A peer-reviewed open-access journal BioRisk 3: 127—136 (2009) doi: 10.3897 /biorisk.3.25 & BioRisk www.pensoftonline.net/biorisk Biodiversity & Ecosystem Risk Assessment Lichens of Israel: diversity, ecology, and distribution Marina Temina, Eviatar Nevo Institute of Evolution, University of Haifa, Israel Corresponding author: Marina Temina (temina@research.haifa.ac.il) Academic editors: L.J. Musselman, FE Krupp | Received 15 March 2009 | Accepted 16 December 2009 | Published 28 December 2009 Citation: Temina M, Nevo E (2009) Lichens of Israel: diversity, ecology, and distribution. In: Krupp FE, Musselman LJ, Kotb MMA, Weidig I (Eds) Environment, Biodiversity and Conservation in the Middle East. Proceedings of the First Middle Eastern Biodiversity Congress, Aqaba, Jordan, 20-23 October 2008. BioRisk 3: 127-136. doi: 10.3897/ biorisk.3.25 Abstract The biota of lichen-forming and lichenicolous fungi of different plant-geographical regions of Israel was analyzed. These areas differ in climatic conditions. A total of 350 species from 16 orders, 52 families, and 117 genera were recorded; among them 5% are endemic to the Levant. The highest species richness was found in the Mediterranean area. Species diversity of other areas with more arid climatic conditions were much poorer in comparison with species diversity found in the more humid and cooler Mediter- ranean region. Saxicolous lichens were the most common species. However, in the Sudanian penetration area, terricolous lichens dominated. Among phytogeographic elements, temperate species were dominant. However, the Sudanian penetration area was characterized by the dominance of Mediterranean and sub- continental species. Most of the lichens studied were xerophytic and photophytic species. They dominated all plant-geographical areas of Israel. In the Mediterranean region, mesophytic lichens were also common. Keywords Ecology of lichens, lichen diversity, Israel, species composition Introduction The first lichens in the Mediterranean region were mentioned in the work of Miiller Argo (1884). His paper included information on lichen species collected in the Negev desert. Later, lichenological investigations of this region were continued in the 1940s by I. Reichert, who published several papers on steppe and desert lichens (Reichert 1937a, b; 1940). He donated his unique lichen collection comprising specimens from Leba- Copyright M.Temina, E. Nevo. This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. 128 Marina Temina & Eviatar Nevo / BioRisk 3: 127-136 (2009) non, Syria, Jordan, Egypt, and Israel along with his lichenological library to the Depart- ment of Botany, Tel Aviv University. The collection and library served as the basis for lichenological studies of Israel, which were continued by M. Galun and her students. Among Reichert’s students, the famous lichenologist Margalith Galun can be rightly considered as the founder of a new era in the history of Israeli lichenology. In 1958 she, together with Reichert, published her first paper about the lichen flora on olive trees (Reichert & Galun 1958). In the 1960s she started thorough floristic stud- ies of all regions of the country. Her well known handbook, “The Lichens of Israel” (Galun 1970), was the first summary of lichen floristic explorations in Israel, which stimulated further lichenological investigations in the country. Her extensive studies including lichen diversity, lichen biology, lichen ecology, and lichen systematics dur- ing a period of 40 years extended the body of knowledge available in many fields of lichenology not only in Israel, but world-wide. From 1969 until 1995, many scientists contributed to the knowledge of Israeli lichens. Among these contributions are the experimental works on photosynthetic and respiratory activity of desert lichens in the central Negev by the German scientist O. Lange and his colleagues (Lange et al. 1977). Lange also was engaged by the lichenom- etric studies in the Negev desert (Lange 1990). A number of interesting studies on the influence of air pollution on various characteristics (ecological, physiological, heavy metal content, etc.) of different lichen species were made by J. Garti and his students and colleagues. In the 1980s and 1990s, many investigations were devoted to the ques- tion of biogenous weathering of rocks by lichens. They were carried out by Danin and other scientists. On the basis of these studies, Danin proposed to use the patterns of biogenic weathering as indicators of paleoclimates in Israel (Danin 1985, 1986). Since 1996, intensive lichenological investigations have been carried out by the Institute of Evolution, University of Haifa, in collaboration with colleagues from other institutes in Israel and abroad. These investigations are based on past achievements and continue up to the present time. They touch on the different fields of lichenology including lichen biodiversity, taxonomy, ecology, biochemistry, genetics, etc. The present study represents a short review of lichens collected in Israel up to 2008. The following characteristics of lichen biota were studied: Systematics, species richness, species composition in different regions of Israel, phytogeographical elements, and ecological peculiarities. Materials and methods The paper is based on the results of expeditions to various regions of Israel from 2000 to 2007, and data collated from literature sources (Galun 1970, Galun & Mukhtar 1996, Temina et al. 2005). Lichen specimens were identified in the Laboratory of Lichenology at the Institute of Evolution, University of Haifa (Israel), applying stand- ard methods. The following references were used to identify the specimens: Galun 1970, Clauzade & Roux 1985, Purvis et al. 1992, Wirth 1995, Boqueras 2000, and Lichens of Israel: diversity, ecology, and distribution 129) Temina et al. 2005. Lichen nomenclature follows Temina et al. (2005). Specimens were deposited in the herbaria of the Institute of Evolution, University of Haifa, Israel. To estimate similarity between lichen vegetations of different regions, the Sorensen index (Mueller-Dombois & Ellenberg 1974) was calculated. To analyze local commu- nity organization, complexes of lichen species were subdivided into groups based on phytogeographical and ecological characteristics for each species according to Wirth (1995), Temina et al. (2005), and Nimis & Martellos (2008). The contribution of each group to the local lichen community was estimated on the basis of relative frequency (Mueller-Dombois & Ellenberg 1974). Results and discussion Species diversity The biota of lichen-forming and lichenicolous fungi of Israel includes 350 species from 16 orders, 52 families, and 117 genera (Table 1). Among the various families, the high- est species diversity is represented by the Teloschistaceae (55 species), Physciaceae (36 species), Lecanoraceae (28 species), and Verrucariaceae (26 species). The dominance of these families in species diversity shows a similarity of Israeli lichenobiota and licheno- biotas of the Mediterranean and Irano-Turanian regions. Table |. Systematic diversity of lichen-forming and lichenicolous fungi of Israel. Number of taxa one Family Genera Species Agyriales 1 1 Arthoniales 3 8 24 Capnodiales 1 1 Dothideales 2 2 2 Lecanorales 20 53 L75 Lichinales 4 10 25 Ostropales 3 10 Peltigerales 3 4 Pertusariales 1 2 8 Phyllachorales 1 1 1 Pleosporales 3 3 3 Pyrenulales 2 2 Teloschistales 5) 56 Trichotheliales 1 1 3 Verrucariales 2 14 27 Order uncertan 3 4 6 Mitosporic fungi ——e 4 4 Total 117 350 130 Marina Temina & Eviatar Nevo / BioRisk 3: 127-136 (2009) Species composition of different regions of Israel Israel is climatically and geographically very variable as indicated by the presence of 29 geo-climatic regions (Rubin & Pick 1994). This extensive climatic variability results from the presence of the mesic/desert border and from the widening of the Saharo- Syrian aridity belt (especially during the Pleistocene and Holocene, Tchernov 1975). Four plant-geographical regions are represented in Israel (Zohary 1973): The Mediter- ranean, the Irano-Turanian, the Saharo-Arabian, and the Sudanian penetration (Fig. 1). The climate in these regions varies from mesic in the Mediterranean area to semi-arid in the Irano-Turanian area, to arid in the Saharo-Arabian area and to extremely arid in the Sudanian penetration area. The largest number of lichens occurred in the Mediterranean area (Fig. 2), which is not surprising, as this area is ecologically more heterogeneous, represents a spatio-tem- porally “broader niche” (Van Valen 1965) and has more habitat patches and subdivisions than other regions of Israel. Analysis of the Sorensen Indices of Similarity (Table 2) showed significant similarity between lichen biota of the Saharo-Arabian and the Irano- Figure |. Plant-geographical regions of Israel. Abbreviations: M Mediterranean region S-A\ Saharo- Arabian region I-T Irano-Turanian region S Sudanian region. Lichens of Israel: diversity, ecology, and distribution 131 Mediteranean territory 87% ra The Saharo-Arabian territory 25% 19% x The Sudanian The Irano-Turanian territory territory Figure 2. The species richness of lichens in different plant-geographical regions of Israel. Turanian regions, apparently reflecting a relatively high degree of similarity of the cli- matic and biotic conditions of these regions. The frequencies of 60 basic species in the different plant-geographical regions are shown in Table 3. Only seven species were recorded in all regions. Caloplaca aurantia, C. citrina, C. holocarpa, Placidium squa- mulosum, Psora decipiens, and Toninia sedifolia are the most common lichens in Israel. Substrates Most lichens occurring in Israel are saxicolous species (Fig. 3A); among them, calci- colous lichens dominate. This may be attributed to the prevalence of limestone sub- strates in Israel. It is also possible to note the high presence of parasitic species in lichen biota. This is a characteristic feature of lichens of the Mediterranean region, which can perhaps be linked to the scarcity of free-living photobionts in this area. The Mediterranean territory is characterized by the dominance of calcicolous and corticolous lichens (Fig. 3B). In the Irano-Turanian and Saharo-Arabian regions, calci- colous species prevail in lichen biota. The terricolous and substrate-indifferent lichens are the dominant species in the Sudanian penetration area, which probably reflects the lack of dew precipitation in this region. Table 2. Sorensen index of similarity of lichen vegetation in different regions of Israel. Mediterranean Saharo-Arabian Irano-Turanian region Saharo-Arabian region Irano-Turanian region Sudanian penetration region 16 132 Marina Temina & Eviatar Nevo / BioRisk 3: 127-136 (2009) Table 3. The frequencies of basic lichen species in different plant-geographical regions of Israel; (vr - very rare; r— rare; rc — rather common; c — common; ve — very common). Mediterranean | Saharo- Irano- Sudanian ; region Arabian Turanian | penetration opeics region region region Commonness-rarity status Acarospora areolata - c rc vr Aspicilia desertorum r c c . Aspicilia calcarea c r r - Aspicilia farinosa rc rc rc - Bacidina phacodes rc - . . Bagliettoa parmigera c - - - Buellia sorediosa r rc r . Buellia subalbula vax. fuscocapitellata - rc r - Caloplaca alociza rc G rc - Caloplaca arenaria rc - Caloplaca aurantia c - Caloplaca circumalbata var. bicolor c - Caloplaca circumalbata var. circumalbata c . Caloplaca citrina c - Caloplaca erythrocarpa - - Caloplaca holocarpa vc c c - Caloplaca lactea c r r - Caloplaca latzelii rc - - . Caloplaca saxicola rc - - . Caloplaca variabilis c r r - Caloplaca velana var. velana - - Candelariella minuta - c 6 - Cladonia convoluta c - - - Cladonia pocillum c - - . Collema crispum rc rc ie r Collema cristatum c vr r . Collema tenax ¢ rc rc r Diploicia canescens rc r ; - Diploschistes candidissimus rc ive tC - Diploschistes diacapsis - rc c . Diplotomma epipolium r c c . Diplotomma venustum - rc rc - Lecania naegelii ¢ - - - Lecanora crenulata r rc rc - Lecanora pruinosa c i r - Lecidella euphorea c - - - Lichens of Israel: diversity, ecology, and distribution 133 Species Lichinella sinaica Lobothallia radiosa Mediterranean region Saharo- Irano- Arabian Turanian region region Sudanian penetration region Commonness-rarity status r - Neofuscelia pulla Peltula obscurans Peltula patellata. Physcia adscendens Physconia venusta Placidium squamulosum Psora decipiens Ramalina lacera Ramalina maciformis Rinodina bischoffii var. aegyptiaca Rinodina dubyana Squamarina cartilaginea vat. cartilaginea Squamarina cartilaginea vat. pseudocrassa r Squamarina lentigera [Cc Cc Teloschistes lacunosus Toninia sedifolia Tornabea scutellifera Verrucaria fuscella Verrucaria marmorea Xanthoria calcicola Xanthoria mediterranea Xanthoria parietina Geographic distribution Most lichens recorded in Israel are globally widely distributed and occur in all or al- most all continents. However, about one-third of them are rare in the rest of the world; among them, 5% are endemic to the Levant. Lichens found in Israel belong to eight phytogeographical categories: temperate, northern temperate, southern temperate, sub-oceanic, Mediterranean, subcontinental, arid, and endemic. The temperate spe- cies dominate in the lichens of Israel. The other phytogeographical elements are much less common than the temperate species. Widespread temperate species prevail in the Mediterranean, Irano-Turanian, and Saharo-Arabian regions. The Sudanian penetra- tion area is characterized by the dominance of Mediterranean and sub-continental lichens. The high abundances of endemic species in the Irano-Turanian and Saharo- Arabian regions are noteworthy. 134 Marina Temina & Eviatar Nevo / BioRisk 3: 127-136 (2009) Saxicolous A 50% Cortucolous 25% ZS ae ~~ Terricolous S% 11% 9% Parasitic Substrate-indifferent B 100% ate —— 80% |” v Ms 60% ~ SEEEeDG § 40% - 50%: 0% + Med S-Arab Ir- Tur Sud E] On calcareous rocks OOn basalt M™ On trees On soil On different substrates | [On other lichen species Figure 3. The frequencies of lichens on different substrates: A In all areas of Israel B in certain plant-ge- ographical regions of Israel. Abbreviations: Med - Mediterranean region, S-Arab - Saharo-Arabian region, Ir-Tur - Irano-Turanian region, Sud - Sudanian region. Ecology of the lichen species studied The main factors influencing the distribution of lichens in arid areas are light and moisture conditions. As Fig. 4 shows, most lichens found in Israel represent xerophytic and photophytic species. The same groups of species dominate in all plant-geograph- ical regions. However, in the Mediterranean region, high abundances of mesophytic and hygrophytic species were also observed. It is important to note that in this region the frequency of photophytic species is lower than the frequency of these species in other plant-geographical areas with higher solar radiation. Lichens of Israel: diversity, ecology, and distribution 135 Moisture conditions Xerophytic 55% Mesophytic r ae hy. Hygrophytic 14% Indifferent Light conditions eae Hoe Photophytic 2% | 9% ~~ 16% Skiophytic M photophytic Indifferent Figure 4. 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