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The Lichens of Ireland: An Illustrated Introduction to Over 250 Species PDF

161 Pages·2011·284.452 MB·English
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Lichens of Ireland An illustrated introduction to over 250 species Paul Whelan 154-The Lichens of Ireland-update.indb 3 19/07/2012 22:24 Lichens of Ireland An illustrated introduction to over 250 species Paul Whelan has a degree in Biology from University College Dublin and studied Computer Science and Art History at Trinity College Dublin. He published several books on Computer Aided Design in the 1990s before returning to his earlier interest in natural history and focusing on Ireland’s lichen flora. In 2008 he became involved in the LichenIreland project and developed an independent website (www. lichens.ie), which susbequently won an Irish Internet Association Net Visionary Award. In 2010 he developed www.ispynature.com for primary school children. Paul lives in east Cork. 154-The Lichens of Ireland-update.indb 1 19/07/2012 22:24 For Anna, Aine, Molly and Pip. Ernst Haeckel (1834–1919), lichens from Kunstformen der Natur (1904). [Cladia retipora, Cladonia perfoliata, Cladonia cervicornis ssp. verticillata, Cladonia squamosa, Cladonia fimbriata, Cladonia cornucopiae, Lobaria pulmonaria, Physcia aipolia, Melanohalea olivacea, Flavoparmelia caperata, Anaptychia crinalis] 154-The Lichens of Ireland-update.indb 2 19/07/2012 22:24 Contents Acknowledgements iv Introduction v Part One – Biology and Ecology What are lichens? 2 Defining a lichen – A little more detail – Damsel–tyrant duality Naming lichens – Mycobionts: fungi – Photobionts: algae and cyanobacteria Morphs – Thallus anatomy Growth forms and colours 7 Growth forms – Crustose lichens – Foliose lichens – Fruticose lichens Squamulose lichens – Lichen colours Lichen sexual reproduction 12 Sexual reproduction in ascomycete lichens: Life cycle of ascomycete lichens – Apothecia – Structure of an apothecium Apothecial variations – Perithecia – Spore size and the micron Basidiomycete lichens Spore dispersal Asexual reproduction (vegetative) 17 Fragmentation – Soredia (and soralia) – Isidia – Schizidia – Pycnidia – Blastidia Non-reproductive bits A practical approach 22 A guide to basic equipment – Setting up your own herbarium Vice-county maps – Lichen substances and spot tests Spot tests and how to apply them – Ultraviolet light (UV) – Microscopes How to section an apothecium – Spore septa and spore size Ecology of lichens 26 Ireland’s geology – Where are these rocks? – Climate and substrates Oceanic climate – Lichen substrates – Saxicolous lichens – Corticolous lichens Bryophytes and dead vegetation Lichen habitat types in Ireland Lichens and pollution – Importance of lichens in an ecosystem Some conservation pointers Lichen communities and habitats: Lobarion community – Graphidion community – Parmelion community The Burren Limestone – Atlantic Woodlands, Killarney – Atlantic hazel woods Rocky seashores Part Two – Alphabetic Photographic Guide to Selected Lichens Species descriptions 42 References 152 Internet Resources 153 154-The Lichens of Ireland-update.indb 5 19/07/2012 22:24 Acknowledgements In particular I would like to thank John Douglass, lichenologist, for many enjoyable and informative field trips throughout Ireland. Thank you also to Howard Fox, lichenologist with the National Botanic Gardens, Glasnevin, Dublin, for his encouragement, and to the LichenIreland team for involving me in field work. LichenIreland is supported by National Parks and Wildlife Service; National Botanic Gardens, Glasnevin; Environment and Heritage Service and the Ulster Museum, National Museums Northern Ireland. I would like to express my thanks to Alan Orange, the National Museum of Wales, for use of his wonderful ink drawings which help to elucidate much of the lichen terminology, and to both the British Lichen Society and LichenIreland for access to their records of Ireland’s lichens at 10km resolution. Special thanks must be given to the fieldworkers who walked the length and breadth of Ireland over many years amassing more than 130,000 records of our rich lichen biodiversity. These lichenologists include Andy Acton, Brian Coppins, Sandy Coppins, Simon Davey, John Douglass, Howard Fox, Vince Giavarini, Joe Hope, Alan Orange, Ivan Pedley, David Richardson, Neil Sanderson, Jenny Seawright, Mike Simms, Robert Thompson and Stephen Ward. Also, thank you to Gill Weyman of the Irish Wildlife Trust, Anna O’Connor and Paula Meenehan, for reading the text and making suggestions and also to Jenny Seawright for helping me locate some species to photograph for the book. And thanks to Nico Nieuwstraten, who wrote a wonderful Photoshop script file to generate the vice-county maps from the vice-county data. Finally, I would like to express my gratitude to the Heritage Council of Ireland for financial support under the 2011 Heritage  Education,  Community  and  Outreach scheme, without whom this book would not have been published. Paul Whelan October 2011 154-The Lichens of Ireland-update.indb 6 19/07/2012 22:24 Introduction Currently, Ireland is home to almost 1,200 recorded lichens and 205 lichenicolous and non- lichenized fungi. Considering that the whole of North America has approximately 3,600 recorded species, Ireland’s lichen biodiversity has a staggering richness for a country of its size and latitude. This hidden richness of our natural world was relatively neglected until recently, when the LichenIreland Project (2005–2010) began to uncover the extent of the biodiversity, particularly of our Atlantic woodlands and hyper-oceanic hazel woods. This book is the result of my involvement in the LichenIreland project. It introduces beginners, armed with a simple hand lens, to the delightful Lilliputian world of lichens. There could be much debate as to which species should be included in a book of this nature and size. Many will feel that species ‘X’ should have been included. However, a selection had to be made and included are some notable species that thrive in our oceanic climate, and some dots-and-squiggles lichens that are rather difficult to identify. All will contribute in some way to challenge beginners, and prepare them for more advanced texts such as Frank Dobson’s Lichens (6th edition) or The Lichens of Great Britain and Ireland, 2009 edition (British Lichen Society). The book lacks a key. For this I make no apology as a key would take the text beyond its current scope, and possibly lose the readership for which it is intended. The book is necessarily brief and sometimes simplified, but will hopefully turn the minds of Ireland’s new generation of young naturalists to the world of lichenology. A downloadable glossary (pdf) to the book is currently available at www.lichens.ie. The lichens are listed alphabetically. This method was chosen after consideration of other possible arrangements. Presentation by body form was rejected because that classification system quickly breaks down (for example Cladonia lichens belong to both squamulose and fruticose forms) and classification by habitat breaks down because many species straddle more than one habitat. Similarly, classification by substrate breaks down although it was a system I gave a lot of consideration to. Alphabetic arrangement provides more immediate rewards because the beginner will quickly learn to recognise the genera and can flick to the relevant section efficiently. All the photographs in the text were taken by the author over a three-year period of lichen hunting around Ireland. The magnified insets on many images act to either show more detail or to concentrate the eye on where to look for characteristic features (not necessarily magnified). With just a hand lens and a willingness to understand lichens, you will open up a miniature world of immense beauty. It is almost inevitable that a beginner using a hand lens for the first time will consider him or herself in the shoes of Alice in Wonderland. Some of the earliest Irish lichen records were made c. 1696 by William Sherard and Caleb Threlkeld (1676–1728). In the twentieth century Matilda Knowles made significant contributions with her work on seashore zonation and her 1929 publication History of Irish Lichens (available on www. lichens.ie). Other twentieth century contributions were made by Lilian Porter, Robert Lloyd Praeger, Annie Lorrain Smith, John Adams and more recently Howard Fox. The LichenIreland project of 2005–2010 made a major contribution to documenting Irish lichen flora. At the time of writing a Lichen Red Data list is imminent. Nomenclature in this text is mainly according to Smith et al. (2009) and Hawksworth (2003). 1 154-The Lichens of Ireland-update.indb 1 19/07/2012 22:24 What are lichens? Lichens contribute to the visual ‘background noise’ of our environment, providing colour and texture to woodlands, parks, buildings, monuments and graveyards. They add the ‘old’ to old walls, gateposts and monuments, their aesthetic value only missed when they are absent. Lichens have been used as medicines, poisons and dyes and have contributed to making many a child’s doll in the form of hair or bulking out material for arms and legs. Even today they are used in the manufacture of perfume, acting as a fixative to ensure the slow release of scent. Evernia prunastri has been used to give some perfumes a moss-like scent. Cladonia pyxidata was once thought to cure whooping cough. Dyes can be extracted from lichens and used to colour wool. Extraction sometimes involved boiling or soaking them in urine. During the nineteenth century lichens became economically valuable, often demanding prices in excess of the cost of rare spices. A large cottage industry developed around lichen dyes. Lichens act as a source of nitrogen for many plants and are frequently the first living forms to inhabit bare rock. Birds use them as nesting material and they are an essential component in food chains, being eaten by snails, slugs, insects and higher animals such as deer. Lichens act as biomonitors, Crustose lichens add texture and colour to headstones in a west- some absorbing heavy metals from their of-Ireland graveyard. They enhance the aesthetic quality of many environment and responding to both buildings and monuments. air and water quality. Defining a lichen Lichens (phonetic pronunciation ‘like en’ or ‘litch en’, both acceptable) are fungi that live with a photosynthesizing partner, usually an alga or cyanobacteria. It is a clever arrangement because many fungi are saprophytes (they feed on dead organic matter such as dead plants or animals) and the algae and cyanobacteria are autotrophs, able to make their own food from sunlight, water and carbon dioxide (photosynthesis). With such a dual food source, they are unlikely to starve. A little more detail A standard textbook definition describes lichens as two and sometimes three organisms living together in symbiosis. This is essentially true. The three organisms in question are a fungus (related to bread mould, although a few are related to mushrooms), algae (related to seaweed or the green slime on ponds) and in some cases photosynthesizing bacteria belonging to a group called cyanobacteria. The relationship is stable and they form a distinct shape and structure (both used as identification aids) and generally live a long time. Some lichens have been estimated to be 4,500 years old. Lichen-type symbiosis has been positively identified in fossil records as old as 410 million years (Rhynie chert in Scotland). Recent studies have pushed this back to 600 million years. 2 what are lichens? 154-The Lichens of Ireland-update.indb 2 19/07/2012 22:25 In 1982 the International Association of Lichenology issued the following definition of a lichen: an ‘association of a fungus and a photosynthetic symbiont resulting in a stable thallus of specific structure’. In summary, a lichen is not a single organism, but an assemblage of two or three different species working together for survival. The fungal part is termed the mycobiont and the photosynthetic partner is the photobiont. Damsel–tyrant duality The dual nature of lichens was first recognized in 1869 by a German biologist, Simon Schwendener. There was great resistance to this idea at the time. He wrote: ‘lichens are ... colonies ... of individuals, of which one alone plays the master, while the rest, forever imprisoned, prepare the nutrients for themselves and their master. This fungus is a fungus of the class Ascomycetes, a parasite which is accustomed to live upon others’ work. Its slaves are green algae, which it has sought out ... and compelled into its service.’ A further description around the same time, by Rev. James Crombie, a Scotsman, described the dual relationship as Simon Schwendener (1829– ‘the unnatural union between a captive algal damsel and tyrant fungal 1919) who first postulated master’. Regardless of which of these descriptions you favour, it was the dual nature of lichens. determined that the fungal partner was the dominant one. Perhaps a This idea was rejected by more concise definition of a lichen is ‘a fungus that associates with a many scientists of the time. photobiont’. A fungus makes up the bulk of the lichen body, occupying 80% or more of the thallus; frequently the algal part is as low as 5%. Whether the fungus enslaves the alga or cooperation is mutual is a matter of conjecture. Both partners seem at ease with one another, although the alga has had to give up its own sex life. Only the ‘tyrant’ fungus is the sexually active partner. Simon Schwendener also suggested that lichens should be classified with fungi, but this was rejected by the scientific community. Since the 1950s lichens are accepted as lichenized fungi and have been accepted into the Kingdom Fungi. Despite this, the study of lichens has been considered a separate study (lichenology) from the study of fungi (mycology). Naming lichens Carl von Linné (Linnaeus) introduced a binomial system for naming plants and animals in the eighteenth century. Names such as Ochrolechia parella or Homo sapiens are binomial. Latin was then in use as the universal scientific language. Binomial names are derived from Latin or Greek words and written in italics when printed and underlined when handwritten. The first part of the name is the genus or group to which the organism belongs and the second part of the name is the species (parella or sapiens in the above examples) and is written in lowercase letters. Now, while Linnaeus made major progress with naming just about every known plant and animal at the time, lichens confounded him, Carl von Linné (1707–1793) although he did manage to describe 109 different species. Overall, he described 109 lichens and referred to them as ‘nature’s had little time for lichens and referred to them using the Latin phrase paupers’. rustici pauperrimi (nature’s paupers). As a lichen is an assemblage of two or three distinct species, each with its own binomial name, how do we name it? The name of a lichen is the name of the fungal partner. Consequentially, a name such as Ochrolechia parella refers to the fungus that makes up the lichen; the photobiont does not usually contribute to the lichen’s name. naming lichens 3 154-The Lichens of Ireland-update.indb 3 19/07/2012 22:25 Mycobionts – fungi Th e fungal partner in a lichen is termed the mycobiont. Fungi are a large and diverse group of organisms, comprising at least 70,000 species worldwide. Some fungal experts say the number is more likely to be 1.4 million species. Fungi are the primary ‘decomposers’ or saprophytes, in the biosphere. Many fungi are also parasites, feeding on living cells and tissues. Both animals and green plants are constructed of individual block-shaped cells. Fungi, on the other hand, are composed of thread-like fi laments. Th is Photograph showing a mould growing on food (jam), illustrating fi lamentous structure has contributed the thread-like nature of fungal cells. In many crustose-type lichens the outer growth area or prothallus is similar. largely to their success as a group, allowing them to grow into crevices, enabling access to carbohydrate and protein molecules of their hosts. Nearly all fungi that form lichens in Ireland come from one group, the Ascomycetes or ‘sac fungi’, so called because they produce sexual reproductive spores in sacs or asci. Only a handful of lichens in Ireland (less than 5%) have the mushroom- type fungi (group Basidiomycetes) as a mycobiont. Th ese fungi do not produce spores in sacs, but instead on club- shaped structures, the basidia. In this book, the terms ‘spores’ and ‘ascospores’ A crustose lichen showing the prothallus or outer growing area. are used interchangeably. Th e orange-red colour is a free-living Trentepohlia alga. Photobionts – algae and cyanobacteria Photobionts use light to make molecules of glucose. Th is process of converting electromagnetic energy into chemical energy is termed photosynthesis. All green plants photosynthesize, as do all algae and cyanobacteria. Many lichens contain both algal and cyanbacterial photobionts in the same thallus; however, one is dominant and is termed the primary photobiont. Algal photobionts contribute sugars to the fungus; cyanobacteria also provide sugars, but additionally supply nitrogen in a form (nitrates) the fungus can use to build up its body tissue (mainly proteins). Algae belong to the Kingdom Protista and those found in lichens belong to the genera Trentepohlia (orange in colour) and Trebouxia (bright green in colour). Th e group Trebouxia are unicellular and by far the most common green algae integrated into lichens, but rarely found free- living. Trentepohlia algae, on the other hand, grow freely outside lichens as fi laments, but when part of the lichen symbiosis they become single cells. Crustose lichens tend to have Trentepohlia algae as their photobiont. Lichens whose only photobiont is green algae are termed chlorolichens. A group called the ‘jelly lichens’ have a predominance of cyanobacteria (their primary photobiont) and belong to the genera Collema and Leptogium. Nostoc cyanobacteria are found in jelly lichens. 4 mycobionts 154-The Lichens of Ireland-update.indb 4 19/07/2012 22:25

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