first time in
Israel and its vicinity. Seven of them belong to "Phaeothallia". They are: Acarospora
assimulans, A. fuscata, A. glaucocarpa var. conspersa, A. impressula, A. laqueata, A.
tominiana, and A. umbilicata. The lichen A. contigua belongs to a group of yellow
Acarospora species. Our new records bring the total number of known Acarospora in our
investigated territory to 13, but this number is probably not final.

Taxonomy
Acarospora contigua
According to Golubkova (1983), this species is morphologically and anatomically very
similar to A. fulva, which occurs on granitic rocks and on the thalli of other lichens and is
distributed in the mountain steppes of Mongolia and the Asiatic part of Russia (Altay
mountains, The Baikal ridge) (Golubkova 1981, 1988). Possibly, it will be more correct to

445
consider A. fulva as a variety of A. contigua, because these species are distinguished from
each other by only the different chemistry. A. contigua contains rhizocarpic acid, and A. fulva
contains rhizocarpic and gyrophoric acids (Golubkova 1979, 1983).
Some lichenologists include saxicolous species A. contigua in a terricolous A. schleicheri.
However, these species are quite distinct in morphological and anatomical characters and
have different ecology.

Acarospora glaucocarpa var. cervina

Lichen A glaucocarpa is very variable, and is often recognized as two distinct species: A.
cervina and A. glaucocarpa. According to Roux & Gueidan (2002), A. cervina is a variety of
A. glaucocarpa, because two taxa have insignificant morphological and anatomical
differences and many transitive forms. We also did not find significant distinctions between
these taxa in our material. According to literature data, these taxa have also similar ecology
and distribution. Therefore, here we consider A. cervina as a variety of A. glaucocarpa.

Acarospora tominiana
This species resembles A. hospitans, which Clauzade & Roux (1981) consider as a variety of
A. impressula. Golubkova (1988) consider A. tominiana as a form of A. impressula. We prefer
to treat the two taxa as distinct species because they have morphological and anatomical
distinctions and also different ecology and distribution. According to literature data, A.
tominiana is an obligate parasite of calcicolous Aspicilia species of the A. calcarea group and
is distributed in areas with a dry continental climate, whereas A. impressula occurs on
calciferous or siliceous rocks, frequently on the thalli of other crustose lichens, especially
Aspicilia polychroma Anzi and has arctic-alpine to boreal-montane distribution (Nimis 1993).

Acarospora nodulosa var. reagens.

The description of the exciple of A. nodulosa var. reagens does not correspond to that by
Magnusson (1929a). According to his description, this species has no exciple or has a very
thin (10 μm) exciple. However, specimens of A. nodulosa var. reagens from Israel have a 15-
20 μm thick exciple. Presence of rather well developed exciple (10-14 μm at the base and
expanding to 30-70 μm above) in some specimens of A. nodulosa var. reagens was also
recorded by Golubkova et al. (1978) and Golubkova (1988).

Substrates

446
Most studied Acarospora species were found on basaltic rocks. They are A. assimulans, A.
contigua, A. fuscata, A. impressula, A. laqueata, and A. umbilicata. The lichens A. areolata,
A. glaucocarpa var. cervina, and A. glaucocarpa var. conspersa were collected on calcareous
rocks. A. bornmuelleri was recorded on both substrates. A. murorum, and A. nodulosa var.
reagens occurred on fine sandy soil. A. tominiana was found on the thallus of Aspicilia
calcarea growing on limestone.
Species A. assimulans, A. bornmuelleri, and A. laqueata were found on atypical substrates.
According to literature data, A. assimulans and A. laqueata occur on calcareous rocks
(Golubkova et al. 1978, Clauzade & Roux 1981, Golubkova 1988, Nimis 1993). In the
studied territory these species were found on basaltic rocks. According to Magnusson
(1929a), A. bornmuelleri occurs on granitic or volcanic rocks. This species was also recorded
on basalt rocks in Mongolia (Golubkova 1981). In the territory of Israel this species was
found on basaltic rocks, but in the Sinai Peninsula A. bornmuelleri was recorded on
calcareous rocks (Galun & Garty 1972).
The remaining species were found on substrates, which are characteristic for them according
to literature data. It is possible to note also that species A. murorum is most usually saxicolous
and only rarely terricolous (Golubkova et al. 1978, Clauzade & Roux 1981, Nimis 1993). In
Israel this species was found on sandy soil in the Negev Desert.

Distribution in Israel and its vicinity

According to Zohary (1973), the territory of Israel and its vicinity is subdivided into four of
the following plant-geographical territories: the Mediterranean, the Irano-Turanian, the Saharo-
Arabian, and the Sudanian penetration. The biggest number of Acarospora species occurred in
the Mediterranean territory of the studied area. In that territory, nine species from this genus
were found. The number of Acarospora species recorded from the Saharo-Arabian territory are
4; from the Irano-Turanian territory 3; and from the Sudanian penetration 1.
Almost all species of the genus Acarospora are rare in the studied area. Only A. areolata
occurs more frequently. This species, described by Galun & Reichert (1960) from the Negev
Desert, has more or less wide distribution in the Judean and Negev Deserts and the Dead Sea
area. It was found also in the Sinai Peninsula (Galun & Garty 1972).
Each of four species: A. assimulans, A. bornmuelleri, A. glaucocarpa var. cervina, and A.
nodulosa var. reagens was found hitherto in three localities, but only A. bornmuelleri is more
widely distributed. It occurs in two plant-geographical territories: the Mediterranean (localities
in Upper and Lower Galilee) and Irano-Turanian (locality in Upper Jordan Valley). This

447
species was recorded also in the Saharo-Arabian Sinai Peninsula (Galun & Garty 1972).
Distribution of A. assimulans and A. glaucocarpa var. cervina is connected to the
Mediterranean territory. A. assimulans was found in the Upper Galilee and in the Golan
Heights. Three localities of A. glaucocarpa var. cervina were recorded at an altitude of 1500
m, of 1650 m, and of 2000 m above sea level in the Mount Hermon area. The distribution of
A. nodulosa var. reagens is limited to the Saharo-Arabian territory. This species occurs in the
Judean and Negev Deserts.
A. contigua, A. fuscata, A. impressula, A. laqueata, A. tominiana, and A. umbilicata are
distributed in the Mediterranean territory. Each of them (excluding A. umbilicata) was
recorded hitherto in one locality. A. contigua, A. impressula, and A. laqueata were found on
the Golan Heights. A. fuscata occurred in the Upper Galilee. A. tominiana was recorded in the
Mount Hermon area. A. umbilicata was found in two localities: in the Golan Heights and in
the Upper Galilee.
A. murorum and A. glaucocarpa var. conspersa occurred in the Saharo-Arabian territory. Both
species were found in the Central Negev, where each of them was found hitherto in one locality.

Distribution in the world

According to Wirth (1995), Acarospora fuscata is a species with wide ecological amplitude in
relation to environment conditions. This lichen has worldwide distribution.
The lichens A. glaucocarpa, A. impressula, and A. nodulosa var. reagens are also widespread
and occur almost in all continents. According to literature data, A. glaucocarpa has wide
holarctic distribution from boreal to Mediterranean regions. According to Golubkova (1983),
this lichen is an obligate calcicolous species and its distribution in various phytoclimatic
zones is limited to the presence or absence of calcareous substrates. A. impressula is an arctic-
alpine to boreal-montane species, and in Italy the distribution of this species is limited to
upland areas of South Tyrol (Nimis 1993). In Mongolia, A. impressula was also found
predominantly in mountainous regions (Golubkova 1981, 1983). In Europe this species has
distribution in the Temperate Zone (Wirth 1995). A. nodulosa var. reagens occurs in warm-
dry areas. According to Poelt (1969), A. nodulosa var. reagens is distributed widely in
southeastern Europe, but very scattered. According to Nimis (1993), the total range of A.
nodulosa var. reagens is holarctic in the Northern Hemisphere and extends to Australia.
The distribution of A. laqueata, A. murorum, and A. umbilicata is mainly in the
Mediterranean region. A. umbilicata and A. murorum have southern distribution in Europe
(Poelt 1969, Nimis 1993, Wirth 1995). These species are also known from North Africa and

448
Israel. A. laqueata occurs in southern Europe, North Africa, and in dry-warm areas of Central
Asia (Golubkova et al. 1978, Golubkova 1988, Nimis 1993).
A. assimulans, A. bornmuelleri, and A. tominiana are the Irano-Turanian lichens. Distribution
of these species is poorly known. They occur in areas with a dry continental climate and are
distributed in Asia (Golubkova et al. 1978, Golubkova 1983, 1988). According to Clauzade &
Roux (1981), A. assimulans was found also in Algeria (North Africa). The only species
known from Europe is A. tominiana. It is known from two localities in Europe. The first
locality of this species is in the southern part of European Russia (Tomin 1956, Golubkova et
al. 1978, Golubkova 1981), and another locality is in Sardinia (Italy) (Nimis 1993).
According to Brodo et al. (2001), A. contigua is probably a widespread species, but its
distribution is not properly documented because of taxonomic difficulties. Some
lichenologists include saxicolous species A. contigua in a terricolous A. schleicheri, and so
precise distribution of these species is unknown.
According to Magnusson (1929b), distribution of A. contigua is in mountainous regions of
South America (Argentina, Bolivia, Chile, Colombia) and southern North America (Arizona).
Magnusson (1940) reported also on finds of this species in China. As A. bella this lichen is
known from Africa (Morocco, South Africa) and Hawaii (Doidge 1950, Magnusson 1955,
Clauzade & Roux 1981). The taxon is poorly understood and in need of further study.
Thus, Acarospora species occurring in Israel are also widely distributed in southern Europe,
North Africa, and dry-warm areas of Asia. In Israel the distribution of species from this genus
is connected mainly to Mediterranean territory.

449
Plate 1. A - Acarospora assimulans. B - Acarospora glaucocarpa var. cervina. C -
Acarospora contigua. D - Acarospora fuscata. E - Acarospora glaucocarpa var. conspersa. F
- Acarospora laqueata.

Acknowledgements

We thank Prof. N.S. Golubkova for her help with the identification of some lichen species and Dr. Michail
Andreev for the loan of the herbarium material. We also thank Prof. Margalith Galun and Mrs. Haya Lehr for the
possibility to work with the lichenological collection of herbaria of Tel-Aviv University. We also thank Mr.
Michael Margulis for his help with the original illustrations. We would like to express our thanks to the Israeli
Ministry of Absorption and the Ancell-Teicher Research for Genetics and Molecular Evolution for financial
support of this research.

450
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