MARCO CONCEPTUAL

Future studies are required in New Zealand to examine epiphytic community patterns on other species of Nothofagus to determine whether the high diversity and patterns of non-vascular epiphytic species is consistent across beech trees. Conditions in Craigieburn of high winds and frequent wetting and drying in combination with thallus morphology may be advantageous for species dispersal. Further studies are required to examine the effective dispersal ability according to size and shape of propagules and how their dimensions may affect dispersal distance and what forces (e.g. electrostatic) are required for propagules to adhere to substrates. Also, as wind is one of the primary forms of disturbance, a better understanding of what effect it has on the dispersal of particular lichen growth forms is necessary. Further investigation is needed to test the

hypotheses that fruticose lichens are better disperses than crustose and foliose lichens. Because the more easily dispersed species are those with higher abundance in primary succession such as in younger stands and in younger regions of the tree, whereas foliose lichens are less easily dispersed and take longer periods of time to establish.

Research in ecology is focused on trying to explain why species are common or rare. In relation to lichens one could ask whether the morphological structure such as numerous perforations and a hollow inner cavity assists with fragmentation and mobility of propagules [e.g. Menegazzia

pertransita] and determines its high abundance. Such fragments could easily be propelled

considerable distances during wind events or dispersed across the bark after wetting, allowing the species to establish and occupy other suitable habitats. To test these hypotheses one possibility involves conducting wind tunnel experiments that expose thalli to environmental conditions and wind events that would simulate conditions in Craigieburn. How different growth forms or morphological structures affect propagule fragments and dispersal distances could then be assessed.

Further studies are required to understand interactions among species in the community such as parasitism, amensalism, mutualism or commensalism (Armstrong 1982). Understanding these interactions can provide knowledge about additional processes involved in structuring epiphytic lichen communities. One such study could examine lichen interaction with other components of the ecosystem such as invertebrates. Various invertebrate species are a part of the beech forest ecosystem and were observed interacting with lichens. Further investigation of how invertebrates such as scale insects influence lichen dispersal and distribution patterns may also explain why M.

molds and sooty molds are clearly an important component of the epiphyte community in

Craigieburn, present in over 90% of the samples in 140yr and 265yr old beech trees. Sooty molds were also able to coexist with M. pertransita, as both species were able to overgrow each other. Further research is required to investigate the outcome of overgrowth between these species as well as other epiphytes in the community to determine if there are any risks of sooty molds invading, outcompeting or inhibiting growth of particular lichens species.

Our knowledge of the dynamic function of secondary metabolites of lichens is limited. For example, does secondary metabolite production provide lichens with a competitive advantage? Atranorin is known to reduce germination of spores of other lichen species (e.g. Caloplaca

citrina) (Huneck 1999) thus it may contribute to M. pertransita’s success by prohibiting growth

of competitive species. An interesting study would be to extract atranorin from M. pertransita thalli and apply to surfaces of bare bark and see if colonization of propagules is slower or faster than an untreated area of similar size.

High abundance and biomass of cyanolichens in older forest suggests that they could be an important contribution to nitrogen budgets and store high concentrations of CO2. Measuring

concentrations and input of nitrogen through active N-fixation could be compared with nitrogen contributed by decomposition of organic matter. Also how much CO2 is being stored by

cyanolichen species particularly in older aged stands. Craigieburn’s environmental conditions showed frequent periods of wet and dry. Understanding the humidity patterns and how they affect cyanolichens P. colensoi and P. faveolata physiologically (e.g. thallus water content and CO2

exchange) to better predict distribution patterns. Additional manipulative experiments are also required to test how important these species are on the cycling of water and other nutrients. What effects will the loss of lichens have on these forests? One possible experiment could include measurements of through flow and nutrient concentrations from lichens and bark over a period of time.

Patterns by lichen according to secondary compounds suggest their potential importance in acclimating to current environmental conditions. The role of UV radiation levels on the

production of usnic acid and whether levels of usnic acid production would be the same. The risk of losing lichens in response to increased atmospheric pollution could also be studied using growth chambers where the growth and thallus health responses to different pollution levels of known concentrations are assessed.

References

Alpin, P.S. and D.J. Hill. 1979. Growth Analysis of circular lichen thalli. Journal of Theoretical Biology 78: 347-363.

Affeld, K., J. Sullivan, S. P. Worner, R. K. Didham and David Bowman. 2008. Can spatial variation in epiphyte diversity and community structure be predicted from sampling vascular epiphytes alone? Journal of Biogeography 35: 2274-2288

Agrawal, A.A., D.D. Ackerly, F. Adler, A. E. Arnold, C. Caceres, D.F. Doak, E. Post, P.J. Hudson, J. Maron, K.A. Mooney, M Power, D. Schemske, J. Stachowicz, S. Strauss, M.G Turner and E. Werner. 2007. Filling the key gaps in population and community ecology. Frontiers in Ecology Environment 5: 145-152.

Antoine, M.E. and B. McCune. 2004. Contrasting fundamental and realized ecological niches with epiphytic lichen transplants in an old-growth Pseudotsuga forest. The Bryologist 107: 163-173.

Archer, A.W. 1991. New species and new reports of Pertusaria (lichenised Ascomycotina) from Australia and New Zealand with a key to the species in Australia. Mycotaxon 41: 223- 269.

Armstrong, R.A. 1975. The influence of aspect on the pattern of seasonal growth in the lichen

Parmelia glabratula ssp. fuliginosa (Fr. ex Duby) Laund. New Phytologist 75: 245-251.

Armstrong, R.A. 1976. The influence of the frequency of wetting and drying on the radial growth of three saxicolous lichens in the field. New Phytologist 77: 719-724. Armstrong, R.A. 1979. Growth and regeneration of lichen thalli with the central portions

artificially removed. Environmental and Experimental Botany 19: 175-178.

Armstrong, R.A. 1981. Competition between three saxicolous species of Parmelia (lichens). New Phytologist. 90: 67-72.

Armstrong, R.A. 1982. Competition between three saxicolous species of Parmelia (lichens). New Phytologist. 90: 67-72.

Armstrong, R.A. 1985. Field experiments on competition between three foliose lichens using the de Wit experimental design. Environmental and Experimental Botany 25: 369-374. Armstrong, R.A. 1990. Dispersal, establishment and survival of the soredia and fragments of

Armstrong, R.A. 1992. A comparison of the growth curves of the foliose lichen Parmelia

conspersa determined by a cross-sectional study and by direct measurement.

Environmental and Experimental Botany 32: 221-227.

Armstrong, R.A. 1993. Factors determining lobe growth in foliose lichen thalli. New Phytologist. 124: 675-679.

Asahina, Y. and S. Shibata. 1954. Chemistry of lichen substances. Tokyo: Japan Society for the Promotion of Science.

Badger, M.R., H. Pfanz, B. Büdel, U. Heber and O.L. Lange. 1993. Evidence for the functioning of photosynthetic CO2- concentrating mechanism in lichens containing green algal and

cyanobacterial photobionts. Planta 191: 57-70.

Bayfield, N.G., U.H. Urquhart, and S.M. Cooper. 1981. Susceptibility of four species of

Cladonia to disturbance by trampling in the Cairngorm Mountains, Scotland. Journal of

Applied Ecology 18: 303-310.

Becker, V.E. 1980. Nitrogen fixing lichens in forests of southern Appalachian Mountains of North Carolina. The Bryologist 83: 29-39.

Bell, A.D. 1991. Plan form: an illustrated guide to flowering plant morphology. Oxford University Press, Oxford.

Benson, S. and D.S. Coxson. 2002. Lichen colonization and gap structure in wet-temperate rainforests of northern interior British Columbia. The Bryologist 105: 673-692. Benzing, D.H. 1990. Vascular epiphytes. Cambridge University Press, Cambridge. Bertness, M.D., and S.W. Shumway. 1993. Competition and facilitation in marsh plants.

American Naturalist 142: 718-724.

Binkley, D. 1986. Forest nutrition management. Wiley, New York.

Bjorn, L.O. 2007. Stratospheric ozone, ultraviolet radiation, and cryptogams. Ecological Conservation 135: 326-333.

Bliss, L.C. and E.B. Hadley. 1964. Photosynthesis and respiration of alpine lichens. American Journal of Botany 51: 870-874.

Boudreault, C., Y. Bergeron, S. Gauthier, and P. Drapeau. 2002. Bryophyte and lichen communities in mature to old-growth stands in eastern boreal forests of Canada. Canadian Journal of Forest Research 32: 1080-1093

Boehm, F. K Clarke, R. Edge, E. Fernandez, S. Navaratnam, W. Quilhot, F. Rancan and T.G. Truscott. 2009. Lichens-Photophysical studies of potential new sunscreens. Journal of Photochemistry and Photobiology B: Biology 95:40-45.

Breslow, N.E. and Clayton, D.G. 1993. Approximate inference in generalized linear mixed models. Journal of the American Statistical Association 88: 9-25.

Brodo, I.M., S.D. Sharnoff and S. Sharnoff. 2001. Lichens of North America. Yale University Press, New Haven USA.

Brodo, I. M. 1961. Transplant experiments with corticolous lichens using a new technique. Ecology 42: 838-841.

Brown, J.H. 1995. Macroecology. The University of Chicago Press. Chicago, USA. Büdel, B. and C. Scheidegger. 2008. Thallus morphology and anatomy. Pp. 40-68

in T. H. Nash III ed., Lichen Biology 2nd edition. Cambridge University Press, Cambridge.

Burns, K.C. and J. Dawson. 2005. Patterns in the diversity and distribution of epiphytes and vines in a New Zealand forest. Austral Ecology 30: 883-891

Cáceres, M.E.S., R. Lücking and G. Rambold. 2008. Efficiency of sampling methods for accurate estimation of species richness of corticolous microlichens in the Atlantic rainforest of northeastern Brazil. Biodiversity Conservation 17: 1285-1301.

Caldiz, M. 2005. Diversity and growth of epiphytic macrolichens in northwestern Patagonian

Nothofagus forests. [dissertation]. Alnarp, Sweden. Swedish University of Agricultural

Sciences.

Caldiz, M. and J. Brunet. 2006. Litterfall of epiphytic macrolichens in Nothofagus forests of northern Patagonia, Argentina: Relation to stand age and precipitation. Austral Ecology 31: 301-309.

Callaway, R.M. 1997. Positive interactions in plant communities and the individualistic- continuum concept. Oecologia 112: 143-149.

Callaway, R.M., and L.R. Walker. 1997. Competition and facilitation a synthetic approach to interactions in plant communities. Special Feature, Ecology 78: 1958-1965.

Campbell, J. and D.S. Coxson. 2001. Canopy microclimate and arboreal lichen loading in subalpine spruce-fir forest. Canadian Journal of Botany 79: 537-555.

Campbell, N.A., J.B. Reece, and L.G Mitchell. 1999. Biology Fifth Edition. Addison-Wessley, CA. USA.

Chase, J. 2005. Towards a really unified theory for metacommunites. Functional Ecology 19: 182-186.

Colwell, R.K., C.X. Mao, and J.Chang. 2004. Interpolating, extrapolating, and comparing incidence-based species accumulation curves. Ecology 85: 2717-2727.

Clements, F.E. 1936. Nature and structure of the climax. Journal of Ecology 24: 252-284. Clinton, P.W., R.B. Allen, and M.R. Davis. 2002. Nitrogen storage and availability during stand

development in a New Zealand Nothofagus forest. Canadian Journal of Forest Research 32: 344-352.

Connell, J.H. 1978. Diversity in tropical rain forests and coral reefs. Science 199: 1302-1310. Connell, J.H. 1980. Diversity and the coevolution of competitors, or the ghost of competition

past. Oikos 35: 131-138.

Cornelissen, J.C. and H. ter Steege. 1989. Distribution and ecology of epiphytic bryophytes and lichens in dry evergreen forest of Guyana. Journal of Tropical Ecology 5: 131-150. Coxson, D.S., M.R. Webber and K.A. Kershaw. 1984. The thermal operating environment of

corticolous and pendulous tree lichens. The Bryologist 87: 197-202.

Coxson, D.S., and S.K. Stevenson. 2007. Growth rate responses of Lobaria pulmonaria to canopy structure in even-aged and old-growth cedar-hemlock forests of central-interior British Columbia, Canada. Forest Ecology and Management 242: 5-16.

Coxson, D., S. Stevenson and J. Campbell. 2003. Short-term impacts of partial cutting on lichen retention and canopy microclimate in an Engelmann spruce-subalpine fir forest in north- central British Columbia. Canadian Journal of Forest Restoration 33: 830-841.

Craig, J., S. Anderson, M. Clout, B. Creese, N. Mitchell, J. Ogden, M. Roberts, and G. Ussher. 2000. Conservation issues in New Zealand. Annual Reviews in Ecological Systematics. 31: 61-78.

Crowley, P.H., H.M Davis, A.L Ensminger, L.C. Fuselier, J.K. Jackson, and D.N. McLetchie. 2005. A general model of local competition for space. Ecology Letters 8: 176-188. Dahlman, L., and K. Palmqvist. 2003. Growth in two foliose tripartite lichens, Nephroma

arcticum and Peltigera aphthosa: empirical modeling of external vs. internal factors.

Functional Ecology 17: 821-831.

Dale, M. 1999. Spatial pattern analysis in plant ecology. Cambridge University Press, Cambridge.

Davis, R. M., R.B. Allen, and P. W. Clinton. 2004. The influence of N addition on nutrient content, leaf carbon isotope ratio, and productivity in a Nothofagus forest during stand development. Canadian Journal of Forest Research 34: 2037-2048.

De Candolle, A.P. 1798. Premier essai cur la nutrion de lichens. Lametherie Journal de Physique, de Chemie et d’Histoire naturelle. 4: 107-116.

DeLucia, E.H., M.H. Turnbull, A.S. Walcroft, K.L. Griffins, D.T. Tissue, D. Glenny, T.M. McSeveny and D. Whitehead. 2003. The contribution of bryophytes to the carbon exchange for a temperate rainforest. Global Change Biology 9: 1158-1170. Denison, W.C. 1988. Culturing the lichens Lobaria oregana and L. pulmonaria on nylon

monofilament. Mycologia 80: 811-814.

Derr, C.C., B. McCune and L.H. Geiser. 2007. Epiphytic macrolichen communities in Pinus

contorta peatlands in southeastern Alaska. The Bryologist 110: 521-532.

Dettki, H. and Esseen P. 1998. Epiphytic macrolichens in managed and natural forest landscapes: a comparison at two spatial scales. Ecography 21: 613-624.

Diamond, J. and T.J. Chase (eds). 1986. Community Ecology. Harper& Row, New York. Dickinson, K.J.M., A.F. Mark, and B. Dawkins. 1993. Ecology of lianoid/epiphytic

communities in coastal podocarp rain forest, Haast Ecological District, New Zealand. Journal of Biogeography 26: 687-705.

Ellis, C.J. and B. J. Coppins. 2006. Contrasting functional traits maintain lichen epiphyte diversity in response to climate and autogenic succession. Journal of Biogeography 33: 1643-1656.

Ellis, C.J., B.J. Coppins, and T.P. Dawson. 2007. Predicted response of lichen epiphyte

Lecanora populicola to climate change scenarios in a clean-air region of Northern Britain.

Biological Conservation 135: 396-404.

Elix, J.A., and E. Stocker-Wörgötter. 2008. Biochemistry and secondary metabolites. Pp. 104- 133 in T. H. Nash III ed., Lichen Biology second edition. Cambridge University Press, Cambridge.

Emmerick, R., I. Giez, O.L. Lange, and P. Proksch. 1993. Toxicity and antifeedant activity of lichen compounds against the polyphagous insect Spodoptera littoralis. Phytochemistry 33: 1389-1394.

Esseen, Per-Anders. 2006. Edge influence on the old-growth forest indicator lichen Alectoria

sarmentosa in natural ecotones. Journal of Vegetation Science 17: 185-194.

Fowler, J., L. Cohen, and P. Jarvis. 2008. Analysis of variance - ANOVA. Pp. 179-209 in J. Fowler, L. Cohen, and P. Jarvis, eds. Practical statistics for field biology second edition. John Wiley & Sons, West Sussex.

Imaging Software to Extract Canopy Structure and Gap Light Transmission Indices From True-Colour Fisheye Photographs, Users Manual and Program Documentation. Simon Fraser University/The Institute of Ecosystem Studies, Burnaby, BC/Millbrook, NY. Galloway, D.J. 1974. A bibliography of New Zealand lichenology. New Zealand Journal of

Botany 12: 397-422.

Galloway, D.J. 1985. Flora of New Zealand lichens. P.D. Hasselberg, Government Printer, Wellington.

Galloway, D.J. 1988. Studies in Pseudocyphellaria (lichens) I. The New Zealand species. Bulletin of the British Museum Natural History (Botany) 17: 1-267.

Galloway, D.J. 1994. A bibliography of New Zealand lichenology. 3 Additions, 1984-1992. New Zealand Journal of Botany 32: 1-10.

Galloway, D.J. 1995a. Líquenes del bosque templado de Chile. Pp. 101-112 in Ecología de los bosques nativos de Chile. J.J. Armesto, C. Villagran and M.K. Arroyo eds. Editorial Universitaria, Santiago de Chile.

Galloway, D.J. 1995b. Lichens in Southern Hemisphere temperate rainforest and their role in maintenance of biodiversity. Pp. 125-135. in Microbial Diversity and Ecosystem Function. D. Allsopp, R.R. Colwell and D.L. Hawksworth eds. CAB International, Wallingford.

Galloway, D.J. 1996. Lichen biogeography. Pp. 315-335. in T.H. Nash II ed., Lichen Biology 2nd edition. Cambridge University Press, Cambridge.

Galloway, D.J. 2007. Flora of New Zealand lichens. Revised second edition including lichen- forming and lichenicolous fungi. Manaaki Whenua Press, Lincoln.

Galloway, D.J. 2008. Godley Review Austral lichenology: 1690-2008. New Zealand Journal of Botany 46: 433-521.

Gaston, K.J. 1994. Rarity. Chapman & Hall, London.

Gauslaa, Y. 1995. The Lobarion, an epiphytic community of ancient forests threatened by acid rain. Lichenologist 27: 59-76.

Gauslaa, Y. 2006. Trade-offs between reproduction and growth in the foliose old forest lichen Lobaria pulmonaria. Basic and Applied Ecology 7: 455-460.

Gauslaa, Y., and K.A. Solhaug. 1996. Differences in the susceptibility to light stress between epiphytic lichens of ancient and young boreal forest stands. Functional Ecology 10: 344- 354.

Gauslaa, Y., and K.A. Solhaug. High- light intensity damage to the foliose lichen Lobaria

pulmonaria within a natural forest: the applicability of chlorophyll fluorescence methods.

Lichenologist 32: 271-289.

Gauslaa, Y., and E.M. Ustvedt. 2003. Is parietin a UV-B or a blue-light screening pigment in the lichen Xanthoria parietina? Photochemical and Photobiological Sciences 2: 424-432. Gauslaa, Y., M. Lie, K.A. Solhaug, and M. Ohlson. 2006. Growth and ecophysiological

acclimation of the foliose lichen Lobaria pulmonaria in forests with contrasting light climates. Oecologia 147: 406-416.

Gauslaa, Y., K. Palmqvist, K.A. Solhaug, H. Holien, O. Hilmo, L. Nybakken, L.C. Myhre, and M. Ohlson. 2007. Growth of epiphytic old forest lichens across climatic and successional gradients. Canadian Journal of Forest Research. 37: 1832-1845.

Gerson, U. 1973. Lichen-arthropod associations. Lichenologist 5: 434-443.

Giordani, P. 2006. Variables influencing the distribution of epiphytic lichens in heterogeneous areas: A case study for Liguria, NW Italy. Journal of Vegetation Science 17: 195-206. Gotelli, N.J. 2001. A primer of Ecology third edition. Sinauer Associates, Inc. Sunderland,

Massachusetts.

Gotelli, N.J. and R.K. Colwell. 2001. Quantifying biodiversity: procedures and pitfalls in the measurement and comparison of species richness. Ecology Letters 4: 379-391.

Gotelli, N. and G. Graves. 1996. Null Models in Ecology. Smithsonian Institution Press, Washington.

Goward, T. 1994. Status report on the oldgrowth specklebelly Pseudocyphellaria rainierensis in Canada. COSEWIC committee on the status of endangered wildlife in Canada.

Ottawa,Ontario.

Goward, T. 1998. Observations on the ecology of the lichen genus Bryoria in high elevation conifer forests. Canadian Field- Naturalist 112: 496-501.

Goward, T. and A. Arsenault. 2000. Cyanolichen distribution in young unmanaged forests: A dripzone effect? The Bryologist 103: 28-37.

Green, T.G.A. and Lange, O.L. 1991. Ecophysiological adaptations of the lichen genera

Pseudocyphellaria and Sticta to south temperate rainforests. Lichenologist 23: 267-282.

Green, T.G.A., J. Horstmann, H. Bonnett, A. Wilkins and W.B. Silvester. 1980. Nitrogen Fixation by member of Stictaceae (Lichenes) of New Zealand. New Phytologist 84: 339- 348.

Green, T.G.A., B. Budel, U. Heber, A. Meyer, A. Zellner, O.L.H. Lange. 1993. Differences in photosynthetic performance between cyanobacterial and green algal components of lichen photosymbiodemes measured in the field. New Phytologist 125: 723-731.

Green T.G.A., T. H. Nash III, and O. L. Lange. 2008. Physiological ecology of carbon dioxide exchange. Pp. 152-181 in T. H. Nash III ed., Lichen Biology 2nd edition. Cambridge University Press, Cambridge.

Griffen, M. and J.G. Conran. 1994. Ecology of the corticolous lichens on Pinus radiata at five sites of increasing age near Linton, Victoria, Australia. Australian Journal of Ecology 19: