23.07.2013 Views

Bulletin of the British Museum (Natural History)

Bulletin of the British Museum (Natural History)

Bulletin of the British Museum (Natural History)

SHOW MORE
SHOW LESS

You also want an ePaper? Increase the reach of your titles

YUMPU automatically turns print PDFs into web optimized ePapers that Google loves.

LICHEN GENUS MICAREA IN EUROPE 61<br />

Paraphyses<br />

The paraphyses in Micarea are characteristically thin and branched. When measured (in 10%<br />

KOH, or ammoniacal erythrosin) at <strong>the</strong> mid-hymenium <strong>the</strong>y may be very thin and only about<br />

0-6-1 fxm wide (e.g. M. anterior, M. botryoides, M. contexta, M. eximia, M. lithinella, M.<br />

misella, and M. prasina), or thin and c. 1-1-5 /xm wide (e.g. M. adnata, M. cinerea, M. denigrata,<br />

M. intrusa, M. muhrii, and M. peliocarpa), or relatively stout and about 1-5-1-8 fjun wide (e.g.<br />

M. assimilata, M. incrassata, M. lignaria, M. osloensis, and M. subnigrata) ; <strong>the</strong>se measurements<br />

relate to paraphyses not coated in pigment. In old, much expanded, apo<strong>the</strong>cia <strong>the</strong> paraphyses<br />

sometimes appear 'stretched' and thinner than normal (especially in <strong>the</strong> lower half <strong>of</strong> <strong>the</strong><br />

hymenium) ; this phenomenon has frequently been observed in collections <strong>of</strong> M. bauschiana, M.<br />

sylvicola, and M. lignaria. In many species <strong>the</strong> paraphyses gradually widen towards <strong>the</strong>ir apices,<br />

but <strong>the</strong> apices are never regularly clavate or capitate. This widening is <strong>of</strong>ten enhanced by <strong>the</strong><br />

deposition <strong>of</strong> closely adhering pigment which sometimes gives <strong>the</strong> appearance <strong>of</strong> a 'hood' (e.g.<br />

M. melaena) ; such coatings or hoods can be detached by gently boiling and <strong>the</strong>n tapping sections<br />

or squash preparations in 50% KOH. In a few cases (e.g. M. melanobola) <strong>the</strong> pigment cannot be<br />

separated in this way and appears to be located within <strong>the</strong> walls <strong>of</strong> <strong>the</strong> paraphyses. Paraphyses<br />

with dark pigmented apical 'caps', like those found in Catillaria s. str. (Killias, 1980: 253),<br />

Buellia, and many species <strong>of</strong> Lecanora, are not known in Micarea.<br />

In all species <strong>of</strong> Micarea a large proportion <strong>of</strong> <strong>the</strong> paraphyses are branched, even if <strong>the</strong><br />

branching is mainly confined to <strong>the</strong> epi<strong>the</strong>cium. Species with sparingly branched paraphyses<br />

include M. assimilata, M. incrassata and M. lignaria. Anastomozing paraphyses have been<br />

observed in all <strong>the</strong> species, but <strong>of</strong>ten <strong>the</strong> anastomoses are ± confined to <strong>the</strong> lower third <strong>of</strong> <strong>the</strong><br />

hymenium. The degree <strong>of</strong> branching and anastomosing is difficult to quantify for <strong>the</strong> practical<br />

purposes <strong>of</strong> identification, but this character can be useful when comparing collections micro-<br />

scopically.<br />

A similarly difficult character is <strong>the</strong> relative abundance <strong>of</strong> <strong>the</strong> paraphyses. The two extremes<br />

can be referred to as: (a) 'numerous' - large in number and immediately discernible when<br />

observing a mount in 10% KOH at x400; (b) 'scanty' - few in number and not immediately<br />

obvious when observed in <strong>the</strong> same way. In hymenia with scanty paraphyses <strong>the</strong> 'extra space' is<br />

taken up by hymenial gel or a higher proportion <strong>of</strong> asci, or a combination <strong>of</strong> both. The situation<br />

in most species lies somewhere between <strong>the</strong> two extremes. Fur<strong>the</strong>rmore, <strong>the</strong> relative propor-<br />

tions <strong>of</strong> hymenial gel and asci to paraphyses sometimes increases as <strong>the</strong> apo<strong>the</strong>cium expands (cf<br />

<strong>the</strong> above example <strong>of</strong> M. bauschiana, M. sylvicola, and M. lignaria). An accurate assessment <strong>of</strong><br />

<strong>the</strong> abundance <strong>of</strong> paraphyses is not usually essential for <strong>the</strong> routine identification <strong>of</strong> Micarea<br />

species, although it can be helpful when comparing difficult, convergent forms <strong>of</strong> M. denigrata<br />

and M. misella (see couplet 11 <strong>of</strong> <strong>the</strong> main key).<br />

In addition to <strong>the</strong> 'normal' paraphyses described above, <strong>the</strong> hymenium <strong>of</strong> several species (e.g.<br />

M. bauschiana, M. botryoides, M. eximia, M. nigella, M. sylvicola, andM. tuberculata) contain<br />

scattered individuals, or small fascicles, <strong>of</strong> ra<strong>the</strong>r stout paraphyses. These 'paraphyses' are<br />

about 2-3 /xm wide and usually more distinctly septate than normal paraphyses. Fur<strong>the</strong>rmore<br />

(especially in species with a dark hypo<strong>the</strong>cium), <strong>the</strong>y are <strong>of</strong>ten coated in pigment throughout<br />

<strong>the</strong>ir length (Fig. 34), with <strong>the</strong> result that <strong>the</strong> hymenium is seen to be intersected by dark vertical<br />

streaks. They are mainly found in species with 'scanty' paraphyses, and appear to extend deep<br />

into <strong>the</strong> hypo<strong>the</strong>cium. The elucidation <strong>of</strong> <strong>the</strong>ir true status and function awaits detailed<br />

ontogenetic studies, but it is possible that <strong>the</strong>y have a streng<strong>the</strong>ning, spacing or protective<br />

function during <strong>the</strong> maturation <strong>of</strong> <strong>the</strong> hymenium from <strong>the</strong> primary corpus.<br />

Hypo<strong>the</strong>cium<br />

The area <strong>of</strong> tissue lying below <strong>the</strong> hymenium, or between <strong>the</strong> hymenium and <strong>the</strong> excipulum (if<br />

present), is generally referred to by lichenologists as <strong>the</strong> 'hypo<strong>the</strong>cium'. In many groups <strong>of</strong><br />

lichenized discomycetes (including <strong>the</strong> Lecideaceae) this area can be divided into an upper,<br />

usually narrow, layer containing mainly ascogenous hyphae, and a lower, <strong>of</strong>ten much deeper<br />

layer <strong>of</strong> structural tissue (hyphae gelatinised to various extents, according to genus or species).<br />

These two layers are <strong>of</strong>ten <strong>of</strong> different colour or colour intensity. Where <strong>the</strong> two layers are<br />

.

Hooray! Your file is uploaded and ready to be published.

Saved successfully!

Ooh no, something went wrong!