Soil and Biota of Serpentine: A World View 2009 Northeastern Naturalist 16(Special Issue 5):341–350 The Lichen Flora of Serpentine Outcrops in the Middle Urals of Russia Alexander G. Paukov* Abstract - Serpentine outcrops in the Middle Urals, Russia, supported 113 species of lichens. These rocks had a more diverse lichen flora and lower specificity (only 9 species) than other rocks types of the region. Pyroxenite's lichen flora was the most similar to that of serpentine. Rocky outcrops along different rivers varied in species richness, the highest being at the southernmost site (River Iset; 81 species) and the lowest at the northernmost site (River Neiva; 57 species). Introduction Serpentine vegetation has been the subject of detailed studies which have revealed the paucity of its flora and a high rate of endemism caused by the presence of heavy metals such as Ni, Cr, Cu, and Fe associated with the base mineral composed of hydrated magnesium silicate (Kruckeberg 1954, Proctor and Woodell 1971). The role of geochemistry and other environmental features of rocks—such as mineral composition, pH of substrate, and water availability—on lithophilous lichens have been demonstrated by several authors (e.g., Brodo 1973, Garty and Galun 1974, Purvis and Halls 1996). Lichens on serpentine have been studied at several locations in Europe and America (Favero-Longo et al. 2004, Harris et al. 2007, Sigal 1989, Sirois et al. 1988). Species diversity on serpentine is highly variable, but unlike vascular plants, lichens show a high species diversity compared to lichens on most other rocks (Favero-Longo et al. 2004, Harris et al. 2007, Sirois et al. 1988). This relatively high lichen diversity is due to chemical features which support calcareous species as well as environmental factors, such as a wide range of favorable microhabitats or a maritime influence (Harris et al. 2007). Some lichen species are regarded as serpentinophytes, and some serpentinophytic lichen communities have been described (Favero-Longo et al. 2004). The heavy metal component of the rocks which affects the diversity of lichens also gives rise to metallophytes (Purvis and James 1985). Lichens can accumulate heavy metals (Puckett et al. 1973), often in high concentrations (Brodo 1973), as in the case of nickel and copper (Nieboer et al. 1976, Purvis et al. 1987, Richardson and Nieboer 1983). However, since much of the metals are bound at extracellular metal-binding sites (Kasama et al. 2001, Nash 1989), the presence of heavy metals may not be a key factor for most lichen species in determining their occurrence on different kinds of *Biological Faculty, Urals State University, Lenin Avenue 51, 620083, Ekaterinburg, Russia; alexander_paukov@mail.ru. 342 Northeastern Naturalist Vol. 16, Special Issue 5 rocks. The interaction of substrate, microclimate, and lichens on ultramafic rocks is thus complicated and needs further investigation. The serpentine lichen flora of the Middle Urals has been studied previously (Kotlov 2003, Paukov and Trapeznikova 2005, Ryabkova 1998), but many species have been added to the list since, and no comparison with the serpentine lichen floras of other regions has been undertaken. Material and Methods The Middle Urals lie between latitudes 55º54' and 59º15' north (Chikishev 1968), and the serpentine outcrops are situated mainly along their eastern slopes, exclusively along riverbanks. Lichen biodiversity was studied on serpentine outcrops along the Rivers Iset (1, 2), Pyshma (3), Rezh (4), and Neiva (5) within the Sverdlovsk region (Fig. 1). The dominant type of serpentine is apoharzburgite, but talc-carbonate slate is found as an intrusion in the serpentine along River Iset. The territory is within the south taiga coniferous forests dominated by Pinus sylvestris L. (Scots Pine) and Picea obovata Ledeb. (Siberian Spruce), but at the southern end, the taiga forests are replaced by forest-steppe vegetation with Betula pendula Roth (European White Birch), B. pubescens Ehrh. (Downy Birch), and Populus tremula L. (Aspen). Forests and steppe patches are dominated by Helictotrichon desertorum (Less.) Nevski, Stipa pinnata L. (Feather Grass), and Echinops ruthenicus M. Bieb. Where the serpentine rocks occur, the climate varies from rather cool and wet on the northern slopes to hot and dry along the southern slopes, with summer temperatures rising above 50 ºC. About 550 specimens of lichens growing on serpentine outcrops within an area of ca. 70 ha were collected from rock, soil, and plant debris and identified using standard lichen floras (Ahti et al. 2007; Oxner 1968, 1993; Timdal and Holtan-Hartwig 1988; Wirth 1995). Lichens were deposited in the herbaria of Urals University and the Komarov Botanical Institute in St. Petersburg (LE). Results Biodiversity and distribution of lichens on serpentine outcrops The list of 113 species found on the outcrops examined, together with their distribution and abundance, is presented in Appendix 1. Spilonema revertens Nyl. and Thallinocarpon nigritellum (Lettau) P.M. Jørg. are new for the Urals. Serpentine and serpentine soils are the richest habitats in terms of lichen species diversity; limestone rocks of the same region bear 96 species, while granites (including diorites) support only 70 species. Serpentine rocks in the Middle Urals have a high number of cyanolichens (21%), compared to 8% worldwide (Sigal 1989). Despite the high lichen diversity of the sampled rocks, soil, and plant debris in the Middle Urals, the number of species restricted exclusively to serpentine rocks was low—only 9 (see Appendix 1). The reason is that 2009 A.G. Paukov 343 many lichens that grow on serpentine can also grow on other rock types; for example, limestone lichens that were also found on serpentine included Anema tumidulum, Aspicilia contorta subsp. hoffmanniana, Collema crispum, C. cristatum, C. flaccidum, C. fuscovirens, C. polycarpon, Diploschistes muscorum, Endocarpon pusillum, Mycobilimbia microcarpa, Peltigera praetextata, Phaeophyscia constipata, Ph. perisidiosa, Thallinocarpon Figure 1. A map of the Sverdlovsk region with the locations of the serpentine outcrops examined. 1 = outcrops near Dvurechensk (56°35'N, 61°03'E), 2 = Uktus mountains in Ekaterinburg (56°46'N, 60°38'E), 3 = outcrops near Staropyshminsk village (56°55'N, 60°54'E), 4 = “Sem’ Bratyev” rock near town Rezh (57°21'N, 61°21'E), and 5 = “Polyakov Kamen” rock near Melkozyorovo village (57°44'N, 61°28'E). 344 Northeastern Naturalist Vol. 16, Special Issue 5 nigritellum, and Thyrea confusa. The presence of some taxa can be attributed to the occurrence of talc-carbonate slates within the serpentine. Only a few species—Chrysothrix chlorina, Micarea erratica, Pertusaria albescens, and Porpidia cinereoatra—were found on both serpentine and granite. The species that could colonize either both serpentine and pyroxenite or serpentine and basalt were Acarospora nitrophila, Amandinea punctata, Aspicilia cf. lapponica, Caloplaca arenaria, C. holocarpa, C. subpallida, Dermatocarpon miniatum, Heterodermia speciosa, Lecidea atomaria, Lichinella stipatula, Peltigera lepidophora, Rinodina confragosa, and Xanthoparmelia conspersa. Ten species were found on all the abovementioned rocks: Candelariella vitellina, Cladonia arbuscula, C. pyxidata, Lecanora muralis, Lecidella stigmatea, Lepraria membranacea, Parmelia sulcata, Phaeophyscia sciastra, Porpidia crustulata, and Xanthoparmelia somloёnsis. A similarity dendrogram of the lithophilous lichen flora on different rocks based on the Sørensen’s coefficient is related to the classification of the minerals based on their SiO2 content. The lichens on ultramafic and basic rocks show a closer cluster that those on acidic granite or limestone (Fig. 2). Some genera, e.g., Rhizoplaca, Dimelaena, Rhizocarpon (except Rh. grande), and Trapelia, or even families, such as the Umbilicariaceae, are known to avoid serpentine in the region. The presence of lichen serpentinophytes could, with care, be used as an indicator of a rock type, as all of the serpentinophyte lichens are rare in the region. However, some may be found on other rock types after more lichen surveys in the Urals have been completed. Sixty-three species were found on serpentine both in the Middle Urals and in Europe and America. Five species that are found on serpentine at other locations are only found on limestone in the Middle Urals: Caloplaca Figure 2. Similarity dendrogram based on the Sørensen’s coefficient of lithophilous lichen flora on five rocks in the Middle Urals. 2009 A.G. Paukov 345 saxicola, Diplotomma alboatrum, Lecania erysibe, Mycobilimbia tetramera, and Placynthium nigrum. Seven serpentine species in Europe and America— Arctoparmelia centrifuga, Caloplaca crenularia, Lecidella scabra, Lepraria neglecta, Parmelia saxatilis, Polysporina simplex, and Trapelia coarctata— are granitophilous in the Middle Urals. Lichen diversity on serpentine varies according to latitudinal position of the outcrops and slope orientation. The most species-rich river was the River Iset (81 species); species numbers decreased northwards, with 70 on the River Pyshma, 62 on the River Rezh, and 57 on the River Neiva. Microhabitat diversity is the key factor in determining species richness on these serpentine rocks. Outcrops on the River Iset are rich in different habitats, from steppe patches to steep and rather wet rocks. The Neiva slopes are cooler, so xerophytic lichens such as Acarospora oligospora, Aspicilia contorta subsp. hoffmanniana, Phaeophyscia constipata, and Physconia muscigena are absent or rare. Conversely, cyanolichens, especially members of the Collemataceae, were more widely distributed along the River Neiva than the River Iset. Peltigera species were found on serpentine outcrops along all four river valleys, but Fuscopannaria was only found along the Pyshma and Rezh rivers. Slopes orientated northwards were richer in lichens (75 species), but many of them are not specifically lithophilous, such as Cladonia and Peltigera; slopes orientated southwards have 67 species, 25 of which are found on slopes of both orientations. Discussion Serpentine, as a silicate rock, provides an opportunity for lichens which prefer granite, basalts, and gabbro to colonize, and on the other hand, being enriched in calcium, it is also a substrate for calcareous species. The diversity of lichen species able to grow on serpentine, compared to vascular plants, reflects the low specificity of lichens for this particular substrate (Harris et al. 2007). This low specificity has also been observed for bryophytes (Briscoe et al. 2009) on serpentine in Maine, USA. Kruckeberg (1971) noticed the “near absence of Rhizocarpon geographicum” on serpentine rocks. This species is not widespread in the Middle Urals, but in the Southern Urals, it avoids serpentine and grows on basalt. In Canada, Umbilicaria is found on amphibolites, but not on serpentine (Sirois et al. 1988), while in California and in Europe, Umbilicaria is present on this rock (Sigal 1989, Takala and Seaward 1978). Rhizocarpon and Trapelia were common on serpentine in Europe (Favero-Longo et al. 2004) and therefore cannot be regarded as “serpentinophobous,” but Rhizoplaca, Umbilicaria, and Dimelaena do seem to avoid serpentine. The similarity dendrogram based on the Sørensen’s coefficient shows that the pH of the substratum, rather than metal availability, plays an important role in defining the affinity of lichens to rocks (Fig. 2). In the present study, nine of the lichens found only grew on serpentine in the Middle Urals 346 Northeastern Naturalist Vol. 16, Special Issue 5 (Appendix 1), but elsewhere these have been recorded from other rock types, such as limestone and basalt (Wirth 1995). The low specificity and high diversity of lichens on serpentine shows that the high heavy metal concentrations and the Mg/Ca ratio in serpentine do not impede lichen growth. These characteristics appear not to have resulted in the evolution of serpentine lichen endemics in the Middle Ural Region. To date, only a few serpentineendemic lichens are known worldwide (Brodo 1973, Wirth 1972). Acknowledgments I am very grateful to Eugene Pushkarev (Institute of Geology and Geochemistry, Russian Academy of Sciences, Ekaterinburg) for the identification of the rock types. I thank Professor M.R.D. Seaward of Bradford University for helpful suggestions on an early draft of this paper and for assistance with English and Grammar. I should like to thank Professor D.H.S. Richardson (Saint Mary's University) whose patience and comments helped me to improve the manuscript. The study was financially supported by grants from the Russian Fund of Basic Research (04-04-96131 and 07-04-96125). Literature Cited Ahti, T., P.M. Jørgensen, H. Kristinsson, R. Moberg, U. Søchting, and G. Thor (Eds.) 2007. Nordic Lichen Flora. Volume 3. Cyanolichens. Museum of Evolution, Uppsala University, Uddevalla, Sweden. 219 pp. Briscoe, L.R.E., T.B. Harris, E. Dannenberg, W. Broussard, F.C. Olday, and N. Rajakaruna. 2009. Bryophytes of adjacent serpentine and granite outcrops on the Deer Isles, Maine, USA. 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Grandtner 1988. Les lichens sur serpentine et amphibolite du plateau du mont Albert, Gaspesie, Quebéc. Canadian Journal of Botany 66:851–862. Takala, K., and M.R.D. Seaward. 1978. Lichens of the Niinivaara serpentinite region, E. Finland. Memoranda Societas Fauna Flora Fennica 54:59–63. Timdal, E., and J. Holtan-Hartwig. 1988. A preliminary key to Rhizocarpon in Scandinavia. Graphis Scripta 2:41–54. Wirth, V. 1972. Die Silikatflechten-Gemeinschaften im ausseralpinen Zentraleuropa. Dissertationes Botanicae 17:1–306. Wirth, V. 1995. Die Flechten Baden-Württembergs. Teil 1 and 2. Eugen Ulmer, Stuttgart, Germany. 348 Northeastern Naturalist Vol. 16, Special Issue 5 Appendix 1. Lichens found on serpentine outcrops at four rivers in the Middle Urals. * signifies lichens found only on serpentine outcrops; 1 = rare, 2 = occasional, 3 = common, r = on rock, s = on soil, and p = parasitic. Rivers Species Iset Pyshma Rezh Neiva Acarospora fuscata (Schrad.) Arnold 3r 3r 3r 3r Acarospora nitrophila H. Magn. 1r 1r 1r 1r Acarospora oligospora (Nyl.) Arnold* 1r Amandinea punctata (Hoffm.) Coppins & Scheid. 1r Anema tumidulum Henssen 1r Aspicilia caeciocinerea (Nyl. in Malbr.) Arnold 1r 1r 1r 1r Aspicilia cinerea (L.) Körb. 3r 3r 3r 3r Aspicilia cf. lapponica (Zahlbr.) Oxner* 1r 1r 1r 1r Aspicilia contorta ssp. hoffmanniana Ekman & Fröberg 1r Aspicilia simoёnsis Räsänen 2r Aspicilia verrucigera (Hue) Zahlbr. 2r 2r 2r 2r Bacidia herbarum (Stizenb.) Arnold* 1s Bacidia inundata (Fr.) Körb. 1r Bellemerea cupreoatra (Nyl.) Clauzade & Cl. Roux 2r 2r Caloplaca arenaria (Pers.) Müll. Arg. 1r Caloplaca flavovirescens (Wulfen) Dalla Torre & Sarnth. 2r Caloplaca grimmiae (Nyl.) H. Olivier 1p Caloplaca holocarpa (Ach.) A. E. Wade 3r 3r 3r Caloplaca lithophila H. Magn. 1r 1r 1r Caloplaca subpallida H. Magn. 3r 3r 3r 3r Candelariella vitellina (Ehrh.) Müll. Arg. 3r 3r 3r 3r Cetraria islandica (L.) Ach. 1s 1s 1s 1s Cetraria laevigata Rassad. 1s 1s 1s 1s Chromatochlamys muscorum (Fr.) H. Mayrhofer & Poelt 1s Chrysothrix chlorina (Ach.) Laundon 1r Cladonia amaurocraea (Flörke) Schaer. 1s 3s 1s 1s Cladonia arbuscula (Wallr.) Flot. 1s 1s 2s 2s Cladonia cariosa (Ach.) Spreng. 1s 1s 1s 1s Cladonia chlorophaea (Flörke) Spreng. 2s 2s 2s 2s Cladonia borealis S. Stenroos 1s 1s Cladonia coniocraea (Flörke) Spreng. 3s 3s 3s 3s Cladonia gracilis (L.) Willd. 1s Cladonia rei Schaer. 1s 1s 1s 1s Cladonia pyxidata (L.) Hoffm. 3s 3s 3s 3s Cladonia rangiferina (L.) F. H. Wigg. 1s 1s 1s Cladonia stellaris (Opiz) Pouzar & Vězda 1s 1s 1s 1s Cladonia stricta (Nyl.) Nyl. 1s Collema crispum (Hudson) Weber ex Wigg. 1r Collema cristatum (L.) F. H. Wigg. 3r, s Collema flaccidum (Ach.) Ach. 2r Collema fuscovirens (With.) J. R. Laundon 1r 1r 1r Collema polycarpon Hoffm. 1r 1r Collema tenax (Sw.) Ach. 1s Dibaeis baeomyces (L.) Rambold & Hertel 1s 2009 A.G. Paukov 349 Rivers Species Iset Pyshma Rezh Neiva Dermatocarpon miniatum (L.) W. Mann. 1r 1r 1r Diploschistes scruposus (Schreb.) Norman 1r 1r 1r 1r Endocarpon pusillum Hedw. 1s Evernia mesomorpha Nyl. 1s Flavocetraria cucullata (Bellardi) Kärnefelt et Thell 1s Fuscopannaria leucophaea (Vahl.) P. M. Jørg.* 1r Fuscopannaria praetermissa (Nyl.) P. M. Jørg.* 1r Heterodermia speciosa (Wulfen) Trevis. 1r 1r Hypogymnia physodes (L.) Nyl. 1r 1r 1r Lecanora campestris (Schaer.) Hue 2r 2r 1r 1r Lecanora dispersa (Pers.) Sommerf. 1r 1r 1r 1r Lecanora frustulosa (Disks.) Ach. 3r 3r 1r 1r Lecanora muralis (Schreb.) Rabenh. 3r 3r 3r 3r Lecanora polytropa (Hoffm.) Rabenh. 2r 2r 2r 2r Lecidea atomaria Th. Fr. 1r Lecidella carpathica Körb. 2r 2r 1r Lecidella stigmatea (Ach.) Hertel et Leuckert 3r 3r 3r 3r Lepraria incana (L.) Ach. 1r 1r 1r 1r Lepraria membranacea (Dicks.) Vain. 3s 3s 3s 3s Leptogium tenuissimum (Dicks.) Körb. 1s Lichinella stipatula Nyl. 1r 1r Melanelixia subargentifera (Nyl.) O. Blanco, A. Crespo, 1r, s 1 r, s 1 r, s Divakar, Essl., D. Hawksw. & Lumbsch Melanohalea exasperatula (Nyl.) O. Blanco, A. Crespo, 1r 1r 1r Divakar, Essl., D. Hawksw. & Lumbsch Melanohalea infumata (Nyl.) O. Blanco, A. Crespo, 1r Divakar, Essl., D. Hawksw. & Lumbsch Micarea erratica (Körb.) Hertel, Rambold & Pietschm. 1r Mycobilimbia microcarpa (Th. Fr.) Brunnb. 1s Parmelia sulcata Tayl. 1 r, s 1 r, s 1 r, s Peltigera canina (L.) Willd. 1s 1s 1s 1s Peltigera didactyla var. didactyla (With.) Laundon 1s 1s Peltigera lepidophora (Vain.) Bitter 1s 1s Peltigera malacea (Ach.) Funck. 1s Peltigera polydactylon (Neck.) Hoffm. 1 1 1 1 Peltigera ponojensis Gyeln. 1s Peltigera praetextata (Flörke ex Sommerf.) Zopf 2s Peltigera rufescens (Weiss) Humb. 2s 2s 2s 2s Peltula euploca (Ach.) Poelt * 1r 1r Pertusaria albescens (Hudson) M. Choisy & Werner 1r 1r Phaeophyscia constipata (Norrl. & Nyl.) Moberg 1s Phaeophyscia nigricans (Flörke) Moberg 1r Phaeophyscia sciastra (Ach.) Moberg 2r, s 2r, s 2r, s 2 r, s Physcia caesia (Hoffm.) Hampe 3r 3r 3r 3r Physcia dubia (Hoffm.) Lettau 3r 3r 3r 3r Physconia detersa (Nyl.) Poelt 2r, s 2r, s 2r, s 2r, s Physconia muscigena (Ach.) Poelt 1s Physconia perisidiosa (Erichsen) Moberg 1s 1s 350 Northeastern Naturalist Vol. 16, Special Issue 5 Rivers Species Iset Pyshma Rezh Neiva Placynthiella uliginosa (Schrad.) Coppins & P. James 1s Porpidia cinereoatra (Ach.) Hertel 1r 1r 1r Porpidia crustulata (Ach.) Hertel 3r 3r 3r Ramalina intermedia (Delise ex. Nyl.) Nyl. 1r 1r 1r Rhizocarpon grande (Flörke) Аrnold 1r 1r Rinodina confragosa (Ach.) Körb. 1r Rinodina milvina (Wahlenb.) Th. Fr. 1r Scoliciosporum umbrinum (Ach.) Arnold 2r 2r 2r 2r Spilonema revertens Nyl.* 1r Stereocaulon tomentosum Fr. 1r, s 1 r, s 1 r, s 1 r, s Thallinocarpon nigritellum (Lettau) P.M. Jørg. 2 r Thyrea confusa Henssen 1r Toninia cinereovirens (Schaer.) A. Massal. 1r Tuckermannopsis sepincola (Ehrh.) Hale 1r Verrucaria caerulea DC.* 1r Verrucaria glaucina Ach. 1r Verrucaria muralis Ach. 2r Verrucaria viridula (Schrad.) Ach.* 1r Xanthoparmelia camtschadalis (Ach.) Hale 1s Xanthoparmelia conspersa (Ach.) Hale 3r 3r 3r 3r Xanthoparmelia somloёnsis (Gyeln.) Hale 3r 3r 3r 3r Xanthoria elegans (Link.) Th. Fr. 2r 2r 2r 2r Xanthoria fallax (Hepp.) Arn. 1r 1r Xanthoria sorediata (Vain.) Poelt 3r 3r 3r 3r