Microbiotic Soil Crusts: Structure and Function
Larry L. St. Clair
Professor of Botany &
Curator of Nonvascular Cryptogams
Brigham Young University
Provo, Utah 84601
In many arid and semiarid areas of the world, soil surfaces are consolidated by complex communities consisting of microorganisms, lichens, and bryophytes. The physical structure of microbiotic crusts varies depending on climate, soil type, and the composition of the biological community. Some crusts are flattened, polygonally cracked, and have a rough, undulating surface; while others are pedicellate and demonstrate a complex vertical microtopography. All soil crust communities contain some combination of cyanobacteria, bacteria, eukaryotic algae, and non-lichenized fungi. More structurally complex soil crusts also contain some combination of lichens and bryophytes.
Biological soil crusts have been known by a variety of names. Fletcher and Martin (1948) first used the term raincrust, but due to the fact that there are raincrusts of nonbiotic origin the term was later abandoned. Others have designated the crust by its dominant life form, i.e., algal crust, lichen crust, or moss crust. Kleiner and Harper (1972) coined the term cryptogamic soil crust, a term that has been widely used over the last 25 years. More recently, other suggestions have been made, including microphytic crust (West 1990), cryptobiotic crust (Belnap 1993), and microbiotic crust (St. Clair and Johansen 1993).
A growing body of data suggests that microbiotic crusts play several important ecological roles in arid and semiarid ecosystems. The most obvious, and likely the most important, role is stabilization of soil surfaces, which effectively controls and reduces erosion (Blackburn 1975, MacKenzie and Pearson 1979, Johansen et al. 1998). Even though the data is somewhat contradictory, there are some cases where seed germination and seedling establishment appear to be enhanced by soil crust development, especially in moist sites associated with the complex microtopography and cracking typical of well-developed soil crust communities (St. Clair et al. 1984, Eckert et al. 1986). Patterns of water infiltration and evaporation differ depending on the structure and biological composition of soil crust communities. Some crusts show increased infiltration with decreased evaporation while others demonstrate the opposite effect (Harper and Marble 1988, West 1990). Finally, microbiotic soil crusts have been shown to improve soil fertility. For example, both free-living and lichenized cyanobacteria fix significant amounts of atmospheric nitrogen (West and Skujins 1977, Jeffries et al. 1992). At least some of this nitrogen is ultimately used by the vascular plant community (Harper and Pendleton 1993). Furthermore, microbiotic crusts also contribute organic matter through primary productivity of cyanobacteria and eukaryotic algae. Due to this increased organic matter and reduced erosion of silts and clays, cation exchange capacity is often higher in crusted soils.
Microbiotic crusts in North America are most prevalent and best developed in the semiarid steppe regions of the Great Basin, Colorado Plateau, and Columbia Basin. They also extend into the hotter, more arid deserts in the southwestern portion of the United States and into Mexico. These areas differ from the semiarid regions east of the Rocky Mountains in that they have developed without the pressure of large herds of grazing ungulates (i.e. bison) and extensive wildfires. Antelope, mule deer, and elk grazed the semiarid Intermountain Area before the arrival of European settlers, but these animals did not graze in large herds and occupied crusted areas only during the colder, wetter months of the year when the crusts were less vulnerable. The prehistoric dominance of native bunch grasses, shrubs, and microbiotic soil crusts in the Intermountain West reflects the general lack of intensive grazing pressure in this region (Mack and Thompson 1982).
With the advent of European settlers and the subsequent introduction of large herds of sheep and cattle, vascular plant communities and microbiotic soil crusts have been significantly impacted. Domestic grazing animals typically do serious damage to soil crust communities through trampling, particularly during the drier, warmer periods of the year (Anderson et al. 1982b, Harper and Marble 1988). More recently, destruction of soil crusts by off-road vehicles and backpackers has become an increasing concern in many areas. Wildfires also cause serious damage to soil crust communities by killing most of the lichens, bryophytes, and algae (Johansen et al. 1982, 1984). The frequency and intensity of wildfires in the Intermountain Area has increased significantly due to the pervasive spread of several introduced annuals, for example, cheat grass (Bromus tectorum).
Several factors influencing the development of microbiotic crusts have been studied (Anderson et al. 1982a). Generally, the data show that silty soils with high electrical conductivity are more likely to have well developed soil crust communities. Several reclamation studies have also been undertaken to evaluate recovery patterns of soil crust components following various types of disturbance (Anderson et al. 1982b, Johansen and St. Clair 1986, Johansen et al. 1982, 1984). These studies indicate that recovery is generally slow, but may be greatly enhanced following periods of unusually high precipitation. Algae generally recover first followed by lichens and mosses. Several studies have shown that application of inoculants can accelerate recovery (St. Clair et al. 1986, Belnap 1993, Johansen et al. 1998).
Literature Cited
Anderson, D.C., K.T. Harper and R.C. Holmgren. 1982a. Factors influencing development of cryptogamic soil crusts in Utah deserts. Journal of Range Management 35: 180-185.
Anderson, D.C., K.T. Harper and S.R. Rushforth. 1982b. Recovery of cryptogamic soil crusts from grazing on Utah winter ranges. Journal of Range Management 35: 355-359.
Belnap, J. 1993. Recovery rates of cryptobiotic crusts: inoculant use and assessment methods. Great Basin Naturalist 53: 89-95.
Blackburn, W.H. 1975. Factors influencing infiltration rate and sediment production of semiarid rangelands in Nevada. Water Resources Research 11: 929-937.
Eckert, R.E., Jr., F.F. Peterson, M.S. Meurisse and J.L. Stephens. 1986. Effects of soil-surface morphology on emergence and survival of seedlings in big sagebrush communities. Journal of Range Management 39: 414-420.
Fletcher, J.E. and W.P. Martin. 1948. Some effects of algae and moulds in the rain crust of desert soils. Ecology 29: 95-100.
Harper, K.T. and J.R. Marble. 1988. A role for nonvascular plants in management of arid and semiarid rangelands. Pages 135-169 in P.T. Tueller, ed., Vegetation Science Applications for Rangeland Analysis and Management. Kluwer Academic Publishers, Dordrecht; Boston; London.
Harper, K.T. and R.L. Pendleton. 1993. Cyanobacteria and cyanolichens: can they enhance availability of essential minerals for higher plants? Great Basin Naturalist 53: 59-72.
Jeffries, D.L., J.M. Klopatek, S.O. Link and H. Bolton, Jr. 1992. Acetylene reduction by cryptogamic crusts from a blackbrush community as related to resaturation and dehydration. Soil Biology and Biochemistry 24: 1101-1105.
Johansen, J.R., A. Javakul and S.R. Rushforth. 1982. Effects of burning on the algal communities of a high desert soil near Walsburg, Utah. Journal of Range Management 35: 598-600.
Johansen, J.R., L.L. St. Clair, B.L. Webb and G.T. Nebeker. 1984. Recovery patterns of cryptogamic soil crusts in desert rangelands following fire disturbance. Bryologist 87: 238-243.
Johansen, J.R. and L.L. St. Clair. 1986. Cryptogamic soil crusts: recovery from grazing near Camp Floyd State Park, Utah, USA. Great Basin Naturalist 46: 632-640.
Johansen, J.R., L.L. St. Clair, D. Evans, V. Flechtner, J. Balczon, and B.L. Webb. 1998. Resilience of Microbiotic Species to Military Training Pressures: Natural and Stimulated Recovery Following Disturbance. Technical Report (Contract DACA88-95-C-0015) USACERL. 210 pages.
Kleiner, E.F. and K.T. Harper. 1972. Environment and community organization in grasslands of Canyonlands National Park. Ecology 53: 299-309.
Mack, R.N. and J.N. Thompson. 1982. Evolution in steppe with few large, hooved mammals. American Naturalist 119: 757-773.
MacKenzie, H.J. and H.W. Pearson. 1979. Preliminary studies on the potential use of algae in the stabilization of sand wastes and wind blow situations. British Phycological Journal 14: 126.
St. Clair, L.L., B.L. Webb, J.R. Johansen and G.T. Nebeker. 1984. Cryptogamic soil crusts: enhancement of seedling establishment in disturbed and undisturbed areas. Reclamation and Revegetation Research 3: 129-136.
St. Clair, L.L. and J.R. Johansen. 1993. Introduction to the symposium on soil crust communities. Great Basin Naturalist 53: 1-4.
St. Clair, L.L., J.R. Johansen and B.L. Webb. 1986. Rapid stabilization of fire disturbed sites using a soil crust slurry: inoculation studies. Reclamation and Revegetation Research 4: 261-269.
West, N.E. 1990. Structure and function of microphytic soil crusts in wildland ecosystems of arid to semi-arid regions. Advances in Ecological Research 20: 179-223.
West, N.E., and J. Skujins. 1977. The nitrogen cycle in North American cold-winter semidesert ecosystems. Oecologia Plantarum 12: 45-53.
Additions to the Lichen Flora of San Clemente Island, California
Charis Bratt
Santa Barbara Botanic Garden
1212 Mission Canyon Rd.
Santa Barbara CA 93105
In the Bulletin of the California Lichen Society 3(2), 1996, Bowler, Weber, and Riefner published a checklist of the lichens of San Clemente Island that included 137 entities: 125 species or infraspecific taxa (3 identified approximately) and 12 entities identified only to genus. They also listed 11 taxa mentioned in Hasse (1903) but not encountered by them. Most of the records in the report were based on collections made by Weber and Santesson in 1966, and some were based on collections by Bowler and Riefner.
Two of the records deserve special comment. The Weber/Santesson collection #42907 that is listed as Toninia tristis has been annotated at COLO by Timdal as Toninia ruginosa ssp. pacifica. According to Timdal's monograph of Toninia (Timdal 1991), T. tristis does not occur in California. All of the specimens cited by Bowler, Weber, and Riefner as Opegrapha sp. in COLO (L-42619, L-42623, and L-42847) have been annotated by Egea as Opegrapha brattiae (an unpublished name).
The present report is based on collections made at San Clemente Island in 1996 and 1997. Fifty-seven species are reported for the first time; no attempt has been made to add new records of species previously listed from the island. Many of the collection sites listed are in the southern third of the island which, because of severe restrictions imposed by the U.S. Navy, had not previously been visited. In this area the canyons on the northeast side of the island are very short and steep. On the southwest side of the island the slope is more gradual although the canyon bottoms are narrow and often more difficult to traverse. Most of the collecting was done near the tops of the canyons. Younger and more agile lichenologists will find much area left to explore.
The most exciting discovery is that of Texosporium sancti-jacobi in two different but adjacent locations in Chenetti Canyon, one at 365 m and the other at 260 m. This is the first report of this species from any of the Channel Islands and only the fourth reported location for this lichen in California. It is interesting that in both of these collections, Texosporium was growing on other lichens, albeit different species. This lichen is otherwise found on detritus or on dried rabbit dung.
The following list is by no means complete as there are many more areas to explore and many more specimens to be identified. All specimens representing original records are stored in SBBG.
Locations:
BC: Box Canyon
CB: China Beach
CH: Chukit Canyon
CL: Canchalagua Canyon
CM: Living Compound
CN: Chenetti Canyon
CP: China Point
CV: Cave Canyon
EC: Eagle Canyon
MA: Camera pad "Malo"
MC: Mosquito Canyon
MR: Middle Ranch
PP: Pot Pond on road to the airport
RW: REWS site between Thirst and Chukit Canyons
SP: Spray
ST: Snapper Tower
VO: Vista Oaks
Collectors:
MD: Marla Daily
SJ: Steve Junak
All others are collections of the author.
Acarospora carnegiei Zahlbr.--CL: 9720, 9730.
Aspicilia caesiocinerea (Nyl. ex Malbr.) Arnold--ST: 9616; RW: 9808; CL: 9723.
Aspicilia calcarea (L.) Mudd --CL: 9702.
Aspicilia cinerea (L.) Körber --EC: 10134, 10149.
Buellia retrovertens Tuck. --RW: 9799.
Buellia turgescens Tuck. --EC: 10164.
Caloplaca brattiae W.A. Weber --ST: 9646; BC: 9742; RW: 9800, 9802; CL: 9725.
Caloplaca cerina (Hedwig) Th. Fr. --PP: 9572. On twigs of Lycium.
Caloplaca epithallina Lynge --CN: 10,271. On Endocarpon.
Caloplaca ferruginea (Hudson) Th. Fr.--CH: 9676. On Prunus.
Caloplaca ignea Arup --MC: 10177.
Candelariella coralliza (Nyl.) H. Magn. --MS: 10200, 10204.
Candelariella rosulans (Müll. Arg.) Zahlbr. --EC: 10161.
Candelariella vitellina (Hoffm.) Müll. Arg. --CL: 9718.
Cladonia pyxidata (L.) Hoffm. --EC: 10151.
Dendrographa leucophaea (Tuck.) Darbish. f. minor (Darbish.) Sundin & Tehler (cited by Sundin and Tehler, 1996 as transitional between f. minor and f. leucophaea)
Diploschistes actinostomus (Ach.) Zahlbr. --ST: 9740; CL: 9729, 9722.
Flavopunctelia soredica (Nyl.) Hale --VO: 9927. On Quercus tomentella.
Fuscopannaria leucophaea (Vahl) P.M. Jørg. --CL: 9711; MA: SJ SCl-993C.
Lecania naegelii (Hepp) Diederich & v. d. Boom --VO: 9926. On Quercus tomentella.
Lecanora meridionalis H. Magn. --CH: 9679. On Prunus.
Lichinella nigritella (Lettau) Moreno & Egea --CH: 9870.
Lichinella stipatula Nyl. --MR: 9770; CH: 9864; CV: 9836.
Melanelia fuliginosa (Fr. ex Duby) Essl. --ST: 9602, 9738, 9751.
Niebla dissecta Spjut --RW: 9830.
Niebla laminaria Spjut --CP: 10258, CM 10130
Niebla sorediata Spjut --EP: COLO Ex #185. Distributed as Ramalina homalea. Isotype.
Niebla sorocarpia Spjut --CM 10128.
Niebla testudinaria (Nyl.) Spjut --CP: 10255.
Opegrapha brattiae Egea & Torrente (nom. ined.), fide Egea annotation of specimen --ST: 9642.
Parmotrema stuppeum (Taylor) Hale --MR: 9789, 9780; CL: 9726.
Peltula omphaliza (Nyl.) Wetmore --CV: 9846.
Peltula patellata (Bagl.) Swinscow & Krog --RW: 9811.
Phlyctis argena (Sprengel) Flotow --VO: 9910. On Quercus tomentella.
Physcia tribacia (Ach.) Nyl. --EC: 10157, 10140, 10148; MA: SJ SCl-994 A.
Placidium chilense (Räsänen) Breuss --CH: 9861.
Placidium lacinulatum (Ach.) Breuss --CV: 9850, 9851; CH: 9859, 9860.
Pleopsidium chlorophanum (Wahlenb.) Zopf --CB: 10249.
Protoparmelia badia (Hoffm.) Hafellner --ST: 9637.
Psora pacifica Timdal --CV: 9856 A.
Psora tuckermanii R. Anderson ex Timdal --CH: 9858; RW: 9817.
Sclerophyton californicum (Tuck.) Hasse --MR: 9793.
Syzygospora physciacearum Diederich --MA: SJ SCl-994 B. On Physcia tribacia on rock face. Note: this is a lichenicolous heterobasidiomycete.
Teloschistes exilis (Michx.) Vainio --ST: 9622; On rock. VO: 9913. On Quercus tomentella.
Tephromela nashii Kalb --PP: 9588.
Texosporium sancti-jacobi (Tuck.) Nádv. --CN: 10312. On a white crustose lichen. CN: 10276; on Peltula patellata. First published record from the California Channel Islands.
Thelomma santessonii Tibell --CL: 9704; RW: 9806; SP: 9660; PP: 9569; MA: 10222; EC: 10155; ST: 9636; BC: 9601.
Usnea esperantiana Clerc --FS: MD 22. On Quercus tomentella.
Vermilacinia acicularis Spjut --EC: 10147, 10132, 10139; RW: 9829; BC: 9606.
Vermilacinia cerebra Spjut --CM: 10129; CP: 10257.
Vermilacinia nylanderi Spjut --MC: 10186
Vermilacinia pumila Spjut --SP: 9657, 9649; ST: 9599; CL: 9696.
Xanthoparmelia coloradoënsis (Gyelnik) Hale --CV: 9837, 9838; CL: 9706, 9693, 9695.
Xanthoparmelia conspersa (Ehrh. ex Ach.) Hale --EC: 10159.
Xanthoparmelia cumberlandia (Gyelnik) Hale --CL: 9699, 9710.
Xanthoparmelia plittii (Gyelnik) Hale --EC: 10153; MA: 10224.
Xanthoria candelaria (L.) Th. Fr. --CH: 9669; RW: 9801; VO: 9908; MR: 9764.
Acknowledgements
I am most thankful to Steve Junak, Santa Barbara Botanic Garden, for taking me to San Clemente Island as his assistant and to Jennifer Stone, U.S. Navy, Southwest Division, who arranged for transportation to the island and housing while we were there. The Navy personnel on the island and the staff at the Environmental Station were most cordial and helpful. I am grateful to Drs. Bruce Ryan, Teuvo Ahti, John Sheard, Jose Maria Egea, Clifford Wetmore, and Othmar Breuss for many of the identifications.
Literature Cited
Bowler, P.A., W.A. Weber and R.E. Riefner, Jr. 1996. A checklist of the lichens of San Clemente Island, California. Bulletin of the California Lichen Society 3(2):1-8.
Hasse, H.E. 1903. The lichen-flora of San Clemente Island. Bulletin of the Southern California Academy of Sciences 2: 54-55.
Sundin, R. and A. Tehler. 1996. The genus Dendrographa (Roccellaceae). The Bryologist 99: 19-31.
Timdal, E. 1991. A monograph of the genus Toninia. Opera Botanica 110: 1-137.
Questions and Answers
Janet Doell
1200 Brickyard Way #302
Point Richmond, CA 94801
When lecturing to the general public about lichens, I field certain questions which are of common interest to those attending. I have chosen three of these for a trial "question and answer" column in the CALS Bulletin with the expectation that some readers may have wondered about these subjects. The column is meant to serve people who are new to lichens and do not have easy access to lichen literature, but also people with more experience who may themselves be asked questions like these. I will be happy to address questions asked by readers.
1. Question
What is the phylogeny of lichens? Where do they fit into the scheme of phylum, order, etc.
Answer
The whole subject of lichen phylogeny is under scrutiny at the moment, but here is some information on the subject. You can see right from the start that what we are talking about here is the fungal partner only, not the algal component of the lichen. Lichens don't really fit anywhere perfectly because they are not an "organism" in the usual sense; rather their component bionts fit separately into classification schemes.
Ainsworth & Bisby's Dictionary of the Fungi, eighth edition (Hawksworth et al., 1995, pp. 169-171, 227-228) includes a discussion about the complexity of "fungi", but it segregates as the Kingdom Fungi four phyla of what have been traditionally regarded as fungi. Two of these phyla are Ascomycota and Basidiomycota, which include the lichen fungi. Like green algae, fungi are eukaryotic, the cell nucleus being enclosed in a membrane; by contrast, the nuclear material in cyanobacteria ("blue-green algae") is not surrounded by a membrane (prokaryotic).
Fungi have no photosynthetic pigments. They absorb nutrients from dead or living organisms; as decomposers of the biosphere, they break down organic products, restoring carbon, nitrogen and other components to soil and air.
Most lichen fungi are ascomycetes (spores borne inside asci); few are basidiomycetes (spores borne externally on basidia). Much more information is needed in order to clarify phylogenetic relationships, and some categories used elsewhere have been discarded in Ainsworth & Bisby's Dictionary of the Fungi of 1995, which includes 46 orders and 264 families, only a few of which include lichens.
Included within the orders (names ending in -ales, such as Lecanorales) are families (names ending in -aceae; thought to be interrelated). The families include what are construed as related genera, and within each genus is included an assemblage of species presumably more closely related to one another than they are to species in another genus. According to Hale (1974, p. 156) "species are recognized by more or less arbitrary combinations of vegetative and ascocarp characters". If we add to this chemical characteristics, DNA and other current microbiological information, the species concept becomes very complex. You will find a current discussion on this subject, with more detailed information in Tehler (1996).
2. Question
What are some ascomycetes that are not lichenized?
Answer
Ascomycetes are a large, widespread group of fungi, but most non-lichenized species are small and inconspicuous. Brewer's yeast, for example, is an ascomycete, as are many plant pathogens, both in the temperate zones and in the tropics. Penicillium is also an ascomycete. A few fleshy, mushroom-like fungi are ascomycetes, but gilled mushrooms belong to the basidiomycetes, in which the spores are borne externally.
3. Question
Corals have a way of controlling the population size of their symbiotic algal partners. Do lichens have a similar mechanism?
Answer
A study of Flavoparmelia caperata was cited by Ahmadjian (1993) as exhibiting some kind of control mechanism; he stated (Ahmadjian, 1993, p. 88) that "The photobiont population appeared to be under some type of limiting control, beyond what might be expected from environmental constraints. Perhaps the mycobiont produces inhibitors in older regions of a thallus that affect photobiont division." He added a quotation from Honegger's paper on pattern formation in the algal layer of lichens with stratified thalli (Honegger, 1987, Bibliotheca Lichenologica 25: 59-71), to the effect that Honegger felt that "the photobiont cells of non-growing areas are slowed down in their metabolic activity by a probably, mycobiont-derived inhibitory complex, the molecular basis of which is not yet understood."
Literature Cited
Ahmadjian, V. 1993. The Lichen Symbiosis. John Wiley & Sons, Inc. New York, i-xv, 250 p.
Hale, M. 1974. The Biology of Lichens. Second Edition. American Elsevier Publishing Co., Inc. New York, 181 pp.
Hawksworth, D.L., P.M. Kirk, B.C. Sutton and D.N. Pegler 1995. Ainsworth & Bisby's Dictionary of the Fungi. Eighth Edition. CAB International, Wallingford.
Tehler, A. 1996. Systematics, phylogeny and classification. Chapter 12. Pp 217-239, in Lichen Biology, ed. T.H. Nash, III. Cambridge University Press, Cambridge.
Preliminary List of Rare California Lichens
David L. Magney
P.O. Box 1346,
Ojai, CA 93024-1346
Very little work has been done to date to identify which lichens occurring in California are rare. A few lichens have been placed on the Preliminary Global Red List of Lichens by the International Committee for the Conservation of Lichens (ICCL), set up by the International Association of Lichenologists (IAL), which is a specialist group connected to IUCN (World Conservation Union). That list is available at the Swedish Threatened Species Unit web-site
(http://www/dha.slu.se/guest/global.htm),
which lists only extinct and critically endangered species. The Global Red List only includes three taxa from California. Clearly, many more than three California lichen taxa are rare.
The California Native Plant Society (CNPS) has spent a great deal of time and put a lot of thought into developing its Inventory of Rare and Endangered Vascular Plants of California, now in its 5th edition. The CNPS list has become an extremely important tool for conserving the California flora, and is the standard which is used by state, federal, and local agencies, and others, when evaluating land management options, such as under the environmental evaluation process of the California Environmental Quality Act (CEQA). However, no such parallel list occurs for the lichens of California, some of which are as rare and endangered as any vascular plant or wildlife species. Unfortunately, the only practical way in which plants are considered under the CEQA process is if a list is published by a respectable group or scientist(s), such as CNPS or CALS.
Therefore, I think there is a great need to start the process of developing a list of rare and endangered lichens for California. I suggest that CALS work with CNPS to gather the necessary data to support our ranking of the California plants that we, the professional, academic, and amateur lichenologists, are most familiar with--this unique group called lichens.
For the past few years I have been pushing consultants and lead agencies in California to evaluate the lichen flora of a project site equally to that of the vascular plant flora for any project evaluated under CEQA. After all, lichens should have the same standing as vascular plants and animals. Recently, good efforts have been made with this for environmental impact reports (EIRs) in Ventura, Santa Barbara, Sonoma, and San Luis Obispo counties, so a precedent has already been set. In fact, a recent (October 1998) court ruling by the Ventura County Superior Court found a project EIR insufficient, according to CEQA Guidelines. The court found that the consultants and lead agency failed to conduct a survey of the lichen flora of the project site, or evaluate project-related impacts on lichens. The court then ordered that such surveys and analysis must be conducted to satisfy CEQA requirements.
I recommend that we apply the same rules to California lichens for determining rarity as those used by CNPS and the IUCN. Both ranking systems can be used to properly place our California rare lichens on such a list based on: number of populations or individuals; level of threat to a taxon's existence; and distribution. As Violeta Atienza of the University of Valencia, Spain pointed out (International Lichenological Newsletter 31(2): 57. 1998), determining lichen population size and viability is challenging at best, when trying to apply methods used for vascular plants.
With these ideas in mind, I have developed a preliminary list of lichens that may be considered endangered, rare, or at least uncommon, in California. Part of the problem we face in developing such a list is that many lichenologists think that we don't know enough about the California lichen flora to be able to accurately assign rarity levels to lichen taxa. It is true that many areas of California have been poorly, or not even, collected, or documented. This is true also for vascular plants, but less so. However, we need to start somewhere and now is as good a time as any. Some taxonomic groups of lichens are poorly understood, with many species often being misidentified. Some lichen species I have considered here are believed to be too common to include, but few records are known for them in California.
This list will be revised on a periodic basis. It is often the case that, when someone publishes a flora, numerous comments are submitted to the author(s) about all those omitted taxa. While floristic and taxonomic work is never done, identifying a task and challenging others to work on it can raise considerable interest. With that in mind, I present here a preliminary list of rare California lichens.
I have suggested rarity status categories following both the CNPS and IUCN designations. Both ranking/listing criteria are provided below in Table 1 - Natural Diversity Data Base Element Ranking System.
The CNPS R-E-D Code is a numerical ranking for each of the three categories (rarity, endangerment, and distribution) that more accurately describes each plant's population levels and is specific for each of the three categories, as described in Table 2 - CNPS R-E-D Code.
CNPS categorizes each rare plant into five lists (1A, 1B, 2, 3, and 4), as described below in Table 3 - CNPS Rare Plant Lists. I find these lists generally good; however, the CNPS categories leave a large gap between List 1B and List 4, making it difficult to properly place some taxa that are clearly rare but not endangered.
I would also suggest that in future lists we consider rarity on a more local level, such as coastal versus inland, northern California versus southern California, and by county, biogeographic, or floristic region.
Each taxon has been assigned to a global and state ranking and to one of the CNPS lists; however, rare lichens will not be published in the CNPS Inventory, at least not in the sixth edition.
This preliminary list identifies 38 lichen taxa that, based on available information at the time of this publication, appear to be rare in California. Some of these may be more common while others not listed here are indeed rare and should be added to this list.
Charis Bratt reviewed this and earlier drafts of this list and together we eliminated many species that we agreed were too common to be included. Darrell Wright also reviewed the manuscript. We applied fairly high requirements for inclusion on this list, deciding not to include lichen taxa that, based on our experience and intuition, we think are seriously undercollected or documented. Unverified records of lichens in California were discounted and not included in this list. Please review this list and provide as much data and criticism as you can to me at:
FAX 805/646-6975 or
email to dmagney@aol.com.
I look forward to your assistance.
Preliminary List of Rare California Lichens
Bacidina californica S. Ekman / Lecideaceae G2/S2.1; CNPS 4 - 3-1-3. Marin Co.; Monterey Co., Big Sur (type locality); Santa Barbara Co., Santa Barbara foothills at SR154 & SR192 Bratt & Magney 10,319 (Bratt herbarium); San Mateo Co.; Channel Islands. On bark of Aesculus californica, Cupressus, and Umbellularia californica. Possibly under-collected.
Bryoria pseudocapillaris Brodo & D. Hawksw. / Parmeliaceae G1/S1.1; CNPS 2 - 2-3-2; Critically Endangered (B1), on Global Red List (Thor 1996). San Luis Obispo Co., Baywood Park; Humboldt Co.; Oregon, Florence.
Bryoria spiralifera Brodo & D. Hawksw. / Parmeliaceae G2/S2.2; CNPS 1B - 2-3-3. Humboldt Co., Samoa Peninsula; Monterey Co., near Pt. Lobos; Santa Clara Co., Happy Valley J.F. McBride s.n. Apr 1946 on Usnea collection (D. Wright unpubl.; F); San Luis Obispo Co. (Baywood Park, Montaña de Oro SP; Los Osos Oaks Reserve); Sonoma Co.
Caloplaca subpyraceella (Nyl. in Hasse) Zahlbr. / Teloschistaceae G3/S3.3; CNPS 4 - 2-2-3. Butte Co.; Marin Co.; Santa Barbara Co., Santa Barbara foothills at SR154 and SR192 Bratt & Magney 10,321 (Bratt herbarium); San Diego Co., Pt. Loma; San Luis Obispo Co.; Ventura Co., Conejo Mtn. Region, Pt. Mugu, north edge of Toland Road Landfill Bratt & Magney (Bratt herbarium). Possibly under-reported.
Cladonia firma (Nyl.) Nyl. / Cladoniaceae G3/S1.1; CNPS 2 - 3-3-2. San Diego Co., Torrey Pines Reserve?; San Luis Obispo Co. ( Los Osos; Baywood Park Magney [Bratt herbarium]); Mediterranean Region.
Cladonia thiersii S. Hammer / Cladoniaceae G2/S2.3; CNPS 4 - 2-2-3. Marin Co., Pt. Reyes Peninsula; Sonoma Co.; Mendocino Co.; San Diego Co., Torrey Pines Reserve?.
Coccotrema pocillarium (Cummings) Brodo [Ochrolechia pacifica H. Magn., Perforaria minuta Degel.] / Pertusariaceae? G1/S1.2; CNPS 1B - 3-2-2. Sonoma Co., SR1 near Fort Ross. Probably under-collected and under-reported.
Cyphelium brunneum W.A. Weber / Coniocybaceae G3/S3.2; CNPS 1B - 2-2-3. Orange Co., Niguel Hill; Channel Islands.
Gyalecta herrei Vzda / Gyalectaceae G3/S3.2; CNPS 4 - 2-2-3. Humboldt Co.; Orange Co., Aliso Pk.; Marin Co., three localities (Wright unpubl.); San Mateo Co.
Lecania cyathiformis Szatala / Lecanoraceae G1.S1.1; CNPS 1B - 3-3-3. San Diego Co., La Jolla Parks 3371 (UC) (Tavares 1997); San Luis Obispo Co. On Rhus.
Lecanora phryganitis Tuck. / Lecanoraceae G3/S3.3; CNPS 4 - 1-1-3. Marin Co., "South Franklin Rocks"/Tomales Bay (West and Doell 1995); Mendocino Co.; Monterey Co.; San Luis Obispo Co., Morro Rock Reserve; Santa Cruz Co.; San Francisco Co.; San Mateo Co.; Sonoma Co.
Phaeophyscia decolor (Kashiw.) Essl. / Physciaceae G2/S2.3; CNPS 4 - 2-2-2. Alpine Co., Clark Fk. Rd., Carson-Iceberg Wild.; Inyo Co.; Los Angeles Co., Eagles Roost Sand Shed-SR2; Madera Co., Rock Cr. Camp; San Diego Co., Palomar Divide Truck Trail; Shasta Co., Manzanita Cr.-SR44; Siskiyou Co., Shackleford Cr.; Tulare Co., Mt. Whitney; Tuolumne Co., Herring Cr. Rd. May be under-collected and -reported.
Phaeophyscia kairamoi (Vainio) Moberg [Physcia kairamoi Vainio]/ Physciaceae G2/S2.3; CNPS 4 - 1-1-2. Inyo Co., Death Valley; Los Angeles Co., Santa Catalina I. (Los Angeles Co.); Monterey Co., Arroyo Seco Cyn. Camp; Santa Barbara Co., Black Mtn., Santa Rosa I. (Santa Barbara Co.); San Luis Obispo Co., Coon Cr., Montaña de Oro; Ventura Co., Pt. Mugu, Conejo Mtn. (Riefner 1992). May be under-collected and -reported.
Phaeophyscia sciastra (Ach.) Moberg [Physcia sciastra (Ach.) Du Rietz] / Physciaceae G2/S2.3; CNPS 4 - 1-1-2. Fresno Co.: Marin Co.; Riverside Co., Pinyon Flats at SR74; Shasta Co., Manzanita Cr. at SR44; Ventura Co., Sespe Cr. at SR33. May be under-collected and -reported.
Physcia halei J.W. Thomson / Physciaceae G1/S1.2; CNPS 1B - 3-2-2. Tulare Co., Jack Flat Camp at SR137. May be under-collected and -reported.
Physcia magnussonii Frey / Physciaceae G1/S1.3; CNPS 1B - 2-1-2. Lake Co., Lake Pillsbury. Identification needs to be checked.
Protoparmelia badia (Hoffm.) Hafellner [Lecanora badia (Hoffm.) Ach.] / Lecanoraceae G4/S2.2; CNPS 2 - 2-2-1. Calaveras Co.; El Dorado Co., Lost Lake/Sugar Pine SP; Inyo Co.; Tulare Co., Kaweah R. at SR198 at 300m; Ventura Co., Pt. Mugu; Colorado and eastward where it is common. California records need to be verified.
Punctelia punctilla (Hale) Krog / Parmeliaceae G1/S1.1; CNPS 1B - 3-3-2. Ventura Co., Conejo Mtn. (Riefner 1992), Pt. Mugu.
Pyrrhospora russula (Ach.) Hafellner [Lecidea russula Ach.] / Lecideaceae G2/S2.2; CNPS 4 - 2-2-2. Lake Co., Lake Pillsbury Tucker 19 APR 1997 (Wright 1997); Santa Barbara Co.?; Trinity Co. Probably under-collected and -reported.
Ramalina thrausta (Ach.) Nyl. / Ramalinaceae G3/S1.2; CNPS 2 - 2-2-2. Sonoma Co., between Cazadero and Fort Ross (Sanders 1997); Oregon; North America.
Rhizocarpon concentricum (Davies) Beltr. / Lecideaceae G1/S1.2; CNPS 1B - 3-2-2. Mendocino Co.; San Bernardino Co., Granite Mountains (Bratt pers. comm.); San Luis Obispo Co., near Montaña de Oro (Riefner); Arizona. California collections need to be verified.
Roccella babingtonii Mont. / Roccellaceae G1/S1.1; CNPS 1B - 3-3-2. San Clemente I. (Los Angeles Co.), Eel Pt. Bowler 83-102 (IRVC; Bowler et al., 1996); San Diego Co.
Roccella fimbriata Darbish. / Roccellaceae G1/S1.1; CNPS 1B - 3-3-2. San Clemente I. (Los Angeles Co.), Eel Pt. Weber & Santesson L-42600 (COLO), Riefner & Bowler 89-109 (IRVC; Bowler et al., 1996); San Diego Co., Pt. Loma (Bratt 1997); San Diego Co.; San Luis Obispo Co., White Pt./Morro Bay SP.
Roccellina conformis Tehler / Roccellaceae G1/S1.1; CNPS 1B - 3-3-2. Santa Catalina I. (Los Angeles Co.), near Avalon (Tehler 1402a S - Type; additional collections from Santa Catalina). On bark.
Roccellina franciscana (Zahlbr. ex Herre) Follman [Dirina franciscana Zahlbr. ex Herre, Schismatomma cupressum Herre] / Roccellaceae G2/S2.2; CNPS 1B - 1-2-3. Monterey Co.; Santa Cruz Co.; San Mateo Co.; San Francisco Co.; San Luis Obispo Co., Morro Rock Reserve, White Pt./Morro Bay SP.
Sulcaria badia Brodo & Hawksw. / Alectoriaceae G1/S1.1; CNPS 2 - 3-3-2; Critically Endangered on Global Red List (Thor 1996). Mendocino Co., on apple and oak in Quercus garryana woodland; Oregon, Benton County; Washington, Clallam County (Olympic Peninsula) (may be extirpated). Mendocino Co. occurrences may be only viable populations. (Peterson et al. 1998.)
Sulcaria isidiifera Brodo / Alectoriaceae G1/S1.1; CNPS 1B - 3-3-3. San Luis Obispo Co., Montaña de Oro SP, Field Ranch, Morro Bay SP, Rancho Cañada de Los Osos, Baywood Park.
Teloschistes californicus Sipman / Teloschistaceae G2/S2.2; CNPS 1B - 2-2-2. Marin Co.; San Mateo Co.; Santa Cruz Co.; Channel Islands except Santa Cruz I. (Santa Barbara Co.).
Teloschistes exilis (Michaux) Vainio / Teloschistaceae G3/S3.3; CNPS 4 - 1-2-1. Los Angeles Co., Pasadena, Claremont; Marin Co., Copper Mine Gulch, McCurdy Trail at Bolinas Ridge, Mt. Tamalpais/Rock Springs (West and Doell 1995); Mendocino Co.; Riverside Co., Santa Rosa Plateau; Santa Barbara Co., Santa Barbara foothills at SR154 and SR192 (Bratt & Magney 10,367 [Bratt herbarium]); San Luis Obispo Co., Cerro Alto SP, Montaña de Oro SP; San Mateo Co.; Sonoma Co.
Teloschistes flavicans (Sw.) Norman / Teloschistaceae G3/S3.3; CNPS 4 - 1-2-1. Los Angeles Co., Santa Monica Mtns.; Marin Co., Mt. Tamalpais, Mill Valley, "South Franklin Rocks"/Tomales Bay (West and Doell 1995); Monterey Co.; Santa Barbara Co., Pt. Sal; San Clemente I. (Los Angeles Co.), Lost Point Canyon Weber & Santesson L-42890 (COLO); San Luis Obispo Co., Montaña de Oro SP, SR46 at Cypress Mtn.; San Mateo Co., Pilarcitos Cr. Cyn.; Channel Islands. On shrubs and branchlets.
Texosporium sancti-jacobi (Tuck.) Nádv. [Cyphelium sancti-jacobi (Tuck.) Zahlbr.] / Caliciaceae G2/S1.1; CNPS 2 - 3-3-2; Critically Endangered, on Global Red List (Thor 1996). San Benito Co., Pinnacles NM; San Diego Co. (likely extirpated); Santa Barbara Co., Aliso Cyn./Cuyama Valley on detritus; San Clemente I. (Los Angeles Co.) parasitic on lichens; Oregon (2 sites 6 km apart) at base of bunchgrasses; Idaho (13 sites within a 30 km radius). California populations are often associated with Artemisia in undisturbed Coastal Sage Scrub-Grassland habitats, on animal (rabbit) dung, and detritus.
Toninia submexicana de Lesdain / Lecideaceae G4/S1.2; CNPS 2 - 2-2-2. Los Angeles Co.; Santa Barbara Co., De la Guerra Springs; San Luis Obispo Co.; Ventura Co.; Baja California. Mostly saxicolous on basalt and serpentine, six localities on the south coast and four in Baja California (Bratt and Wright 1995).
Toninia verrucarioides (Nyl.) Timdal / Lecideaceae G5/S1.2; CNPS 2 - 2-2-1. Santa Clara Co.; Rockies; w. Europe. Apparently always on cyanophilic lichens, especially Placynthium, on basic rock or soil on rock; one California locality on central coast, temperate and boreal in the Rockies and western Europe (Bratt and Wright 1995).
Verrucaria mucosa Wahlenb. / Verrucariaceae G1/S1.1; CNPS 1B - 1-2-1. San Luis Obispo Co., Morro Rock Reserve.
Verrucaria tavaresiae R. Moe / Verrucariaceae G1/S1.1; CNPS 1B - 3-3-3. San Francisco Co., Fort Mason Moe & Silva UC1512286 (Holotype); San Mateo Co.; Marin Co. (Moe 1997).
Xanthoparmelia angustiphylla (Gyelnik) Hale / Parmeliaceae G1/S1.2; CNPS 1B - 3-2-2. San Diego Co., Torrey Pines Reserve; San Luis Obispo Co., Coon Cr., Field Ranch; Ventura Co., Conejo Mtn. (Riefner 1992).
Xanthoparmelia californica Hale / Parmeliaceae G2/S2.2; CNPS 1B - 2-2-2. Sacramento Co.; Santa Barbara Co.; San Luis Obispo Co., Camarillo Peaks, Morro Bay SP.
Xanthoparmelia mougeotii (Schaerer) Hale [Parmelia mougeotii Schaerer] / Parmeliaceae G1/S1.2; CNPS 1B - 3-2-2. Humboldt Co.; Plumas Co.; San Luis Obispo Co., Coon Cr. at Montaña de Oro SP; Siskiyou Co.
____________________________
Notes:
Nomenclature follows Esslinger, T.L. and R.S. Egan (1995) and original rarity determinations follow Hale and Cole (1988), Riefner et al. (1995), and Thor (1996). Many southern California localities are from Hasse (1913).
Abbreviations:
Cr. = Creek, Cyn. = Canyon, Fk. = Fork, km = kilometers, SB = State Beach Mt. = Mount, Mtn(s). = Mountain(s), P = Park, Pks. = Peaks, Pt. = Point, R = River, RA = Recreation Area, Res. = Reserve, SF = State Forest, SP = State Park, SR = State Route,Tr. = Trail. Abbreviations of herbaria follow Holmgren et al. (1990).
Table 1. Natural Diversity Data Base Element Ranking System
|
Global Ranking (G) |
|
G1 |
Less than 6 viable element occurrences (populations for species) OR less than 1,000 individuals OR less than 809.4 hectares (ha) (2,000 acres [ac]). |
| G2 | 6 to 20 element occurrences OR 809.4 to 4,047 ha (2,000 to 10,000 ac). |
|
G3 |
21 to 100 element occurrences OR 3,000 to 10,000 individuals OR 4,047 to 20,235 ha (10,000 to 50,000 ac). |
|
G4 |
Apparently secure; this rank is clearly lower than G3 but factors exist to cause some concern (i.e., there is some threat, or somewhat narrow habitat). |
|
G5 |
Population or stand demonstrably secure to ineradicable due to being commonly found in the world. |
|
GH |
All sites are historic; the element has not been seen for at least 20 years, but suitable habitat still exists. |
| GX | All sites are extirpated; this element is extinct in the wild. |
| GXC | Extinct in the wild; exists in cultivation. |
| G1Q | The element is very rare, but there is a taxonomic question associated with it. |