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Non-ultramafic rock types studied. A, silica-carbonate: Clear Creek Mine just below horizon (Site 5); B, shale and sandstone: San Carlos Peak Mine (Site 9).
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Although several lichen inventories exist for European ultramafic sites, only four surveys of serpentine lichens for North America have been published to date. Of those, only one has been conducted in California. We conducted a survey of saxicolous lichens from ultramafic rocks (including nephrite, partially serpentinized peridotite, and serpentini...
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... typically hard, but can be crumbly, and generally has a coarse granular surface texture. Boulders of nephrite (very hard; granular surface texture) are distributed throughout the serpentinite mass. The New Idria serpentinite mass also contains massive in- clusions of silica-carbonate rocks, many of which con- tain cinnabar (mercury ore) deposits (Fig. 4A). Silica- carbonate rocks are typically hard with a vacuolar porous surface texture and have a dominant mineral com- position of quartz, chalcedony, opal, ankerite, magnesite, and dolomite (Van Baalen 1995). Silica-carbonate forms from the precipitation of minerals from hydrothermal fluids of ultramafic origin within the serpentinite ...
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... Terricolous lichens cover more than 6% of the land surface of the earth [23], and are found in areas such as tropical forests, desert, alpine regions, polar regions, urban areas, as well as highly polluted areas [26,27]. Lichens found in naturally heavy metal–enriched sites, such as serpentinite and other ultramafic rocks [28][29][30]and metal mines [31,32], may be good model systems for biomonitoring. Accordingly, lichens may have the potential to be broadly used as biomonitors in various terrestrial habitats worldwide. ...
Levels of trace element pollution in surface soil can be estimated using soil analyses and leaching tests. These methods may reveal different results due to the effect of soil properties, such as grain size and mineral composition, on elemental availability. Therefore, this study advocates an alternative method for monitoring and assessment of trace element pollution in surface soil using terricolous fruticose lichens. Lichens growing at abandoned mine sites and unpolluted areas in southwest Japan and their substrata were analyzed using inductively coupled plasma-mass spectrometry and X-ray fluorescence spectrometry to clarify the relationships between Cu, Zn, As, and Pb concentrations in lichens and soils, including their absorption properties. Concentrations of these elements in the lichens were positively correlated with those in the soils regardless of lichen species, location, habitat, or conditions of soils. The analyzed lichens had neither competitive nor antagonistic properties in their elemental absorption, which made them good biomonitors of trace element pollution in surface soil. The distribution maps of average Cu, Zn, As, and Pb concentrations at each sampling region detected almost all of the Cu, Zn, and As pollution of the soils. Therefore, lichens could be used in practical applications to monitor Cu, Zn, and As pollution in surface soils.
... Rock roughness and area varied independently of the grassland type, showing that both studied areas do not vary in substrate availability for lichens to establish and grow. Some authors have found that rock surface structure can determine saxicolous lichen community structure (Rajakaruna et al. 2012). Since the studied sites had both smaller and structurally less complex rocks and also bigger rocks with more heterogeneous surface, we did not note this influence. ...
Background and aims – The use of fire as a managing tool influences and maintains many types of vegetation and may determine landscape physiognomy and species composition. Fire has a strong effect on lichens but studies about their recolonization or resistance are rare. Considering this, the objectives of this study are to investigate how fire impacts the structure of saxicolous lichen communities on rocks in grassland, considering richness, cover and lichen taxa composition and to verify if there are changes on functional diversity and redundancy among sites with different fire events. Methods – The study was undertaken in an area composed of mosaics of native Araucaria forests and grasslands, in southern Brazil, where two sites were sampled. One of them experienced a recent intense fire episode, after being fifteen years without fire or grazing influence, and the other was regularly subjected to grazing and annual fire episodes. Key results – The sites differed in lichen cover, lichen taxa richness and composition. Regarding the environmental and structural variables of the surrounding vegetation, significant influence on saxicolous lichen communities was also observed, with taller vegetation and greater vegetation cover on the rocky outcrops from the site with only one and intense recent fire event. Functional diversity and functional redundancy was also significantly different between both sites. Conclusion – Based on these results it is possible to affirm that managing southern Brazilian natural grasslands with annual fire and grazing, unless it is not too intense, does not prevent the development of saxicolous lichen communities. The effect of fire on abandoned grasslands, with a high quantity of organic matter and taller vegetation, is importantly negative to saxicolous lichen community. Lichens take more time to grow and occupy the rock than the surrounding vegetation recovers, which can cause microclimate changes. The recolonization of these rocky outcrops can therefore take a long time and the new lichen community will probably never be the same as the former one. © 2015 Botanic Garden Meise and Royal Botanical Society of Belgium.
... Therefore, it is hypothesized that the element composition will correlate with the thallus cover and sexual reproduction of lichen species. Lichen species and assemblages have already been correlated with rock type (Barnett and Lilly 1966; Zonneveld 1975; Bates 1978; Clayden and Bouchard 1983; Hauck et al. 2001; Giordani et al. 2002; Favero-Longo et al. 2004; Purvis and Pawlik-Skowron´skaSkowron´Skowron´ska 2008; Rajakaruna et al. 2012; Spitale and Nascimbene 2012). In a previous study by , community types were named after the most prevalent vegetation and two Xanthoparmelia species (Xanthoparmelia cumberlandia (Gyelnik) Hale, and X. viriduloumbrina (Gyelnik) Lendemer) occurred in all community types, whereas Arctoparmelia centrifuga (L.) Hale was limited almost exclusively to a single community . ...
Substratum plays an important role in saxicolous lichen communities, providing a stable growth surface for long-term attachment and influencing the relative abundance of species. Saxicolous lichen communities consisting of Xanthoparmelia and Arctoparmelia were previously characterized using thallus cover on the substratum as well as lichen fecundity, but it was not known whether the substratum influenced lichen fecundity. The objectives of this paper were to compare variation in element concentrations among lichen communities, and to examine the correlation between elemental composition and percent cover and fecundity (numbers of apothecia, ascospores, and percent germination) of each of three lichens Arctoparmelia centrifuga, Xanthoparmelia viriduloumbrina, and X. cumberlandia. Percent cover and fecundity features were compared with the rock type and 21 elements from 37 transects. Element concentrations varied among communities, but the grassy rock community had significantly higher concentrations of elements than the mossy or treed rock communities. Positive correlations were present between fecundity and particular elements for the three species. This study provides insight into the complexity of the relationship between the geological composition of the substratum and saxicolous lichen biology. © 2015, National Research Council of Canada. All rights reserved.
... Kruckeberg (2002) proposed that the microbial biota of serpentine soil would be species poor, paralleling the scanty vegetation, but admitted that very little is known about the microbes of serpentine soils. Since Kruckeberg (2002), numerous studies have documented the diversity of bacteria (Oline 2006; Rajkumar et al. 2008), fungi (DeGrood et al. 2005; Daghino et al. 2012; Southworth et al. 2014) and lichens (Favero-Longo et al. 2004; Rajakaruna et al. 2012) in serpentine soil, yet investigations of algae and cyanoprokaryotes in ultramafic soil (Terlizzi and Karlander 1979; Couté et al. 1999; Hauer 2008) are still scarce. The aim of the present study was to conduct a preliminary survey of the algal and cyanoprokaryote composition in mafic-and ultramaficderived soil to improve our understanding of the microbial diversity of serpentine and related soils. ...
Despite a large body of work on the serpentine-substrate effect on vascular plants, little work has been undertaken to describe algal communities found on serpentine soils derived from peridotite and other ultramafic rocks. We report a preliminary study describing the occurrence of algae and cyanoprokaryotes on mafic and ultramafic substrates from South Africa. Results suggest that slope and aspect play a key role in species diversity and community composition and, although low pH, nutrients and metal content do not reduce species richness, these edaphic features influence species composition. Further, typical soil genera such as Leptolyngbya, Microcoleus, Phormidium, Chlamydomonas, Chlorococcum and Hantzchia were found at most sites. Chroococcus sp., Scytonema ocellatum, Nostoc linckia, Chlorotetraedron sp., Hormotilopsis gelatinosa, Klebsormidium flaccidium, Pleurococcus sp. and Tetracystis elliptica were unique to one serpentine site. The preliminary survey provides directions for future research on the serpentine-substrate effect on algal and cyanoprokaryote diversity in South Africa.
... Soil is an intermediary between bedrock and vegetation. The chemical composition of this biologically important layer is largely determined by the composition of underlying parent materials (Rajakaruna and Boyd 2008). Soil composition is a significant factor in plant ecology and evolution: various plant species have evolved specific traits, such as the ability to tolerate heavy metals or nutrient imbalances, as a result of natural selection in populations on particular soils (Kruckeberg 1986;O'Dell and Rajakaruna 2011). ...
... A brief note concerning terminology is warranted here. Although ''gabbroic soils'' would be a better term grammatically, we use the phrase ''gabbro soils'' here to be consistent with other geoecological literature, in which ''serpentine soils'' is the accepted term for soils derived from serpentinite and other related rocks (see Rajakaruna et al. 2009). ...
... No comprehensive work has been done on the lichens of gabbro outcrops in California. As serpentine rocks in the CFP have been shown to support unique lichen assemblages (Rajakaruna et al. 2012), a study of gabbro lichens could be worthwhile. Comparisons should also be made between mycorrhizal communities on gabbro and other unique substrates, such as serpentine (Southworth et al. 2014). ...
Gabbro is a mafic plutonic rock that is common in the Sierra Nevada, Klamath
Mountains, and Peninsular Ranges of the California Floristic Province. Its mineralogical
and chemical compositions span the range between those of peridotite, an ultramafic
rock, and diorite, a rock more silicic than gabbro. A broad range of nutrient element
compositions makes "gabbro soils" extremely diverse substrates for plants; it is not
surprising that they harbor numerous rare and endemic plant species, particularly at
the Pine Hill complex in El Dorado County, California. These rare and endemic species
include Ceanothus roderickii (Rhamnaceae), Galium californicum subsp. sierrae
(Rubiaceae), Wyethia reticulata (Asteraceae), and Calystegia stebbinsii
(Convolvulaceae). More research is required to discern the factors affecting endemism
and plant distributions on gabbro soils. Attention should be paid to vegetational
differences between olivine gabbro, which chemically borders serpentine, and gabbro
lacking olivine or containing more hornblende than olivine.
... In contrast, in the case of Area B, the occurrence of species restricted to one of either the ultramafic or mafic substrates corroborates reports by several authors suggesting L Australian Journal of Botany S. E. Favero-Longo et al. that lichen communities on serpentinites may have unique species assemblages relative to those of other lithologies (e.g. Sirois et al. 1988; Rajakaruna et al. 2012). However, the fact that lichen communities on the ultramafic substrate Ser B displayed higher similarity with those on the mafic substrate MMg relative to those on the ultramafic substrates of Area A suggests that environmental factors other than substrate may prevail in selecting lichen functional traits and driving the community composition. ...
... Although rock chemistry together with texture and moisture capacity, has long been suggested as the main factor driving the diversity of saxicolous lichen communities on different lithologies (Brodo 1973), studies which correlate rock mineralogy and chemistry with lichen communities are rare (Rajakaruna et al. 2012). Rajakaruna et al. 2012 correlated lichen community composition on ultramafic and nonultramafic rocks with XRF chemical analyses directly performed on the rock substrates, but only a weak correlation was detected. ...
... Although rock chemistry together with texture and moisture capacity, has long been suggested as the main factor driving the diversity of saxicolous lichen communities on different lithologies (Brodo 1973), studies which correlate rock mineralogy and chemistry with lichen communities are rare (Rajakaruna et al. 2012). Rajakaruna et al. 2012 correlated lichen community composition on ultramafic and nonultramafic rocks with XRF chemical analyses directly performed on the rock substrates, but only a weak correlation was detected. However, the report did not perform XRF analyses on lichen thalli and high levels of metals in the rock substrates can only be considered a proxy me ...
There is a long history of studies on lichens found in ultramafic habitats, but comparisons between lichen communities on different ultramafic lithologies are scant, and potential metabolic adaptations to the multiple edaphic stresses of ultramafic substrates have been widely neglected. The present work is the first to characterise differences in the abundance and structure of saxicolous lichen communities on different ultramafic lithologies (dunite, lherzolite, and serpentinite), analysed in two areas of the Western Alps (NW Italy). Differences between communities on various ultramafic lithologies were observed, including differences between a mafic control (Mg-Al metagabbro); however, factors other than the substrate were observed to drive more remarkable differences between lichen communities on ultramafics of alpine and pre-alpine areas. XRF analyses demonstrated that the mineral composition of different lithologies is reflected by metal contents in crustose lichens, with weathering processes accounting for relative shifts in elemental abundances between rocks and thalli. A thin layer cromatography screening of lichen secondary metabolites (LSMs), which are thought to regulate metal and pH homeostasis in thalli, revealed lithological vicariance among dominant lichen species with different LSM patterns and intraspecific variability in LSM production associated with differences in lithology and location. In particular, the presence or absence of norstictic acid in species or lineages/individuals on the different lithologies, in relationship to concentrations of Fe, Mg, and Ni in lichen thalli, was recognised as a metabolic adaptation to metal stress. Pull-up tests revealed that physical factors such as a differential surface disaggregation may contribute more towards differences observed in lichen abundance on the different lithologies investigated.
... Lichens on serpentinite have attracted the attention of lichenologists for several decades and have been studied in many locations in Europe and America. One of the features of serpentinite lichen flora is its impoverished nature (Wirth 1972), but many recent studies have characterised it as species-rich (Takala and Seaward 1978; Sirois et al. 1988; Sigal 1989; Favero-Longo et al. 2004; Harris et al. 2007; Paukov 2009; Rajakaruna et al. 2011 Rajakaruna et al. , 2012), primarily because of serpentine harboring many species growing on acid or calcareous rocks (Gilbert and James 1987; Purvis and Halls 1996). In contrast to limestone, another species-rich rock, serpentinites, contain substantial quantities of metals such as Ni, Cr, Cu and Fe (Kruckeberg 1954; Proctor and Woodell 1971) and tolerance to them as well as ultrabasic nature of serpentinite may be another reason of high diversity of lichens on this substrate. ...
Accumulation of metals in four crustose lichens with different secondary chemistry growing on serpentinite was studied. Aspicilia cinerea and A. blastidiata contain depsidone norstictic acid, A. goettweigensis contains stictic acid, and A. contorta ssp. hoffmanniana contains aliphatic compound aspicilin. The highest concentrations in lichens compared to serpentinite were found for Ca (average – 11 times, maximum – 20 times). Sr, Cu, Na, Zn and Cr were 2.8–9 times greater in lichens compared to rocks and other elements such as Ni, Fe, Co, Mn, and Mg were equal or lower in the thalli compared to the substrate. Three species showed little differences in concentrations of the same metals but Aspicilia blastidiata which is obligate to serpentinite had statistically higher levels of most elements. This implies that difference in secondary chemistry does not strongly influence accumulation rates of metals in selected species on serpentinite but that lichens have both mechanisms of accumulation and avoidance that may be related to ‘lichen acids’.
Aspicilia blastidiata Paukov, A. Nordin & Tibell is described as new to science.
... These effects of serpentine on bryophyte assemblages in Californian grasslands resemble the effects seen in other regions (Briscoe et al. 2009; Cleavitt et al. 2009; Rajakaruna et al. 2009). Unlike vascular plants (Kruckeberg 2002) or saxicolous lichens (Rajakaruna et al. 2012), bryophyte species and assemblages do not appear strongly specialized on serpentine in our data (see also Lepp 2001 ). Serpentine substrates were mainly occupied by widespread epigaeic or epilithic species that are found on a variety of different non-acid bedrock types. ...
Bryophyte communities of semi-arid grassland ecosystems are little studied,
and it is poorly known how they vary in relation to soil quality, disturbance and the
invasion status of vascular plants. By analogy with short-statured, competitively inferior
native vascular plants, we hypothesized that bryophytes would show high beta diversity
and increasing alpha diversity along a gradient from productive non-serpentine grasslands
dominated by exotics to low productivity serpentine grasslands dominated by native
vascular plants. We also predicted that disturbance by gophers would benefit bryophyte
alpha and beta diversity, especially at the productive end of the soil fertility gradient. We
sampled bryophytes in 48 sites in a 3,000 ha landscape in the Inner Coast Range of
California, USA. We used several multivariate (non-metric multidimensional scaling,
PERMDISP) and univariate analysis methods (generalized linear models). As predicted,
we found high beta diversity of bryophytes along the soil gradient, and higher cover and
richness in unproductive rocky serpentine grasslands than productive and exotic-dominated
non-serpentine grasslands. Gopher burrowing had a unimodal influence on bryophyte alpha
diversity. Our results show that bryophyte species composition and diversity respond
strongly to the same soil gradient that controls vascular grassland species. Bryophytes are
likely imperiled by the continued spread of exotic vascular plants, but promoted by
moderate disturbances such as gopher burrowing.
... Calcium increased the most after the addition of BC to serpentine soil. Given that Ca is often considered to be the most limiting nutrient for plant growth in serpentine soil (O'Dell and Rajakaruna 2011; Rajakaruna et al. 2012 ), this observed increase in response to BC application is worthy of note. The available amounts of K, Ca, and Mg in the 5 % BC amendment were multiplied by 1.2–2.0-fold ...
Purpose: Serpentine soils derived from ultramafic rocks release elevated concentrations of toxic heavy metals into the environment. Hence, crop plants cultivated in or adjacent to serpentine soil may experience reduced growth due to phytotoxicity as well as accumulate toxic heavy metals in edible tissues. We investigated the potential of biochar (BC), a waste byproduct of bioenergy industry in Sri Lanka, as a soil amendment to immobilize Ni, Cr, and Mn in serpentine soil and minimize their phytotoxicity.
... These areas can be anthropogenic in origin, such as mines where waste accumulation on the surface has dramatically changed the landscape, or naturally occurring, such as outcrops of ultramafic rock. Decades of research have suggested that these ''edaphic islands'' may harbor a high proportion of endemic plants that have evolved to tolerate edaphic stresses (Kay et al. 2011;Rajakaruna and Boyd 2008). Often-quoted examples include the flora of the island of New Caledonia, which has abundant ultramafic rock outcrops and far more endemic species than many larger islands (Anacker 2011), and California, which supports a high number of species endemic to specific geologies, including ultramafic rocks such as serpentinite (Kay et al. 2011;Safford et al. 2005). ...
... 116 2006;O'Dell and Rajakaruna 2011), providing opportunities for the study of ecological speciation (Kay et al. 2011). Compared to vascular plants, lichens on serpentine outcrops have received less attention (Favero-Longo et al. 2004;Rajakaruna et al. 2012) and, as with vascular plants, very little of the research has been carried out in eastern North America (Harris et al. 2007;Rajakaruna et al. 2009;Reed 1986;Sirois et al. 1988). This is somewhat surprising considering that New England has received extensive lichenological study, including investigations of sites with unique geologies (see Miller et al. 2005). ...
... 116 2006;O'Dell and Rajakaruna 2011), providing opportunities for the study of ecological speciation (Kay et al. 2011). Compared to vascular plants, lichens on serpentine outcrops have received less attention (Favero-Longo et al. 2004;Rajakaruna et al. 2012) and, as with vascular plants, very little of the research has been carried out in eastern North America (Harris et al. 2007;Rajakaruna et al. 2009;Reed 1986;Sirois et al. 1988). This is somewhat surprising considering that New England has received extensive lichenological study, including investigations of sites with unique geologies (see Miller et al. 2005). ...
Geochemistry and mineralogy of rocks play important roles in the occurrence of individual lichen species and assembly of lichen communities. Whereas lichens of metal-enriched settings have been a focus of study for many decades, only a few such lichen inventories exist for North America. We reexamined the lichen biota of Pine Hill, a serpentine outcrop on Little Deer Isle, Maine and Callahan Mine, a copper- and zinc-enriched Superfund site in Brooksville, Maine, by conducting additional field surveys and reexamining unidentified taxa from previous collections. To better characterize the substrates upon which the lichens were found, we conducted elemental analyses via x-ray fluorescence and inductively coupled plasma-mass spectrometry on rock samples collected at Pine Hill and recorded pH, electrical conductivity, and elemental concentrations via inductively coupled plasma mass spectrometry on soil samples from Callahan Mine. The re-investigation of lichens of the two metal-enriched sites resulted in the addition of 20 taxa to Pine Hill and 10 taxa to Callahan Mine. These include Dermatocarpon leptophyllodes, Placynthiella hyporhoda, Pyrenocarpon thelostomum, and Vezdaea acicularis, all recorded for the first time from New England. In addition, we report the first documented records since the late 19th to early 20th century for New England of Porocyphus coccodes, Sarcosagium campestre, and Steinia geophana, and the first such record for Maine for Coccocarpia palmicola. Stereocaulon condensatum and S. subcoralloides, both considered rare in New England, were also collected from Callahan Mine.
