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Bulletin of the British Museum (Natural History)

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LICHEN GENUS MICAREA IN EUROPE 19<br />

eluded from <strong>the</strong> genus and <strong>the</strong> new combinations, Psilolechia clavulifera (Nyl.) and Bacidia prasinata<br />

(Tuck.), are proposed. Keys for <strong>the</strong> identification <strong>of</strong> all <strong>the</strong> accepted European taxa are given. The taxonomic<br />

parts are preceded by an outline <strong>of</strong> <strong>the</strong> historical background to <strong>the</strong> study <strong>of</strong> Micarea, and details <strong>of</strong><br />

materials and methods employed in this study. Detailed accounts <strong>of</strong> <strong>the</strong> morphology, chemistry, and<br />

ecology in <strong>the</strong> genus are provided , and a discussion <strong>of</strong> distributions is supported by maps for <strong>the</strong> <strong>British</strong> taxa.<br />

All Micarea species occur on acidic, nutrient poor substrata, and most are confined to cool-temperate,<br />

boreal, or oceanic regions; a few occur in arctic-alpine areas but <strong>the</strong> genus is poorly represented in<br />

dry, lowland, Mediterranean regions. Prior to this study, most <strong>of</strong> <strong>the</strong> species were little-known or<br />

misunderstood; clarification <strong>of</strong> <strong>the</strong>ir taxonomy has been achieved by paying particular attention to <strong>the</strong>ir<br />

anamorphic states, chemistry (including pigmentation), and detailed anatomy. Consideration <strong>of</strong> <strong>the</strong><br />

distribution and ecology <strong>of</strong> <strong>the</strong> species has proved invaluable in ordering <strong>the</strong> taxonomic chaos which previously<br />

surrounded <strong>the</strong> notoriously variable species <strong>of</strong> <strong>the</strong> genus.<br />

Lecideaceae<br />

Historical background<br />

Until recently <strong>the</strong> circumscription <strong>of</strong> <strong>the</strong> Lecideaceae (and <strong>of</strong> <strong>the</strong> genera within it) had changed<br />

httle from that adopted by Zahlbruckner (1926). It included most Uchens with <strong>the</strong> following<br />

combination <strong>of</strong> characters: a crustose to squamulose thallus, a 'grass-green' phycobiont<br />

(excluding Trentepohlia and Phycopeltis) , ± immersed to sessile, disc-like apo<strong>the</strong>cia without a<br />

thalhne margin, mainly colourless spores, and an absence <strong>of</strong> parietin (or related pigments) and<br />

(or) polarilocular spores. The principal genera in <strong>the</strong> family (e.g. Lecidea, Catillaria, Bacidia,<br />

Biatorella, Mycoblastus, Lopadium, Bombyliospora, and Toninia) were separated mainly on<br />

<strong>the</strong> basis <strong>of</strong> spore characters, i.e. size, septation, and number per ascus. This classification<br />

largely ignored many features which (according to modern mycological concepts) now merit<br />

careful consideration, although <strong>the</strong>y were used to varying degrees for <strong>the</strong> delimitation <strong>of</strong> species<br />

or infrageneric categories above this rank. In brief, <strong>the</strong>se features involve <strong>the</strong> structure <strong>of</strong> asci,<br />

excipular and hypo<strong>the</strong>cial tissues, paraphyses and anamorphs, ontogeny, finer aspects <strong>of</strong> thallus<br />

structure, and nature and location <strong>of</strong> pigments and lichen substances. In addition, investigations<br />

<strong>of</strong> <strong>the</strong> phycobiont(s) and considerations <strong>of</strong> ecology and phytogeography <strong>of</strong>ten provide valuable<br />

supplementary information. However, <strong>the</strong> use <strong>of</strong> some <strong>of</strong> <strong>the</strong>se features in an attempt to define<br />

more natural genera is not a purely recent phenomenon. Several lichenologists working in <strong>the</strong><br />

1850s and 1860s made bold attempts in this direction. With regard to <strong>the</strong> Lecideaceae s. lat., two<br />

lichenologists deserving special mention are G. W. Korber (who introduced Lecidella, Lopadium,<br />

Pyrrhospora, Schaereria, Schadonia, and Steinia) and A. B. Massalongo (who introduced<br />

Catillaria, Psilolechia, Sarcosagium, Scoliciosporum, Strangospora, and Toninia). In <strong>the</strong> latter<br />

half <strong>of</strong> <strong>the</strong> 19th century lichenology came under <strong>the</strong> almost monarchical influence <strong>of</strong> William<br />

Nylander, whose simplistic generic concepts gained precedence over <strong>the</strong> more far-sighted works<br />

<strong>of</strong> Korber, Massalongo, and o<strong>the</strong>rs. From <strong>the</strong> 1870s right up to <strong>the</strong> 1950s <strong>the</strong>re were few<br />

attempts to reassess <strong>the</strong> generic concepts <strong>of</strong> Nylander or <strong>the</strong> slightly more complex, but no less<br />

artificial, system <strong>of</strong> Zahlbruckner. Between about 1929 and 1954 <strong>the</strong> French lichenologist M. G.<br />

B. Choisy resurrected many <strong>of</strong> <strong>the</strong> old and more or less forgotten genera, and created several<br />

new ones (e.g. Haplocarpon [= Huilia], Hypocenomyce, Trapelia, and Tremolecia). Unfortu-<br />

nately, Choisy's works made little impact at <strong>the</strong> time and it was not until <strong>the</strong> mid-1960s that<br />

lichenologists began to look more carefully at <strong>the</strong> delimitation <strong>of</strong> genera. Recent investigations<br />

have led to <strong>the</strong> reinstatement (although <strong>of</strong>ten with emendations) <strong>of</strong> many <strong>of</strong> <strong>the</strong>se genera and<br />

many new genera have had to be described (e.g. Fuscidea, Herteliana, Melanolecia, Trapeliop-<br />

sis, Tylothallia, and Vezdaea). Most are included in <strong>the</strong> key to European lichen genera in Poelt<br />

& Vezda (1981). Despite <strong>the</strong> many advances made during <strong>the</strong> last 15 years, it will be several<br />

decades before a reasonably natural generic classification within <strong>the</strong> Lecideaceae s. lat. is<br />

achieved. The enormity <strong>of</strong> <strong>the</strong> task can be appreciated from <strong>the</strong> fact that Zahlbruckner<br />

(1921-40) accepted no less than 1450 species in <strong>the</strong> genus Lecidea alone! In addition to <strong>the</strong> high<br />

number <strong>of</strong> taxa involved , fur<strong>the</strong>r problems arise from <strong>the</strong> locating and obtaining on loan suitable<br />

(including type) material, and <strong>the</strong> many difficulties in observing and interpreting many<br />

microscopical, morphological, and ontogenetic features.

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