key to lichen genera in thailand with special reference … · nat. hist. bu比.siams配.43:...

34
NAT. HIST. Bu .SIAMS .43:303-335 1995 KEY TO LICHEN GENERA IN THAILAND WITH SPECIAL REFERENCE TO EPIPHYTIC TAXA PART 1:MACROLICHENS P.A.Wolselel and B. Aguirre-Hudson 2 ABSTRACT An introduction to lichens and e 旬ロ凶nologyused to describe them with illus ated plates is followed by an tificialkey to 59 macrolichen genera ofτ'h ailand.Key characters anddis butionof eachgenusaredescribedandsomecommonorch acteristicspecies mentioned. TRODUC ON Li chens 'ebi 'O l 'O gically distinct entities c 'O mp 'O sed 'O f an algal 'O r cyan 'O bacterial p tner (ph 'O t 'O bi 'O nt) and a fungal (myc 'O bi 'O nt) p 紅白erliving in a symbi 'O tic state. The ph 'O t 'O bi 'O nt mayals 'O 'O cc ina 'ee-livingstate and thes nealga may be f 'O und in a wide range 'O f lichens. Th e ph 'O t 'O bi 'O nt isn 'O t kn 'O wnt 'O repr 'O ducesexuallyina lichens 匂.t e.H 'O wever efungalp tnerisspecifict 'O thelichentax 'O n S 'O出at eclassificati 'O n 'O f lichensis based 'O nthe sexual characteristics 'O f efungal partner. H 'O wever the sexual state is r e in m ylichensand vegetative repr 'O ducti 'O niswidespread in m ygenera all 'O wing the algalandfungalp nerst 'O bedispersedt 'O gether. Inthefieldlichentaxac be circumscribed by a number 'O fm 'O甲h 'O l 'O gicalcharacters that are the result 'O f the p tnership 'O f b 'O血 symbi 'O nts ,叩din 'O rdert 'O distinguishgeneradspeciesthesecharacterse widely used in an artificial key. The ph 'O t 'O bi 'O nt issensitive t 'O a range 'O f envir 'O nmen ta1 c 'O nditi 'O ns and w i1l determine the dis buti 'O n 'O f the lichen tax 'O n. In 'O rder t 'O protect the ph 'O t 'O bi 'O ntandextenditsec 'O l 'O gicalr geemyc 'O bi 'O ntshaveev 'O lvedadiversity 'O f chemicalsubstances. Al th 'O ughthese have been used dyeingagentsandasmedicinal remedies(RICHARDSON 1975) f 'O r a l 'O ngtime theunderlyingchemis yhas 'O nlybeen S diedinthelast 120 years anddevel 'O ped仕'O m simpletestsf 'O r c 'O l 'O urreacti 'O nst 'O chemical analysis using s 'O lvents and chr 'O mat 'O gr hyt 'O identi 命曲e 'O rg cc 'O mp 'O nents. Th ese have n 'O wbec 'O me an imp 'O rtant tax 'O n 'O mic t 'O'O lt 'O distinguish genera and species in difficult familiessuchasParmeliaceaeinadditi 'O n t 'O m 'O rph 'O l 'O gicalfeatures. usgeneric key uses a c 'O mbinati 'O n 'O f m 'O中h 'O l 'O gicalcharacters and simple chemical reacti 'O nst 'O identify ec 'O mp 'O nent genera. 百lI sish 'O wever 組側出cialkey and related generad 'O n 'O t necessarilykey 'O ut near each 'O ther. Ther ge 'O f genera included 泊血iskeyisbased 'O nliteraturerec 'O rds om S 'O uth- EastAsia(AGUIRRE-HuDSON & WOLSE Y 1994) '0町'O wnc 'O llecti 'O ns 仕''O m Thailand andmaterialc 'O llected by ese 凶'O rau 血'O r between 1988-1993. Thec 'O llecti 'O ns 'en 'O t I Department of Botany Th e Natural Hi story Museum Cromwell Rd London SW7 5BD 2 Present address: 58 Willowhayne Drive Walton -o n-Thames Surrey KT12 2NR U. K. 303

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Page 1: KEY TO LICHEN GENERA IN THAILAND WITH SPECIAL REFERENCE … · nat. hist. bu比.siams配.43: 303-335,1995 key to lichen genera in thailand with special reference to epiphytic taxa,

NAT. HIST. Bu比 .SIAMS配 .43:303-335,1995

KEY TO LICHEN GENERA IN THAILAND WITH SPECIAL REFERENCE TO EPIPHYTIC TAXA, PART 1: MACROLICHENS

P.A. Wolselel and B. Aguirre-Hudson2

ABSTRACT

An introduction to lichens and出e旬ロ凶nologyused to describe them with illus町atedplates, is followed by an紅 tificialkey to 59 macrolichen genera ofτ'hailand. Key characters and dis凶butionof each genus are described and some common or ch紅 acteristicspecies mentioned.

町TRODUC百ON

Lichens釘'ebi'Ol'Ogically distinct entities c'Omp'Osed 'Of an algal 'Or cyan'Obacterial p紅tner

(ph'Ot'Obi'Ont) and a fungal (myc'Obi'Ont) p紅白erliving in a symbi'Otic state. The ph'Ot'Obi'Ont

may als'O 'Occ町 ina企'ee-livingstate, and the s創nealga may be f'Ound in a wide range 'Of

lichens. The ph'Ot'Obi'Ont is n'Ot kn'Own t'O repr'Oduce sexually in a lichen s匂.te. H'Owever

由efungal p紅 tneris specific t'O the lichen tax'On, S'O出at白eclassificati'On 'Of lichens is

based 'On the sexual characteristics 'Of血efungal partner. H'Owever the sexual state is r紅 e

in m組 ylichens and vegetative repr'Oducti'On is widespread in m佃 ygenera, all'Owing the algal and fungal p制 nerst'O be dispersed t'Ogether. In the field lichen taxa c組 be

circumscribed by a number 'Of m'O甲h'Ol'Ogicalcharacters that are the result 'Of the p紅 tnership

'Of b'O血 symbi'Onts,叩din 'Order t'O distinguish genera阻 dspecies these characters紅 e

widely used in an artificial key. The ph'Ot'Obi'Ont is sensitive t'O a range 'Of envir'Onmenta1 c'Onditi'Ons and wi1l determine the dis凶buti'On'Of the lichen tax'On. In 'Order t'O protect the ph'Ot'Obi'Ont and extend its ec'Ol'Ogical r佃 ge出emyc'Obi'Onts have ev'Olved a diversity 'Of

chemical substances. Alth'Ough these have been used錨 dyeingagents and as medicinal

remedies (RICHARDSON, 1975) f'Or a l'Ong time, the underlying chemis位yhas 'Only been

S加diedin the last 120 years, and devel'Oped仕'Om simple tests f'Or c'Ol'Our reacti'Ons t'O

chemical analysis using s'Olvents and chr'Omat'Ogr叩hyt'O identi命曲e'Org削 cc'Omp'Onents.

These have n'Ow bec'Ome an imp'Ortant tax'On'Omic t'O'Ol t'O distinguish genera and species in

difficult families such as Parmeliaceae in additi'On t'O m'Orph'Ol'Ogical features.

百usgeneric key uses a c'Ombinati'On 'Of m'O中h'Ol'Ogicalcharacters and simple chemical

reacti'Ons t'O identify血ec'Omp'Onent genera.百lIsis h'Owever組側出cialkey and related

genera d'O n'Ot necessarily key 'Out near each 'Other.

The r叩 ge'Of genera included泊血iskey is based 'On literature rec'Ords企omS'Outh-

East Asia (AGUIRRE-HuDSON & WOLSE日Y,1994), '0町'Ownc'Ollecti'Ons仕''OmThailand,

and material c'Ollected by由ese凶'Orau血'Or,between 1988-1993. The c'Ollecti'Ons紅'en'Ot

I Department of Botany, The Natural History Museum, Cromwell Rd, London SW7 5BD 2 Present address : 58, Willowhayne Drive, Walton-on-Thames, Surrey KT12 2NR, U.K.

303

Page 2: KEY TO LICHEN GENERA IN THAILAND WITH SPECIAL REFERENCE … · nat. hist. bu比.siams配.43: 303-335,1995 key to lichen genera in thailand with special reference to epiphytic taxa,

304 P.A. WOLSELEY AND B. AGUlRRE.HUDSON

representative of the geographical or ecologica1 diversity of Thai1and, and it is hoped that this preliminary key to macrolichen genera in Thailand and adjacent紅 easof South-East Asia wi11 stimulate more research in由isarea. In layout and much of its content the key follows The Macrolichen Flora 01 East Africa (Sw町 scow& KROG, 1988) which has contributed to血eidentification of some tropica1 macrolichen genera and species using a combination of morphological, reproductive and chemical characters. A key to the macrolichens of India and Nepa1 (AWAST凹, 1988) has provided much information仕omm 叫acentgeographica1 region. Generic concepts within part of白eParmeliaceae have been based on the recent publication on generic delimitation of Parmeliαsensu lato (ELIX, 1993). Descriptions of key characters used,組dof ecological distribution, where known,

for each genus in Thailand紅 einc1uded in a section at the end of the key. All terr凶nologyas bold type in text is i1lustrated in plates I-IV.

LICHEN STRUCTURE

Thallus Morphology

Lichens fall into morphologically distinct groups which provide a basis for most lichen keys.

1. Fruticose lichens are shrubby, erect or pendulous, and may be attached by a disc or holdfast, or be unattached to a substrate. They may be terete or flattened, hollow or solid and with or without a centra1 cord-like出 is.Branching pattem may be sympodial or dichotomous. Podetia are erect cup-like to branched s回 cturesbearing fruits and arising from a primary tha11us. Found in Cladoniaceae.

2. Foliose lichens have erect or horizonta11y spreading leaf-like lobes, with upper and lower surfaces often differing in colour. Attachment to出esubstrate may be by rhizin回,hapters, or a holdfast. Rhizines may be simple, branched (dichotomous or squarrose) or fasciculate.百leseare distinguished from cilia that occur on出emarg凶sor出eupper surface of the tha11us.

3. Placodioid lichens with c10sely attached lobes radiating from a crust-like centre, without distinct organs of attachment e.g. Dirinaria.

4. Squamulose lichens with scattβred or compact, often overlapping leafy outgrowths or peltate discs attached to the substrate by a hypotha11us, e.g. Phyllopsora.

5. Filamentous lichens with the filaments of the phycobiont surrounded by the hyphae of the mycobiont,出ewhole being found as a loosely arranged matt e.g. Dic砂onema.

6. Crustose lichens are c10sely appressed to the substrate, and attached to it by the hyphae of the medulla. They may grow in a continous crust on出esubstratum, or as rounded or angular areolae. A marginal hypotha11us of unlichenised hyphae may or may not be present. If pr,田ent,出isdistinguishes the area occupied by each individua1 tha11us. The thalli of crustose lichens may also be endolithic (pen巴tratinga rock surface) or endophloeodal (below the bark surface). 百leselichens are the subject of a separate key to be published sep紅 atelyas P制 11.

Within groups 1-5 a range of s回 cωres紅 eused as a basis for the description of lichens, and it is important to be familiar with the terrninology used in order to use the key.

Page 3: KEY TO LICHEN GENERA IN THAILAND WITH SPECIAL REFERENCE … · nat. hist. bu比.siams配.43: 303-335,1995 key to lichen genera in thailand with special reference to epiphytic taxa,

KEY TO LJCHEN GENERA IN THAILAND

Fruticose shrub-like cup-like sympodial branching (Cladonia furcata) (Cladonia humilis) (Usnea baileyii)

podetia

5mm -Foliose (Physconia) sp

~~ 5mm

Squamulose (Phyllopso団 furfuracea)hypothallus

1mm

Rhizines

Placodioid (Dirinaria) sp

時与

1mm

t branched squarrose fasciculate (Physconia sp.) (Peltigera sp.)

Plate I. Thallus morphology

branched dichotomous (Hypotrachyna sp.)

dichotomous branching (Everniastrum nepalense)

クiJW

305

Page 4: KEY TO LICHEN GENERA IN THAILAND WITH SPECIAL REFERENCE … · nat. hist. bu比.siams配.43: 303-335,1995 key to lichen genera in thailand with special reference to epiphytic taxa,

306 P.A. WOLSELEY AND B. AGUIRRE-HuDSON

Filamentous Foliose h削 ers~字翠戸

ーーーーーーー・ 100~m paraplechtend、ymafrom Poelt (1966)

hetromerous

prosoplechtenchyma

- 1∞ドm

Vegetative reproductive structures Soralia

バ!将ぱ;;mpM t

喰;ijjj保~:r:み;.~-.!;/r~笠23宗主・ヰニ一二~ご三二一一一-一 一一Position on thallus

Plate II. Thallus struct山巴。

Isidia

孔{ヤ

川U

ぷ一一v

fungal hyphae

lower coはex

cyphella (5tida sp)

金:ご二ーコ

て主主〉

Page 5: KEY TO LICHEN GENERA IN THAILAND WITH SPECIAL REFERENCE … · nat. hist. bu比.siams配.43: 303-335,1995 key to lichen genera in thailand with special reference to epiphytic taxa,

KEY TO L1CHEN GENERA TN TT-TATLAND 307

Basiodiomycetes Basiodiocarp 智 、話 。靭J!r;.;;.-b田伽pore

圃圃圃 10ドm

gyrose (Umbilicaria sp)

_..basidium

Ascomycetes

:-_;.....,.t: ):六下人文,

lecideine, lacking a thalline exciple (Porpidia sp) 圃園田園圃圃

Plate III. Sexual r巴production-themycobiont

scales = 1mm

lecanorine, with thalline exciple (Lecanora sp) 園田園圃・圃

主:ν '"-1.二口ゴ.て勺!

Structure of ascus

Page 6: KEY TO LICHEN GENERA IN THAILAND WITH SPECIAL REFERENCE … · nat. hist. bu比.siams配.43: 303-335,1995 key to lichen genera in thailand with special reference to epiphytic taxa,

308 P.A. WOLSELEY AND B. AGUIRRE-HuDSON

ルIyrmecia

(Chlorophyceae)

Trentepohlia

(Chlorophyceae)

Plate IV. Lichen photobionts

Trebouxia (Chlorophyceae)

Stigonema (Cyanobacteria)

Page 7: KEY TO LICHEN GENERA IN THAILAND WITH SPECIAL REFERENCE … · nat. hist. bu比.siams配.43: 303-335,1995 key to lichen genera in thailand with special reference to epiphytic taxa,

KEY TO LICHEN GENERA IN THAILAND 309

Thallus Structure (microscopic)

Thalli may be heteromerous and stratified (in section) with differing upper and lower

surfaces a distinct dorsiventral structure, or homoiomerous with similar upper and lower surfaces, photobiont and mycobiont occぽ ringmore or less randomly throughout the thallus.

Cortex: consists of tightly packed hyphae which may have a distinct chemis仕yand colour, occuηing in two characteristic tissues; paraplectenchyma-as hyphae growing from the innerp紅 tof the thallus at right angles to出esurface, giving血ecortex a cellular appe紅'ance

in both cross and longitudinal section, or prosoplechtenchyma-as cortical hyphae p紅 allelto the surface.

Photobiont layer: usually lies between the upper cortex and the medulla in foliose and

crustose species, and as a layer紅 oundthe medulla in fruticose species. The contact between the photobiont cells and the mycobiont cells may be developed by haustoria

(swollen tip of hyphae).

Medulla: a more or less compacted cottony layer of hyphae forming the main body of the

lichen. It also contains characteristic lichen compounds. It may be white or coloured.

Hypothallus (also known as prothallus by some au血ors):the initial structure of hyphae without algae from which a lichenised thallus develops is often visible along the margins

of the thallus. Found in crustose, squamulose and fruticose genera.

Thalline Features (use 10x hand lens)

The upper surface of the thallus may be smooth, wrinkled, cracked or reticulate; shiny, pruinose or hail)人

Cyphellae紅 es住uc刷redperforations through the lower cortex, with a raised rim around a cup-like s凶 ctureand a distinct membrane bounding the medulla.

Pseudocyphellae are shallow, round, long, or effigurate openings in the cortex exposing the medulla.百leycan occur on upper or lower surface or on the margins of lobes. When on the upper surface maculae and pseudocyphellae may not be easy to distinguish.

Irregularities of dis位ibutionof algal cells c叩 givea white spotted or reticulate pattern of

maculae, which may become fissured and cracked, when they訂'ehard to distinguish from pseudocyphellae.

Cephalodia紅 ebodies of cyanobacteria and fungal hyphae growing on the surface or within the thallus of a lichen with a green photobiont. Al白oughthe fungal p紅 tneris血e

same,白eappearance may be quite different.

Page 8: KEY TO LICHEN GENERA IN THAILAND WITH SPECIAL REFERENCE … · nat. hist. bu比.siams配.43: 303-335,1995 key to lichen genera in thailand with special reference to epiphytic taxa,

310 P.A. WOLSELEY AND B. AOUIRRE-HUDSON

Vegetative Reproductive Structures

Soralia訂'ef'Ormed by a rup加ring'Of血ec'Ortex t'O exp'Ose a p'Owdery 'Or granular m出 s'Of

s'Oredia. each c'Onsisting 'Of ph'Ot'Obi'Ont cells l'O'Osely enmeshed in fungal hyphae and with'Out

組 Yc'Ortical s回 C加re.百leymay be laminal, mar脚 al'Or arise fr'Om dactyls 'Or 抑制les,(irregul紅 c'Orticatedpro加ber;佃 ces,usually eroded), and may v紅y金omfine powder to a

granular appe釘加ce,and紅'eusually paler in c'Olour白anthe thallus. Isidia have a c'Ortex

組 d'Originate槌 extensionsof the upper surface of the thallus. They may be p泊ーlike,globose or flattened, branched, corall'Oid 'Or simple and may be situated加国nally'Or

marginally on血eupper surface 'Of the thallus, appearing血esame c'Olour or darker白血

血.ethallus. They may break 'Off企''Om 血ethallus leaving a pale紅白, and occasionally

become sorediose.

Phyllidia紅'esmall dorsiventral structures, cons凶ctedat base, appearing on血em紅g泊'Or

加国na'Of the lichen thallus. More complicated complanate cortical outgrowths 'Occuring

in the genus Stereocaulon are called phyllocladia.

LICHEN MYCOBIONTS

Sexual Reproductive Structures

Sexual repr'Oducti'On is 'Only found in白efungal symbi'Ont in lichens S'O白atreleased

sp'Ores will only reproduce出efungal partner. Al出oughthe fungal p紅側eris f'Ound 'Only

in the lichenised state,出esame algal partner can als'O 'Occ町 ina f同e・livingsぬ.te,姐din

'Other lichens in different genera. When a fungal spore gerr凶natesit h邸 toincorporate an

algal P紅 tnerin order to form the symbiotic organism, a lichen. Wi血in也efungal kingd'Om

lichenised genera are found in出easc'Omycetes and basidiomycetes. Ascomycetes紅ethe

m'Ost frequently 'Occurring group producing characteristic fruiting bodies, apo曲ecia組 d

perith目ia.The m'Ost c'Ommon f'Orm of出eapo曲目iumis a disk-like 紺 ucturefound on

the margins, 'Or usually upper surface of the thallus, it c'Ontains the reproductive佃 d

accessory tissues which provide important diagnostic features. Lecideine apo曲 目iahave

no血all泊em紅ginto the apo出ecium,出eouter edge of血eapo曲目iumf'Orrning a rim

known儲白eproper exciple, lecanorine apo血eciahave a thalline m釘 g泊 (c'On鈎泊担galg鵠)

visible around the apo也氏ia.The apothecia may be elongated forrning Iirellae. Peri曲目ia

are gl'Obular or pear-shaped recept配 lesenclosing the sp'Ore bearing hymenium, with the sp'Ores released through an ostiole.

τ'he ascopor回 arecontained泊踊ci,sac-like b'Odies within the hymenium and v紅y

in shape, size, septation and colour.τ'here are usually eight spores, but也ismay be reduced

to one 'Or greater由加 eight.In some lichens the asci degenerate at maturity leaving the spores in a p'Owdery mass termed a mazaedium. Apothecia may als'O be bom on stalked

or cup-like podetia. C'Onidia (asexual spores) may develop in a flask-like pycnidium

(mayappe紅 likesmall perithecia), which may be immersed, sessile, 'Or pedicillate on血e

白allus.百lese紅'eproduced by budding 'Off simple or branched hyphae wi出inthepyc凶dium.

Basidiocarps pr'Oduce b錨 idiospor,田 extemallyat血eapex of血ereproductive cell or

b舗 idium.

Page 9: KEY TO LICHEN GENERA IN THAILAND WITH SPECIAL REFERENCE … · nat. hist. bu比.siams配.43: 303-335,1995 key to lichen genera in thailand with special reference to epiphytic taxa,

KEY TO LICHEN GENERA IN THAILAND 311

LlCHEN PHOTOBIONTS

The photobionts that provide the carbohydrates on which the fungal partner depends belong to two types of algae: the Chlorophyta (green algae) and the Cyanobacteria (formerly called blue-green algae). Within the Chlorophyta the commonest photobionts紅'eunicellular Trebouxia and filamentous Trentepohlia, the latter occurring more frequently in血etropics. Within the cyanobacteria the commonest photobionts are Nostoc, Scytonema, and Stigonema. These two main groups are readily distinguished in the field by thallus colour when wet. Lichens contai凶ngTrebouxia tum grass-green when wet, Trentepohlia appe紅 S

orange when the合ylichen surface is scratched, and lichens with cyanobacteriaωrn or remain blue-grey when wet, when they may also become gelatinous. Lichens with cyanobacteria紅 eable to synthesise nitrates from atrnospheric nitrogen, but they are also more sensitive to drought and occur more frequently in wet or moist environments. Within the monsoon cIimate of Thailand they are more frequent in damp rnist-drenched montane communities over 1000 m.

This apparently simple symbiosis is more complex in some cases; a single lichen species may contain more than one photobiont, or may change photobionts and thallus form at different stages of its life or in different habitats (JAMES AND HENSEEN, 1976)

LlCHEN CHEMISTRY USED悶 THEKEY

Lichens contain many unique compounds which訂'ederived from也氏em司jormetabolic pathways, and have been used in chemotaxonomy to identify groups of related species. Several techniques of chromatography are used to extract these substances. However, the use of simple readily-obtained chemicals and colour reactions still remains a basis for field identification. As many of the genera in the lichen fa凶 lyParmeliaceae訂 edistinguished by their chemis仕y,these tests become important for identification of the genera level.

Three chernicals are commonly used: potassium hydroxide (K), bleach or calcium hypochlorite (C),組dparaphenylenediarnine (Pd). K is used in a 10-159もaqueoussolutions,

組 dPd is made up freshly in 95% alcohol. K is stable and lasts indefinitely, but C must be fresh and smelling of bleach or it wi1l not produce a colour reaction (test on a known C + lichen e.g. Parmotrema tinctorum). All血reeare corrosive and can ruin paper or cIo白血g.They should be applied with great c紅 eto small pieces of the lichen thallus that can be discarded after the test.

Compounds may be found in the cortex or the medulla, so that colour reactions are tested directly on the cortex, and on the medulla by cutting away a small訂 eaof the photobiont layer to expose the medulla. Colour reactions and the constituent compounds are indicated in the key. Some lichen compounds fluoresce when exposed to the long-wave UV from an ultraviolet lamp (360μm). Fluorescense may be observed in the whole lichen, or following separation of lichen compounds on a chromatographic plate. Only substances which紅 eUV + in the cortex or medulla of the whole specimen訂 eused in出ekey. Xanthones are found in the cortex and may fluoresce various shade of yellow, red and or;組 ge(e.g.some species of丹'xine),while depsides and depsidones found in the medulla fluoresce blue to white (e.g. Dirinaria).

Page 10: KEY TO LICHEN GENERA IN THAILAND WITH SPECIAL REFERENCE … · nat. hist. bu比.siams配.43: 303-335,1995 key to lichen genera in thailand with special reference to epiphytic taxa,

312 P.A. WOLSELEY AND B. AGUJRRE-HUDSON

KEY TO MACROLICHEN GENERA

The key is presented in dichotomous form, where possible using a combination of morphological and sexual characters. However some fami1ies血at訂 eimportant叩 dcharacteristic components of lichen communities in monsoon forests, such as Parmeliaceae and Physciaceae are frequently found in an unfertile state. In order to accomodate these specimens genera are keyed out several times in the key.

Terminology is described in the gloss釘y(see Appendix) and is illustrated in Plates I-IV.

Genera in bold type have been found in Thailand, others are found in adjac四 t紅 'eas組 d紅 'eexpected to occ町 in官lailand.

la. Thallus fruticose or filamentous, erect, pendulous, or prostrate (including podetia 叩 dpseudopodetia on a squamulose or crustaceous or filmy base),山uc加remore or less radial to isolateral, although a slight colour difference between the physiological upper and under side may occur…H ・H ・...・ H ・-…....・ H ・..… H ・H ・...・ H ・....2

lb. Thallus foliose, placodioid or squamulose, structure usually clearly

dorsiventral …...・ H ・...・ H ・...・ H ・.......・ H ・....・ H ・-……・…...・ H ・....・ H ・...・ H ・.....・ H ・...・ H ・...・ H ・H ・H ・...16

Filamentous Genera

2a. Thallus composed of fine filaments clearly visible with a 10x lens …....・H ・-…..32b. Thallus with a cortical s住uc飢re,not composed of fine filaments …...・H ・....・H ・...63a. (2a) Thallus blue-green, sessile, shelf-like, often forming rosettes, sometimes ribbon-

like. Texture fibrillose, soft when wet brittle when世y;basidiocarps white or creamish on underside of thallus ....・ H ・-…....・ H ・....・ H ・-…....・ H ・....・ H ・.....…Dictyonema

3b. Thallus greyish, green or yellowish-green, forr凶ngcushions or sheets or :t foliose-like; soft wet orむy;apothecia scattered on upp巴rsurfac…....・ H ・-…...・ H ・....・ H ・...4

4a. (2b) Thallus whitish-grey and crust-like, tex旬rewoolly; apothecia lecanorine,

grey to black ...・ H ・.....・ H ・-…....・ H ・...・ H ・.....・ H ・....・ H ・....・ H ・....・ H ・....・ H ・-…...・ H ・-…..Crocynia4b. Thallus green to black, branched and shrubby or felt-like.…...・ H ・-… H ・H ・...・ H ・-…..55a. (4b.)τ'hallus green to yellowish-green, felt-like, apothecia lecideine, or姐 ge.

…....・H ・..…Coenogonium5b. Thallus brown to black, repeatedly dichotomously branched to form small cushions

...・H ・Polychidium

Fruticose genera

6a. (2b).百lallUScitrine-yellow to deep orange (K+ purple) or rarely grey; apothecial

disc or粗野 (K+purple)…...・ H ・.....・ H ・....・ H ・....・ H ・...・ H ・....・ H ・...・ H ・....・H ・-…Teloschistes6b. Thallus and apothecial disc not citr加eyellow or or叩 ge....・H ・H ・....・ H ・-……....・...77a. (6b). Thallus (at least upright parts) with a cen回 l鉱 isor cylinder of carti1aginous

tissue.....・H ・-….....・ H ・...・ H ・.....・ H ・....・ H ・......・ H ・....・ H ・-……....・ H ・...・ H ・....・ H ・.......・ H ・....・ H ・...…87b. Thallus without central axis of carti1aginous tissue ..,・ H ・.....・ H ・....・ H ・....・ H ・....・ H ・...128a. (7a). Thallus of branching pseudopodetia covered with 1巴proseecorticate granules

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KEY TO LICHEN GENERA IN THAILAND 313

...・H ・..…Leprocaulon8b. Thallus with a well-developed cortical structure on squarnules or branches ….9

9a. (8b). Thallus ofupright podetia with a central, hollow, rigid cartilaginous cylinder; squarnules often present on podetia or at their base, fibrils absent; apothecia

without thalline exciple ....・ H ・....・ H ・....・ H ・.......・ H ・.........・ H ・...・ H ・....・ H ・.....・ H ・...・ H ・...・ H ・..109b. Thallus an erect or pendulous shrub with a hollow or solid central axis of elastic

or flexuous cartilaginous tissue; branches bearing few to nurnerous fibrils, squarnules absent; apothecia with thalline exciple and cilia.…...・H ・.....・H ・...Usnea

lOa. (9a) Thallus sirnple or spぽinglybranched, often squ氾nulose,branches tapering or terrninated by cups with convex to conglornerate brown or red apothecia

...・ H ・H ・H ・..…CladonialOb. 百lallusrepeatedly branched without squarnules, apices without cups or conspicuous

apothecia ....・ H ・....・ H ・-…...・ H ・....・ H ・....・ H ・.....・ H ・....・ H ・....・ H ・....・ H ・-…・・ H ・H ・...・ H ・-…....・ H ・...11lla. (10b) Thallus conspicuously and consistently perforate, surface srnooth and shiny,

paler above and brown in older parts .............・ H ・-… H ・H ・-…...・ H ・.Cladia aggregata 11 b. Thallus irnperforate except at the axils of branches, surface uneven, rnatt or

arachnoid ....・ H ・...・ H ・.....・ H ・-…...・ H ・.....・ H ・.....…....・ H ・...・ H ・-…・…CladoniaSect Cladina 12a. (7b) Thallus dark brown, or dark grey, pendulous with filarnentous branches

…Bryoria 12b. Thallus white to pu中lish,or green to grey, erect or if pendulous with flattened

strap-like branches ..........・ H ・-…....・ H ・-… H ・H ・....・ H ・-… H ・H ・..…・.....・ H ・...・ H ・-…… H ・H ・-…...1313a. (12b) Thal1us pale grey to lilac-grey; cortex C + bright red; photobiont

(Trentepohlia); rnainly coastal …...・ H ・....・ H ・-… H ・H ・....・ H ・-…...・ H ・.........・ H ・-…..RocceUa13b. Thallus pale green-grey, cortex C-; photobiont green ...・ H ・H ・H ・..…...・ H ・.....・ H ・..…1414a. (13b). Thallus silvery or ash grey; branches bearing phyllocladia and cephalodia;

saxicolous or terricolous in rnontane and alpine regions ....・H ・.....・H ・.Stereocaulon 14b. Thallus shades of green, grey, or straw, lacking phyllocladia, phyllocladioid

branchlets,組dcephalodia; substrate and distribution various.…...・H ・-….....・H ・..1515a. (14b). Thallus gree凶shto brownish grey; apo出eciacornrnon, disc black powdery;

on rnossy e紅白 orrnossy tree trunks in rnontane forest ........・H ・...Sphaerophorus 15b. Thallus various, if filarnentous, pendulous on tree trunks and branches; usnic acid

present, rnazaediurn absent, distribution and substrate various (widespread)

....................・・・・・・…........Ramalina

Foliose and Squamulose Genera

16a. (1b). Thallus hornoiornerous, colour slate grey, brown, brown-black, or green-

black ...・ H ・.....・ H ・..…...・ H ・....・ H ・.....・ H ・...・ H ・..…...・ H ・.....・ H ・H ・H ・..…...・ H ・.....・ H ・..…...・ H ・..…・1716b. 百lallusheterornerous, colour green, grey or yellow-orange, various …....・H ・..1917a. (16a). Thallus tough, thick and swollen, cartilaginous and ridged even when wet,

spores s面lple…....・ H ・....・ H ・.....・ H ・....・ H ・....・ H ・....・ H ・...・ H ・.....・ H ・...・ H ・....・ H ・..…...・...Physma17b. 百lallussoft, pliant when wet; rnore or less crisp, fragile when dry, spores rnurifoロn

or septate…...・ H ・.....・ H ・.....・ H ・H ・H ・.....・ H ・.....・ H ・....・ H ・-… H ・H ・.....・ H ・H ・H ・.....・ H ・...… H ・H ・...1818a. (17b). Thallus brown-black or green-black, often swollen when wet, without a

cellular upper cortex ...・ H ・-…....・ H ・....・ H ・........・ H ・.......・ H ・-…....・ H ・...・ H ・.....・ H ・..Collema

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314 P.A. WOLSELEY AND B. AGUIRRE-HUOSON

18b.τ'hallus blue-grey,brownish or greyish-black, not usually swollen when wet, cortex present as a single layer of cells ....・H ・-…....・H ・...・H ・.............・H ・...・H ・.......Leptog畑m

19a. (16b).τ'hallus of small squ組lUlesor lobes > 5 mm long and wide .......・H ・...2019b. Thallus of larger foliose or ribbon-like lobes ....・H ・...・H ・-…....・H ・....・H ・....・H ・....・H ・.2820a. (19a). 官lallUSci凶ne-or or,加ge-colo町 ed....・H ・...……・・H ・H ・-…….......Candelaria 20b. 百lallusof other colour …H ・H ・...・H ・-…・H ・H ・-…...・H ・-…....・H ・....・H ・...・H ・.....・H ・…...・H ・.2121a. (20b)官lalluswith blue-green photobiont (dark grey when wet) ...・H ・-…...・H ・..2221b. Thallus with green photobiont (grass green when wet) ……....・H ・...・H ・-…....・H ・.2322a. (21a) Thallus without chemistry, apo由民iausually without thalline exciple

...・H ・.....・H ・....・H ・...ParmelieUa 22b. Thallus Pd+orange (pannarin), apothecia with thalline exciple ...・H ・-….Pannaria23a. (21b) Hypothallus present ...・H ・.....・H ・...・H ・....・H ・....・H ・.....・...・H ・...・H ・.....・H ・-…...・H ・-….2423b. Hypothallus absent ………....・H ・...・H ・-…・H ・H ・-…・H ・H ・-…....・H ・....・H ・...・H ・-…...・H ・-…2524a. (23a)百lallussquamulose (occasionally crustose), apo由民iabiatorine ..・H ・H

... PhyUopsora 24b. Thallus foliose, with over1apping lobes; apo曲目iawith thalline m紅 g泊…・H ・H ・..

・・…...・H ・.....・H ・.....Physcidia wrightii 25a. (23b) Thallus of pale red adjacent squamu1es with a white rim (montane)…・・

...・H ・.......・H ・.Psora25b. Thallus of blue-grey to green squ阻lUles..・H ・...・H ・-…・H ・H ・....・H ・.....・H ・...・H ・.....・H ・..2626a. (25b)官lallusof irregul紅 paperythin squamules with cortex 1 cell由ick....・H ・..

…・H ・H ・-…....・H ・...・H ・-……・・H ・H ・..Escha旬gonia26b. Thallus of regu1ar orbicular or leaf-like squamules with cortex more由加 1cell

出ick・…・H ・H ・....・H ・-…............・H ・...・H ・....・H ・....・H ・.....・H ・...・H ・-……・H ・H ・-…・............・H ・...2727a. (26b) Thallus of bl凶sh-grey or gl叩 .cous-greyorbicular squamules with neatly

raised rim, often sorediate, upper and lower surfaces concolorous, lower surface

pale or white ...........・H ・....・H ・-…・H ・H ・....・H ・.....・H ・...・H ・-・・H ・H ・...・H ・....・H ・....Normandina27b. Thallus of over1apping or erect leaf-like squ創nules…...・H ・.....・H ・....・H ・-・Cladonia28a. (19b)百lallUSattached to subs回.teon1y by a cen凶 1holdfast …...・H ・.....・H ・-….2928b. Thallus attached to substrate by rhizines, scattered hapters, the under side, or

arising 企oma basal holdfast ...・H ・....・H ・....・H ・-……...・H ・-…・H ・H ・.....・H ・...・H ・....・H ・....3029a. (ο28aω). 官1凶allusw制i血vesiculate swelling 0叩nits包u叩ppeぽ:rsurface and with c∞OI町τe白:spo叩n叫岨.di曲din暗1喝g

indentations below …................…...・H ・-…H ・H ・...・H ・...・H ・...・H ・.........・H ・...・H ・-…Lasallia29b. Thallus without vesicular swellings ...・H ・-…...・H ・....・H ・-…...........・H ・-….Umbi1icaria 30a. (28b) Cyphellae or pseudocyphellae on under side of thallus ...・H ・.....・H ・...・H ・...3130b. Cyphellae and pseudocyphellae absent仕omunder side of thallus...・H ・.....・H ・...3431a. (30a) Under side with cyphellae (compounds absent) ...・H ・.....・H ・...・H ・.....・H ・.Sticta31b. Under side with pseudocyphellae (compounds present)………....・H ・.....・H ・...・H ・.3232a. (31b) Lower cortex tomentose, thallus regularly branched lobes …

…PseudocypheUaria

32b. Lower cortex naked …....・H ・....・H ・...・H ・....・H ・...・H ・-…・....・H ・.....・H ・-…・H ・H ・.........・H ・..….3333a. (32b) Lower surface brown, margins with many stalked verruciform pycnidia,

apothecia if P問 sent,on lower surface... .....……...........・...Nephromopsisorna旬33b. Lower surface pale to white, m紅 ginswithout cilia or conidia, apothecia lar凶nal

... Cetrariopsis wallichiana

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KEY TO LICHEN GENERA IN THAILAND 315

34a. (30b). Lower cortex absent.…...・ H ・.....・ H ・....・ H ・....・ H ・......・ H ・.....・ H ・.....・ H ・...・ H ・....・ H ・...3534b. Lower cortex present (1ook carefully at sma11 firrnly appressed species ....・H ・..4235a. (34a). Lower surface usually distinctly veined, often with projecting rhizines;

apothecia partly immersed at apices of lobes ........・ H ・-…・ H ・H ・.....・ H ・....・ H ・.Peltigera35b. Lower cortex not veined, without projecting rhizines (marginal cilia may be

present); apothecia sessile to stalked, laminally or submarginally situated.…..36 36a. (35b). Photobiont a chlorophyte (thallus green when wet)…...・ H ・.....・ H ・....・ H ・..3736b. Photobiont a cyanobacterium (thallus dark grey when wet)….....・ H ・-… H ・H ・-…..4037a. (36a) Tha11us with margina1 ci1ia; spores brown, 2-celled ....・H ・-…Heterodermia37b. Thallus without marginal cilia; spores colour1ess, simple ........・ H ・-… H ・H ・...・ H ・...3838b. (37b) Hypothallus absent, apothecia m紅gina1, stalked ....・H ・....・H ・..Gymnoderma 38a. Hypotha11us present; apothecia larnina1, :f: sessi1e.…・・ H ・H ・-…...・ H ・...・ H ・...・ H ・...・ H ・...3939a. (38b). Tha11us crustose, squamulose, or foliose, apothecia biatorine ....・H ・

..... ........ Phyllopsora 39b. Thallus foliose, apothecia with a thalline margin ...・H ・....・H ・-…Physcidiawrightii 40a. (36b). Upper side tomentose; apothecia submargina1, stipitate; hypotha11us absent

(rhizines may be pigmented bluish grey) ....・ H ・....・ H ・....・ H ・...・ H ・..…....・ H ・.Erioderma 40b. Upper side glabrous or weakly pub巴scent;apo曲目ialaminal, sessi1e to substipitate;

hypotha11us bluish black ....・H ・...・H ・........................................................................4141a. (40b) Tha11us without chemistry, apothecia usua11y without tha11ine exciple …・・

.. ....... Parmeliella 41b. Tha11us Pd+orange (pannarin), apothecia with thalline exciple ....・H ・-….Pannaria42a. (34b). Photobiont a cyanobacterium (tha11us dark grey when wet) ...・H ・.....・H ・..4342b. Photobiont a chlorophyte (thallus bright green when w巴t)....・ H ・....・ H ・-…....・ H ・-…4543a. (42a). Under side with a co紅 S巴 darkreticulate pattem of short brown to black

hairs, with bare pa1e areas in reticulations ....・ H ・.......・ H ・-….....・ H ・....・ H ・-……Lobaria43b. Under side naked or uniformly finely pubescent, lacking a dark reticulate pattem

of hairs …...・ H ・.....・ H ・-…....・ H ・.....・ H ・-…....・ H ・...・ H ・.......・ H ・...・ H ・...…....・ H ・....・ H ・...・ H ・...・ H ・.4444a. (43b). Tha11us medium to dark brown with short brown to grey tomenωm ofshort

hairs on under side; apothecia on the under side of lobe margins .

…. Nephroma 44b. Tha11us slate-blue, bluish grey, grey, or yellow, with numerous rninute para11el

ridges from centre to margins, often with blue-black hypothallus on underside; apothecia adnate to sessile on upper side of thallus........・H ・........・H ・.Coccocarpia

45a. (42b). Underside with a tomentum of short hairs which may be evenly distributed or form a reticulate pattem with bare areas in reticulations; intema1 cepha10dia

present ...・ H ・.....・ H ・..…...・ H ・....・ H ・...… H ・H ・.....・ H ・...・ H ・H ・H ・.....・ H ・H ・H ・..…...・ H ・....・...Lobaria45b. Under side naked or with rhizines, without short tomentum of hairs ....・H ・-…..4646a. (45b) Rhizines absent …H ・H ・...……...・ H ・..…...・ H ・..……………...・ H ・..… H ・H ・...・ H ・.....・ H ・...4746b. Rhizines present, at Ieast on the margins or in the centraI p紅白 ofthe thallus ..

……50 47a. (46a) Lobes hollow, :f: inflated (mainly montane forest species) ....・H ・........・H ・.4847b. Lobes soIid, thaIIus directly fastened to substrate or by scattered hapters (mainly

lowland and savanna species)..・H ・...・ H ・H ・H ・...........................................................49[Anzia not yet recorded from ThaiIand has soIidly inflated Iobes with a spongy

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316 P.A. WOLSELEY AND B. AGUJRRE..HUDSON

cushion of anastomosing hyphae on the lower surface] 48a. (47a) Hollow lobes with round perforations in upper cortex.

........・.Menegazzia terebrata 48b. Hollow lobes without round perforations in the upper cortex ..・H ・..Hypogymnia49a. (47b)百la11us白rkgrey to dark brown; atranorin absent, cortex K-, medulla UV

…Hyperphyscia 49b. Thallus pa1e grey or yellow; a回 norinusua11y present, cortex K+, medulla UV+.

...Dirinaria 50a. (46b) Upper co託exwith pseudocyphellae or maculae (photobiont layer unevenly

dis住ibuted,or surface cracked exposing medulla) ........…............・H ・....・H ・-……5150b. Upper cortex lacking pseudocyphellae or maculae (evenly dis住ibutedphotobiont

layer), surface not cracked …・H ・H ・....・H ・.....・H ・...・H ・-…・H ・H ・.....・H ・...・H ・-……...・H ・-…5751a. (50a) Tha11us lobes usua11y less白血 2mmwide;鎚 cosporesseptate..・H ・-町対即

日b. Tha11us lobes usua11y more由an2mm wide; ascospores s加ple…............・H ・..5252a. (51b) Upper surface with line紅 toreticulate pseudocyphellae or maculae, lower

surface black …・H ・H ・-…・H ・H ・....・H ・.....・H ・'"・H ・-…....・H ・...・H ・.....・H ・-…..,・H ・-…・H ・H ・-…..5352b. Upper surface with orbicular pseudocyphell蹴;lower surface pa1e tan to black

……55 53a. (52a)官la11usgreen to yellow-green, lobes linear, narrow, and仕組sverselycracked

..,・H ・..Relicinopsis 53b. 百la11usgreen to grey or buff, lobes rounded and finely reticulately cracked or

white-maculate ....・H ・...・H ・....・H ・-…....・H ・-…....・H ・...・H ・....・H ・....・H ・.....・H ・...・H ・.........・H ・-・5454a. (53b) Cilia simple, rhizines simple or squarrose, fine reticulate pa仕.emof maculae

sometimes developing泊tocracks, conidia sublageniform........・H ・...・H ・-…Rimelia54b. At least some cilia sp釘 selybranched, rhizines of two旬pes,maculae effigurate,

conidia filiform..・H ・.....・H ・...・H ・....・H ・.....・H ・...・H ・-…...・H ・.........………....・H ・.Rimeliella 55a. (52b)百la11usyellowish green; rhizines企'equent........………........……Punctelia 55b.τ'ha11us grey-green to brown; rhizines sp紅 seor absent ...・H ・.....・H ・....・H ・-…・H ・H ・.5656a. (55b)百la11usridged-rugose, isidiate, caperatic acid present... Platismatia erosa 56b. Tha11us smooth, punctate on lower surface, with or without isidia and soredia,

caperatic acid absent ....... ..... ..... ...... ..... .......... ........... ......... ......................... Cetrelia 57a. (50b) Margins of tha11us ciliate, sometimes only in lobe axils ....・H ・............・H ・.5857b. Margins of tha11us without cilia (in some species rhizines紅'econtinued to出e

margins)…...・H ・-………・H ・H ・-………....................・H ・........・H ・-…・H ・H ・....・H ・....・H ・-…7058a. (59a) Cilia bulbate at base...・H ・....・H ・....・H ・-…...・H ・-………......・.....................・H ・...5958b. Cilia not bulbate at base ..・H ・...…..........................................................................6059a. (58a) Upper s町 facegrey, cortex K+ yellow (a仕組orin)...・H ・-….........B叫bo曲rix59b. Upper surface straw-yellow to yellow green (us凶cacid), cortex K-…

…Relicina 60a. (58b) Lower surface corticate....・H ・............・H ・H ・....・H ・.........………・.Heteroden凶a60b. Lower surface ecortica旬…・H ・H ・....・H ・....・H ・....・H ・............・H ・.....・H ・..,・H ・....・H ・...........6161a. (62a) Upper surface brown, gree凶shwhen wet …...・H ・-……...・H ・..Phaeophyscia 61b. Upper surface grey to green to 紺 awyellow ………...・H ・....................・H ・.....・H ・.6262a. (63b) Lobes s回 p-likeand cana1iculate, ascending and shrub-like, dichotomously

branching …...・H ・-…………....・H ・....・H ・....・H ・....・H ・....・H ・........................Everniastrum

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KEY TO LICHEN GENERA IN THAILAND 317

62b. Lobes of thallus not strap-like or canalicu1ate ..・ H ・-… H ・H ・.....・ H ・...・ H ・-……...・ H ・...6363a. (62b) Lower cortex white to pale brown....…....・ H ・...........・ H ・.....・ H ・...・ H ・...…・ H ・H ・.6463b. Lower co此exdark brown to black ........・ H ・.....・ H ・....・H ・...・ H ・...・ H ・-…....・ H ・....・H ・.......6564a. (63a) Thallus matt, usually pruinose at least at lobe tips, upper cortex

P紅 aplechtenchymatous....・ H ・...・ H ・-…....・ H ・....・ H ・....・ H ・....・ H ・...・H ・-…....・ H ・....・…Physcia64b. 百lallussmooth to shiny, not pruinose, upper cortex prosoplechtenchymatous…

…Heterodermia 65a. (63b) Medulla yellow to orange ....・ H ・...・ H ・....・ H ・...・ H ・....・H ・.....・H ・.....・H ・.Myelochroa 65b. Medulla white ........・ H ・....・ H ・....・ H ・...・ H ・....・ H ・....・ H ・....・ H ・....・ H ・-…...・ H ・...・ H ・...・ H ・.....・ H ・..6666a. (65b) Conspicuous conical verruciform pycnidia present along margins of thallus

... Parmelaria 66b. Conical verruciform pycnidia absent企omm紅 ginsof thallus ...・H ・-……・ H ・H ・...6767a. (66b) Lobe tips broadly rotund with a broad naked zone on lower surface margin

・H ・H ・.....・ H ・...Parmotrema 67b. Lobe tips narrow,住uncateto sub-rotund, with rhizines to margins....・H ・...・H ・..6868a. (67b) Cilia evenly spread on lobe margins; lobes with住uncateapices...・H ・........

............ Parmelinopsis 68b. Cilia mainly in lobe axils; lobes with sub-rotund apices …....・ H ・...・ H ・.........・ H ・..6969a. (68b) Lobes broad (5-10mm); medulla K+ yellow turning red (salazinic acid) .

……H ・H ・...・ H ・.Parmelinella 69b. Lobes narrow (l-6mm); medulla K・ H ・H ・-…....・ H ・-…H ・H ・-…H ・H ・-….....・ H ・..P紅melina70a. (57b)τ'hallus straw-yellow to yellow-green ....・ H ・...・ H ・-…...・ H ・...・ H ・...・ H ・.....・ H ・.....7170b. Thallus grey to green…...・H ・....・ H ・-…・ H ・H ・-…...・ H ・-…....・ H ・...・ H ・.....・ H ・-…........・ H ・...7271a. (70a) Lower surface rhizinate to margins, occu町ingon rocks and soil, spores

5-12μm long…....・ H ・-…...・ H ・-…...・ H ・....・ H ・....・ H ・-…........・ H ・...・ H ・....Xanthoparmelia 71b. Lower surface naked at margins, occurring on bark and rocks, spores 12-18仰n

long ....・ H ・-…...・ H ・-….....・ H ・...・H ・....・ H ・-…....・ H ・........・ H ・-…....・ H ・.......・ H ・-….Flavoparmelia 72a. (70b) Rhizines dichotomously branched and prominent, often pro佐udingbeyond

the lobe margin ...・ H ・.....・ H ・-…....・ H ・...・ H ・....・ H ・....・ H ・.........・ H ・...・ H ・.........Hypotrachyna 72b. Rhizines simple, squarrose or reduced to hapters, not pro住udingbeyond lobe

margin ..........・ H ・.....・ H ・.........・ H ・-… H ・H ・....・ H ・....・ H ・........・ H ・.........…...・H ・-……....・ H ・....7373a. (72b)百lalluslobes usually more than 5mm wide, broadly rotund at tips, erect or

patent but not appressed to substrate; rhizines sp紅 seor absent in a broad naked zone ne紅 lobetips ...・ H ・.........・ H ・.....・ H ・...・ H ・....・H ・.....・ H ・-…....・ H ・...・ H ・-…Parmotrema

73b. Thallus lobes usually less白血 3mmwide,住uncateor line民 appressedto subs回 te;rhizines extending to lobe tips or reduced to hapters ....・ H ・-…....・ H ・....・ H ・...・ H ・-…74

74a. (73b) Thallus attached by sparse to abundant simple rhizines.... ….. Canoparmelia

74b. 百lallusattached to subs回 teby branched rhizines or by hapters ...・H ・.....・H ・-…7575a. Upper surface with pseudocyphellae; apo也氏ialecideine ...・H ・....・ H ・.....・ H ・.Pyxine75b. Upper surface without pseudocyphellae; apothecia lecanorine ....・H ・-…..Dirinaria

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318 P.A. WOLSELEY AND B. AGUIRRE-HuoSON

LICHEN GENERA: KEY CHARACTERS AND DISTRIBUTION IN 1百AlLAND(Families according to Systema Ascomycetum ( E阻 KSSON& HA WKSWOR叩, 1991)

BULBOTHRIX Ha1e (PARMELIACEAE Zenker)

Key characters: tha11us foliose, grey, lobes土adnate,cilia bu1bate at b邸 e,rhizines sirnple to dichotomous. Distinguished from Relicina with bulbate cilia, by grey colour = absence of usDIc acid.

Distribution: epiphytic and widespread in deciduous and evergr巴enforests, c. 7 species recorded in Thail組 d.Characteristic species of deciduous dipteroc紅pforests include B.pigmenωcea (Ha1e) Ha1e (K-) andB. isidiza (Nyl.) Ha1e (K+ reの,白elatter being prese凶but less frequent in evergreen forests.

CANDELARIA Massal (CANDELARIACEAE Hakul.)

Key characters: tha11us squamulose, bright yellow; photobiont Chlorococcus; apo血.ecialecanorine, asci 20-50 spored. Distinguished from other yellow lichens by sma11 crust-like squ創nules.

Distribution: C. concolor (Dickson) B. Ste泊, a new record inτbailand, epiphytic on branches in deciduous dipterocarp forest.

CANOPAR島IELIAElix & Ha1e (PARMELIACEAE Zenker)

Key characters: thallus foliose, lobes adnate or overlapping, yellowish to grey gr閃 n,ciliaabsent (occasionally in lobe axils), rhizines simple; apothecia laminal, eperforate. Distinguished企omclosely related genera by simple rhizines and absence of cilia.

Distribution: epiphytic姐 dfrequent泊 well-litsi加ations,c. 6 species found in百lailand,C. ecaperaω(Mull. Arg.) Elix & Ha1e (thallus yellowish, lower cortex black) being仕'equentin血edipterocarp forest, and C. salacinifera (Ha1e) Elix & Ha1e (出a11usgrey, lower cortex pa1e tan) being associated with montane forests.

CETRARIOPSIS Kurok. (P ARMELむ主CEAEZenker)

Key characters: tha11us foliose, er即 tand lettuce-like, yellow-green above, pale below with pseudocyphellae on the lower surface, rhizines infrequent, only present on白eanchored p紅 t.Distinguished from other cetrarioid genera by pa1e lower surface with white punctate pseudocyphellae.

Distribution: a monotypic genus, C. wallichiana (Taylor) Kurok. is found on twigs in montane forests above 1500 m, known in Thailand from Doi Suthep.

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KEY TO LICHEN GENERA IN THAILAND 319

CETRELIA W. Culb.& C. Culb. (PARMELIACEAE Zenker)

Key characters: thallus foliose, green to grey-brown with punctiform pseudocyphellae on upper surface, rhizines simple usually sp訂 se;apothecia perforate. Distinguished from other ce位制oidgenera by black lower surface and pseudocyphellae on upper surface.

Distribution: 2 species, epiphytic on twigs佃 dcanopy branches, re町 ictedto montane forests above 1500 m in the north of Thailand.

CLADIA Filson (CLADIACEAE Filson)

Key characters: thallus fruticose, yellow above to pale brown below. of hollow, perforate podetia. with an extemal cartilaginous layer, fertile podetia larger; apothecia terminal. lecideine. Distinguished by yellow colour and hollow podetia.

Distribution: te凶colousin mont組 evegetation above 1500 m. C. aggregata (Swartz) Nyl. is recorded in Thailand (AHTI, 1984).

CLADONIA Browne (CLADONIACEAE Zenker)

Key characters: prim訂yhorizontal thallus. green to grey green. with vertical podetia arising from it. primary thallus crustose, soon disappe紅 ing(subgenus Cladina) or of basal squamules. corticate above. ecorticate below (subgenus Cladonia); podetia. simple to richly branched. hollow. with or without squamules. ecorticate (Cladina), continuous to discontinuous, apothecia without thalline exciple.

Distribution: Epiphytic and terricolous in montane communities over 1000 m. Species recorded in Thailand include Cladonia siamea des Abbayes, C. transindica Ahti. C. rangiferina (L.) Nyl. (subgenus Cladina) recorded in adjacent coun位iesof India and China in montane forests.

COCCOCARPIA Pers. (COCCOCARPIACEAE (Mont. ex K.Muller) Henssen)

Key characters: thallus foliose :f: adnate. thallus grey blue tUIτ1Ing dark grey on wetting. upper surface frequently with concentric ridges on lobe tips. rhizines squarrose frequently white-tipped; photobiont a cyanobacterium-Scytonema; apo白ecialecideine. red to dark brown. Distinguished from other genera with cyanobacteria by squarrose rhizines.

Distribution: c. 4 species in Thailand. epiphytic and characteristic of evergreen forests at lower and higher altitudes. but persisting into more recentIy established dipterocarp or pine forests. absent from long established deciduous dipteroc紅pforest.

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320 P.A. WOLSELEY AND B. AGUIRRE-HUDSON

COENOGONIUM Ehrbg. (GYALECTACEAE Stizenb.)

Key characters: thallus filamentous, green to brown-black, threads branched mat-like; photobiont Trentepohlia; apo曲目ialecideine orange, bom on the tips of白efilaments.

Distribution: genus app訂 entlyrestricted to加 nksof moist-barked trees in undisturbed evergreen forests at low altitudes.

COLLEMA Wigg. (COLLEMATACEAE Zenker)

Key characters: thallus foliose attached to substrate by hapters, olive-green to brown-black, homoiomerous frequently wrinkled or pustulate when dry, gelatinous when wet; photobiont a cyanobacterium-Nostoc; apothecia lecanorine, sessile or stipitate, spores muriform (see Physma). Distinguished from Leptogium by absence of upper and lower cortex and from Physma by the absence of a densely short司 hairylower surface.

Distribution: mainly epiphytic and found in evergreen forests at lower and higher altitudes, but persisting into recently altered deciduous dipterocarp and pine dipterocarp forests in Thailand.

CROCYNIA Nyl. (CROCYNIACEAE M. Choisy ex Hafellner)

Key characters: thallus squamulose-placodioid, thick, white, with an abundant dark hypothallus, ecorticate giving a woolly appe紅 ance,black pigmentation present in白eprothallus and apothecia; apothecia lecideine, grey-black.

Distribution: C. pyx仇oidesNyl. is frequently associated with Phyllopsora species on佐unksand branches in the lowland and montane evergreen forests.

DlCTYONEMA Agardh (MERULIACEAE P.Karsten)

Key characters: thallus matt.司liketo foliose, upper side blue-to grey-green, fibrillose; photobiont cyanobacterial filaments surrounded by hyphae.

Distribution: a characteristic species of true rain forest, D. sericeum (Swartz) Berk. recorded in South Thailand on Kho Tao by PAULSON (1930), not recently recorded.

DIRINARIA (Tuck.) Clements (PHYSCIACEAE Zahlbr.)

Key characters: thallus foliose, yellow帽 greyto white, of closely appressed radiating lobes attached by hapters, rhizines absent; apothecia lecanorine, spores brown, thick walled, 1・septate. Distinguished from Pyxine and related genera by absence of rhizines and lecanorine apothecia.

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KEY TO LICHEN GENERA副官IAILAND 321

Distribution: a c'Osm'Op'Olitan genus 'Occurring m'Ost frequent1y in dry, well-lit and 'Openl disturbed sites. T'Olerant 'Of disturbance and p'Olluti'On, s'O出atit is企equent1yf'Ound泊町bansites. C'Omm'On species are D. consimilis (S伽'On)Awas.肌N-)and D. picω(Sw紅白)

Clements & Shear (UV +).

ERIODERMA P,白 (PANNARIACEAETuck.)

Key characters: thallus f'Oli'Ose, grey t'O grey-br'Own, upper surface with t'Omentum 'Of hairs. l'Ower surface ec'Orticate with dense tufts 'Of blue-black rhizines, ph'Ot'Obi'Ont cyan'Obacterial-Scytonema; ap'O血ecialecideine. Distinguished合om'Other genera副司凶landwith cyan'Obacteria by naked white l'Ower surface wi出 tufts'Of blue-black rhizines.

Distribution: E. sorediatum D.Gall'Ouay & P.M.J'Org. is f'Ound l'Ocally 'On twigs and branches in undisturbed m'Ontane f'Orests 'Over 1500 m.

ESCHATAGONIA Trevisan (f:創凶lyuncertain)

Key characters: thallus 'Of血inpapery el'Ongated and imbricate squamules usually > 5 mm l'Ong, bright green when wet, rhizines lacking. Resembles algal fr'Onds, but c'Ontains myc'Obi'Ont.

Distribution: 'On m'Ossy住unks泊 m'Oistundisturbed evergreen f'Orests at l'Ow altitudes.

EVERNIASTRUM Hale ex Sipman (PARMELIACEAE Zenker)

Key characters: thallus f'Oli'Ose, grey, l'Obes narr'Ow, linear, dich'Ot'Om'Ously divided,

canaliculate with dark l'Ower surface and m紅ginalcilia; ap'Othecia lecan'Orine with h'Oll'Ow sta1k. Distinguished仕omHeterodermia leucomelos (L.) P'Oelt by presence and dark c'Ol'Our 'Of l'Ower c'Ortex.

Distribution: a characteristic and abundant genus 'Of twigs in m'Ontane evergreen f'Orests 'Over 1000 m,血em'Ost c'Omm'On sp民 iesbeing E. nepalense (Tayl.) Hale ex Sipman. E. vexans,組 isidiatespecies, appe紅 sab'Ove 1600 m in undisturbed f'Orests.

FLAVOPAR勘IELIAHale (P ARMELIACEAE Zenker)

Key ch町 'acters:f'Oli'Ose, c'Ortic'Ol'Ous 'Or saxic'Ol'Ous, with br'Oad yell'Ow-green subr'Otund l'Obes, l'Ower c'Ortex with erhizinate z'One at margin; ap'Othecia eperf'Orate, sp'Or,回 large

(14ー20μm). Distinguished fr'Om Xanthoparmelia by 1釘'gersp'Ores.

Distribution: F. caperata (L.) Hale is rec'Orded frequently in出e阻malayas,n'Ot rec'Orded 泊官lailand.

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322 P.A. WOLSELEY AND B. AGUIRRE-HUDSON

GY勘INODERMANyl. (CLADONIACEAE Zenker)

Key characters:血allusfoliose, lobate, upper cor飽xyellow-brown, lower surface ecorticate, white, without rhizines, photobiont ch1oroccocoid, apo血eciam紅ginal,stalked, simple or aggregated, light brown.

Distribution: G.coccoca中umNyl. recorded in百凶land(YOSHIMURA & SHARP,1968) not recently.

HETERODERMIA Trevisan (pHYSCIACEAE Zah1br.)

Key characters: thallus foliose c10sely adpressed or ascending, grey to green-grey, lobes usually ~i1iate, upper suface with periclinal hyphae, lower surface with or without a cortex; photobiont chlorococcoid; apo由民ialecanorine,邸cosporesbrown,血ickwalled,l-septate.

Distribu舗on:l∞allydominant on twigs and仕出血 in出.emon:組問 forests,but also occuring on twigs in the lowland evergreen and dipteroc釘pforests.τ'his genus contains manY species in百 ailand.

HYPOGYMNIA (Nyl.) Nyl. (PARMELIACEAE Zenker)

Key characters: thallus foliose, pale gr芭:y,attached by adhesive discs below, lobes inflated, hollow, rhizines absent; photobiont chlorococcoid. Distinguished from Menegazzia by absence of rounded perforations on upper surface of hollow thallus.

Distribution: c.3 species res凶ctedto high altitudes on twigs and branches of佐ees泊

forests above 1500 m,白emost common sp民 iesbeing H. pseudobitteriana (Awas.) Awas.

HYPOTRACHYNA (Vainio) Hale (PARMELIACEAE Zenker)

Key characters: thallus foliose, pale glaucous grey, with凶 ncatelobe ends, lobes without m紅 ginalci1ia, but with dense dichotomously branched rhizines; photobiont trebouxioid. Distinguished from other Parmeliaceous genera by dense dichotomous rhizines often continued to m紅 gins.

Distribution: a characteristic and dominant genus in血eCanOPY of mon旬neforests above 900 m,江 isr紅 'elyfound in lowland forests. Common species include H. 10mω'sana (Zal止)Hale and H. orientalis (Hale) Hale.

LASALLIA Merat何MBILICARIACEAEChev.)

Key characters: thallus foliose, brown, umbi1icate, corticated above and below, with 抑制la旬 swellingson血.eupper surface and corresponding indentations below, rhizines

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KEY TO LICHEN GENERA町 τ百AILAND 323

absent, apo也氏ialecideine.

Distribution: saxicolous, occurs in Himalayan regions, Yunnan, and Taiwan; not found泊

百組landyet.

LEPROCAULON Nyl. (f:創凶lyuncertain)

Key characters: thallus仕uticose,pale gr,田nwhite, much branched, surface and apices leprose granul低

Distribution: 1 sp回 iesin Thailand, L. arbusculum (Nyl.) Nyl., associated with mossy trunks in montane rain forests over 1500 m.

LEPTOGIUM (Ach.) Gray (COLLEMATACEAE Zenker)

Key characters: thallus foliose, slate grey to brown, homoiomerous, lower cortex with or without hairs; photobiont a cy組 obacterium-Nostoc;apothecia lecanorine (sometimes not app紅 entto the nak:ed eye). Section Mallotium (Ach.) Vainio with has hairy thalli. Distinguished from Collema by pr,巴senceof upper and lower cortex l-cell也ick.

Distribution: widespread and more frequent in evergreen forests at low and high altiωdes, species of section Mallotiu,!, being associated with montane rain forests above 1500 m.

LOBARIA (Schreber) Hoffm. (LOBARIACEAE Chev.)

Key characters: thallus foliose, boadly lobed, green to grey to brown, lower surface continuouslyωmen加seor in a reticulate pattem in thalline depressions; photobiont a cy佃 obacteriaor green alga; apothecia laminal, inf尚quent.

Distribution: epiphytic on trunks, twigs and branches, res凶ctedto montane fagaceous forests above 1500 m. 3 species recorded in百lailand;L retigera (Bory) Trevisan occ町 S

most frequently.

島fENEGAZZIAMassal. (PARMELIACEAE Zenker)

Key characters: thallus foliose, pale grey green, lobes inflated, hollow, with round perforations on白eupper surface, lower surface dark, n紘ed,without rhizines. Dis血伊ished仕omHypogymnia by round perforations on upper s町 face.

Distribution: M. terebraω(Hoffm.) Massal is found on twigs near the summit of Doi In出anon,a new species for Thailand.

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324 P.A. WOLSELEY AND B. AGUIRRE.HUDSON

MYELOCHROA (Asah.) Elix & Hale (PARMELIACEAE Zenker)

Key characters: thallus foliose, grey, wi血simplecilia, medulla yellow to yellOW-Orange, rhizines simple or squarrose. Distinguished from other closely related Parmeliaceae with

cilia by the coloured medulla.

Distribution: locally frequent on trunks and branches in montane forests over 1000 m.

NEPHROMA Ach.例EPHROMATACEAEWetm.)

Key characters: thallus foliose, brownish, lower surface pale, finely pubescent; photobiont a cyanobacterium in Thailand; apo出eciaon lower surface of lobe apices, reniform, reddish-brown. Distinguished企omNephromopsis by the pale lower surface and absence of pseudocyphellae.

Distribution: N. helveticum Ach. (includes N. tropicum (Mull Arg.) Zahlbr.) is restricted

to twigs and branches in montane forests over 1000 m.

NEPHROMOPSIS (P ARMELIACEAE)

Key characters: thallus foliose with dark lower surface, with pseudocyphellae on白elower surface; photobiont Trebouxia; conidiomata marginal on raised stalks, apo血eciaon lower surface of lobe apices. Distinguished 合omNephroma by the dark lower surface

with pseudocyphellae.

Distribution: N. ornata (M甘IIArg.) Hue has been recorded occasionally in montane forests

above 1000 m.

NORMANDll可ANyl. (Family unknown)

Key characters: thallus of rounded shell-like squamules 1-2mm diameter, glaucous blue-grey, photobiont trebouxioid, with or without soredia.

Distribution: a monotypic genus, N. pulchella (Bo汀'er)Nyl. is restricted to undisturbed montane rainforest and usually found on old mossy岡田.

PANNARIA Delise (pANNARIACEAE Tuck.)

Key characters: thallus foliose, squamulose or crustose, pale grey to blue-grey, with or without a conspicuous blue-black hypothallus, photobiont a cyanobacterium-Nostoc; apothecia with persistent thalline exciple. Distinguished 合omParmeliella by the prescence

of pannarin (Pd + Or).

Distribution: reported by Vainio but not recently recorded.

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KEY TO LICHEN GENERA INτ百AILAND 325

PAR勘IELARIAAw出.(PAR民ffiLIACEAEZenker)

Key characters: thallus foliose, maculate with black lower surface and few rhizines, margins ciliate with conical black conidia along margin of thallus, apothecia marginal.

Distribution: P. thomsonii (Stirton) Awas. was collected by Tsuyama at Migothang in 百 ailand.

PARMELIA Ach. (pARMELIACEAE Zenker)

Key characters: thallus foliose with linear or effigurate pseudocyphellae on白eupper surface, rhizines simple or squ町 oseusually continuous to出em紅 gins.

Distribution: m組 yold literature records of出isgenus have been recently transferred to other genera in Parmeliaceae (ELIX, 1993).

PARMELIELLA Ach. (PANNARIACEAE Tuck.)

Key characters: thallus squamulose to foliose, greyish brown to lead grey, corticate above ecorticate below; rhizines grey to bluish black; hypothallus usually present, bluish black; photobiont-Nostoc; apo由ecialecideine; no lichen products detected.

Distribution: P. brisbanensis (C.Knight) P.M. Jorg. & D. Galloway is仕'equenton住unksand branches in well-lit open situations in合yevergreen forest and in mangroves.

PAR島IELINAHale (P ARMELIACEAE Zenker)

Key characters: thallus foliose, grey, lobes narrow, subrotund apices, maculate upper surface, simple cilia and rhizines, medulla white.

Distribution: not recorded from Thailand, but occurs in adjacent coun肘es.

PAR勘IELINOPSISElix & Hale (PARMELIACEAE Zenker)

Key characters: thallus foliose adnate with imbrica飽 lobesand truncate apices, grey, emaculate, cilia simple, often res凶ctedto lobe axils, medulla white, rhizines simple to sparsely dichotomously branched.

Distinguished 合omParmelinella by presence of at least some dichotomous rhizines and absence of salazinic acid.

Distribution: P. expallida (K町 ok.)Elix has been collected in百1ail佃 d泊 ChiangMai Province (specimen seen inτNS herbarium in Japan).

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326 P.A. Wo凶町YAND B. AGUIRREιHUDSON

PARMEL町ELLAElix & Hale (PARMELIACEAE Zenker)

Key characters: thallus foliose, pale grey green, adnate with imbricate lobes and inconspicuous marginal cilia in the sinuses, medulla white, lower cortex with naked marginal zone, simple white-tipped rhiz泊es,isidia abundant on lamina and on apo出ecial血allineexciple. Dis出19uishedfrom Parmelinopsis by s加.plerhizines and presence of salazinic acid.

Distribuuon: P. wallichiana (Taylor) Elix & Hale is widespread on trunks and branches 泊 deciduousdipterocarp forests and in open disturbed and modified habitats.

PELTIGERA Willd. (PEL'τlGERACEAE Dumort.)

Key characters: tha1lus foliose, brown, corticate above ecorticate below with network of raised veins and simple to fasciculate rhizines, apothecia without thalline exciple on upper surface of lobe.

Distribution: infrequent and only found on e紅白 banksabove 2000 m.

PHAEOPHYSCIA Moberg (pHYSCIACEAE Tuck.)

Key characters: thallus foliose, appr,田sedto substrate, lobes brown-grey without pruina, marginal cilia absent, upper and lower cortex present, rhizines simple; photobiont 住ebouxioid;cortical substances absent. Distinguished 合omPhyscia by brown colour of tha1lus and absence of lichen substances.

Distribution: found infrequently担金yevergreen and mon旬neforests.

PHYLLOPSORA M凶1.Arg. (BACIDIACEAE W.Watson)

Key ch町 'acters:thallus squamulose, green to white-green, on a red, black or white hypo由allus,squ釘nulesappressed佃 dplacodioid (occasionally appearing crustose) or branching and overlapping appearing foliose, corticate above, non-corticate below; photobiont gr田 n;apothecia lecideine. Distinguished from other squamulose genera by green phobiont and presence of a conspicuous felted hypotha1lus.

Distribution: epiphytic, a widespread and variable genus characteristic of the moist evergreen lowland and montane forests, also a rapid coloniser following disturbance, it is found as an indicator of former evergreen sta旬sin species-poor deciduous dipteroc紅pforests出athave been frre managed.

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KEY TO LICHEN GENERA町 THAILAND 327

PHYSCIA (Schreber) Michaux (PHYSCIACEAE Tuck.)

Key characters: thallus foliose of radiating appressed to erect lobes, pale white-grey, frequently pruinose; apothecia lecanorine: atranorin and other lichen substances present.

Distinguished from Phaeophyscia by pale grey thallus and presence of lichen substances.

Distribution: infrequent on rocks and trees in deciduous and evergreen forests.

PHYSCIDIA Tuck. (PHYSCIACEAE Tuck.)

Key characters: thallus foliose of radiating lobes, corticate above, non-corticate below with a white hypo出allus,photobiont green, isidia present or absent, apothecia lacking thalline exciple, reddish. Distinguished from Physcia which it closely resembles by the absence of a lower cortex and lecideine apothecia, and from Phyllopsora by the foliose lobes白紙紅enot attached to the substrate by the hypothallus.

Distribution: epiphytic, one species P. wrightii Tuck. is restricted to relatively undamaged evergreen lowland and montane forests.

PHYSMA Massal (COLLEMATACEAE Zenker)

Key characters: thallus foliose, lilac grey to brownish grey, homoiomerous, gelatinous when wet, apothecia lecanorine, spores simple. Distinguished from Collema and Leptogium by the simple spores.

Distribution: P. byrsaeum (Ach.) Tuck. was found at Huay Kha Khaeng, Uthai Thani Province.

POLYCHIDIUM (Ach.) Gray (PLACYNTHIACEAE D油1)

Key characters: thallus fruticose, shi凶ngblue-green, densely dichotomously branched with photobiont a cyanobacterium Scytonema.

Distribution: P. dendriscum (Nyl.) Henssen is found on mossy branches near the summit

of Doi Inthanon.

PSEUDOCYPHELLARIA Vainio (LOBARIACEAE Chev.)

Key characters: thallus foliose, grey, corticate above and below, under side pubescent with white or yellow pseudocyphellae; photobiont a cyanobacterium-Nostoc or chlorococcoid. Distinguished from Sticta by the presence of pseudocyphellae (see plate)

on白elower surface and presence of lichen substances.

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328 P.A. WOLSELEY AND B. AGUlRRE-HuoSON

Distribution: epiphytic 'On twigs and branches, P.junghuhniana (Mull. Arg.) Awast. (white

medulla, upper s田 faceshiny) and P. aurata (Ach.) Vaini'O ( yell'Ow medulla, marginal s'Oralia)紅eres住ictedt'O m'Onぬnesites泊 relativelyundisturbed fagace'Ous f'Orest 'Over 1000 m, (a species 'Of the wet tr'Opics is likely t'O 'Occ町 泊 血es'Outh; P. argyracea (Delise) Vaini'O (isidiate, white medulla).

PSORA H'Offm. (PSORACEAE Zahbr.)

区eycharacters:血allussquamul'Ose, reddish-br'Own, c'Orticate ab'Ove and bel'Ow, tha11us attached t'O substrate by hyphae, ap'Othecia lecideine br'Own t'O black.

Distribution: terric'Ol'Ous, Psora decipiens (Hedwig) H'Offm. is c'Osm'Op'Olitan and can be

expected in百lailand.

PYX町EFr. (PHYSCIACEAE Zahlbr.)

Key characters: thallus f'Oli'Ose 'Of radiating l'Obes, pale grey (t'O dark grey), l'Obes佐uncate,c'Orticate ab'Ove and bel'Ow, pseud'Ocyphellae仕equentlyprl巴sent;ph'O旬bi'Ontchl'Oroc'Occ'Oid; rhizines present; ap'Othecia lecideine. Distinguished fr'Om Dirinaria by presence 'Of rhizines

佃 dlecideine ap'O曲目ia.

Distribution: widespread in well-lit situati'Ons in l'Owland f'Orests, P. coccifera (Fee) Nyl. (red effigurate pseud'Ocyphellae devel'Oping 悶 edia)is frequent in decidu'Ous dipter'Ocarp

f'Orests, whereas P. consocians Vai凶'0(whi旬 effiguratemaculae and dactyls) is 合'equent

白r'Ough'Outincluding sec'Ondary, disturbed and urban sites, c. 8 species in百lailand.

RAMALINA Ach. (RAMALINACEAE Ag.)

Key characters: thallus frutic'Ose, pale green t'O yell'Ow-green, shrubby 'Or pendul'Ous, ph'Ot'Obi'Ont grl田 n,branches terete 'Or c'Omplanate (flattened), h'Oll'Ow 'Or s'Olid. Distinguished fr'Om Usnea when branches tere飽 byabsence 'Of cartilagen'Ous砿 is.

Distribution: epiphytic 'On can'Opy twigs, abundant in exp'Osed si同泊 m'Ontanef'Orest and c'Oastal scrub, elsewhere 'On c'Oastal r'Ocks but n'Ot 'Observed 'On也issurvey.

RELICINA (Hale & Kur'Ok.) Hale (PARMELIACEAE Zenker)

Key characters: tha11us f'Oli'Ose, l'Obes appressed, yell'Ow-green, emaculate, wi白 bulbatecilia, rhizines simple t'O sp紅 selybranched; ap'O曲目ialar凶naleperf'Orate. Distinguished 合omBulbothrix by the yell'Ow-green c'Ol'Our 'Of usnic acid.

Distribution: R. sublimata (Nyl.) Hale is epiphytic 'On can'Opy branches and saxic'Ol'Ous

ab'Ove 1000 m.

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KEY TO LICHEN GENERA町 τHAILAND 329

RELIcn可OPSISElix & Verdon (PARMELIACEAE Zenker)

Key characters: foliose, lobes narrow,叩pressed,yellow-green,出ncate阻 dsimple or agglutinated rhizines, apo白ecialaminal eperforate. Distinguished企omRelicina which also has a yellow-green出allusby the absence of cilia.

Distribution: epiphytic on well-lit trunks and branches, R. rahengensis (Vain.) Elix &

Verdon is characteristic of deciduous dipterocarp forests.

RIMELIA Hale & Fletcher (PARMELIACEAE Zenker)

Key characters: foliose, lobes broad, rounded, ciliate, upper cortex grey green with reticulate maculae becoming cracked and fissured, lower surface rhizinate to m紅'gins,rhizines simple to squarrose. Distinguished 合omRimeliella which also has a cracked and fiss町 'edupper cortex by the simple cilia.

Distribution: the sorediate species R. reticulata (Taylor) Hale & Fletcher occ町 son佐山武sand branches in evergreen lowland and mont姐 .eforests, but is most abundant and dominant in白emontane evergreen forest over 1000 m, especially in disturbed montane rainforest.

RIMELIELLA Kurok. (P AR勘ffiLIACEAEZenker)

Key characters: foliose, lobes broad rounded, cilia frequently branched, upper cortex with effigurate maculae, lower cortex with short marginal rhizines and long agglutinated rhizines. Distinguished from Rimelia by at least some cilia being forked.

Distribution: epiphytic, a specimen of R. subsumpta (Nyl.) Kurok.食omChi叩 gMaiw部

studied atτNS herbarium in Jap組.

ROCCELLA DC. (ROCELLACEAE Chev.)

Key characters: thallus shrubby,ωfted, p泊kish-grey,attached by basal holdfast, medulla within holdfast or叩 ge-yellow;photobiont Trentepohlia; apo出ecialecanorine, lru:国nalor marginal, subhymenial layers black.

Distribution: R. phycopsis Ach. recorded inτbailand on rock at Kaw Tao (Koh Tao) (PAU凶 ON,1930).

SPHAEROPHORUS Pers. (SPAHAEROPHORACEAE Fr.)

Key characters:血allus仕uticose,pale grey to grey-green, much branched, branches usually solid, white to grey-brown; photobiont田 bouxioid;apothecia terminal or subterminal on branches. This genus is currently under revision and it is likely白紙thebulk of白especies

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330 P.A. WOLSELEY AND B. AGUIRRE-Huo哩ON

belong to the genus Bunodophoron Massal.

Distribution: saxicolous and terricolous in montane紅巴asover 1500 m in adjacent coun岡田

of Vietnam, India and Malaysia. Not recorded in百 ailand.

STICTA (Schreber) DC. (LOBARIACEAE Chev.)

Key characters: thallus foliose, grey to green to brown, loosely adnate or shortly stalked and ascending, corticate above and b巴low,lower surface glabrous or pubescent with distinctly shaped cyphellae, no lichen substances, but thallus often smelling of fish when wet; photobiont a cyanobacterium -Nostoc or chlorococcoid; apo也氏ialecanorine.

Distribution: epiphytic on twigs and branches, S. nylanderiana Z油lbr.is local and res凶ctedto montane rain forests over 1500 m.

TELoscmSTES Norman (τ'ELOSCHISTACEAE Zahlbr.)

Key characters: thallus fruticose with intricate dichotomous branching, yellow to orange, glabrous to pubescent, branches terete to complanate, corticate, photobiont位ebouxioid.

Distribution: epiphytic on twigs and canopy branches, T. flavicans (Swartz) Norman is found in adjacent areas of Vietnam and Yunnan above 1000 m, not recorded in 百lailand.

UMBILICARIA Hoffm. (UMBILICARIACEAE Chev.)

Key characters: thallus foliose, single-or multi-lobed, grey-brown to black, umbilicate and attached by central strand, corticate above and below, photobiont chlorococcoid, rhizines present or absent.

Distribution: saxicolous, recorded in temperate紅'easof the Himalayas, Yunnan, T泊W加

and Malaysia, not recorded in Thailand yet.

USNEA Adans. (PARMELIACEAE Zenker)

Key characters: thallus fruticose, shrubby to pendulous, greenish grey to yellowish, attached to substrate by a basal holdfast, outer cortex present, medulla prl回 ent佃 da cartilagenous cen佐al回 is; photobiont trebouxioid. Distinguished from terete Ramalina species by出epresence of the cartilagenous central axis.

Distribution: widespread and dominant in the upper c叩 opyof montane forests over 1000 m, many species recorded in Thailand.

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KEY TO LICHEN GENERA町 THAILAND 331

XANTHOPARMELIA (Vainio) Hale (PARMELIACEAE Zenker)

Key characters: thallus foliose, lobes narrow, yellow-green, eciliate, rhizines simple.

Distinguished from other saxicolous Parmeliaceae by its yellow-green colour and absence

of cilia.

Distribution: usually saxicolous, frequent on boulders and rock surfaces.

ACKNOWLEDGMENTS

We would like to acknowledge Alan Orange and the British Lichen Society for the use of illustrations from The Lichen Flora of Great Britain and Ireland, Sean Cunningh創nfor

additional illustrations of百1aimaterial and preparation of the plates, Peter James and Per Magnus Jorgensen for discussions and suggestions, and Stephen Blackmore for provision of facilities and support at The Natural History Museum, London. We訂 ealso grateful to

the Leverhulme Trust who funded the work in Thailand and to the many百laistudents who

have s回 ggledwith the terminology and tested the key.

REFERENCES

AOUIRRE-HUDSON, B. AND P.A. WOLSELEY. 1994. Bibliography of Southeast Asian Lichenology. J. Hattori Bot.

Lab. 76: 313-350 AHTI, T. 1984.百les組側sof Cladina as a genus segr官gatedfrom Cladonia. Nova Hedwigia 79: 25-61. AWAS叩 1,D.D. (1988) A Key to the Macrolichens of India and Nepal. Hattori Bot. Lab. No 65: 207-302 CLAUZADE, G. AND C. Roux. 1985. Likenoj de okcidenta Europo Bulletin de la Societe Botanique do Centre-

Ouest Nouvelle serie-Numero Specia1e: 7-1985

ELIX J.A., J. JOHNSTON AND D. VERDON. 1986. Canoparmelia, Paraparmelia and Relicinopsis,出reenew genera in出eParmeliaceae (Lichenized Ascomytena). Mycota.xon 27: 271-282

ELIX J.A. AND M.E. HALE. 1987. Canomaculina, Myelochroa, Parmelinella, Parmelinopsis and Parmotremopsis,

five new genera in恥 P訂 meliaceae(Lichenised Ascomytina). Mycota.xon 29: 233-244. ELIX J.A. 1993. Progress in白eGeneric Delimi匂tionof Parmelia sensu lato Lichens (Ascomyco出 a:P,叩neliaceae)

組 da synoptic Key to Parmeliaceae. The Bryologist 96 (3): 359-383.

ERIKSSON, O.E. AND D.L. HAWKSWOR叩.1991. Notes on Ascomycete systematics Systema Ascomycetum 9: 1-271. GEOROE A.S., ED. 1992. Lichens-In加 duction,Lecanora1es. Flora of Australia Vol 54. Aus回 lianGovemment

Publishing Service, C組 berra.HALE M.E. 1961. Lichen Handbook. Smithsonian Institution, Washington, D.C. JAMES, P.W. AND A. HENsSEN. 1976. The Mo中hologica1and Taxonomic Significance of Cepha1odia. Pages

27-77 in Lichenology:・Progressand Problems. D.H., Brown, D.L. Hawksworth and R.H. Bailey,eds.

Systematics Association Specia1 Volume No. 8. Academic Press, London and New York. ORCHARD, A.E., ED. 1994. Lichens-Lecanora1es 2, Parmeliaceae Flora of Australia Vol 55. Austra1ian

Govemment Publishing Service, Canberra PAULSON, R. 1930. Lichens fぬmKaw Tao, an island in the Gulf of Siam. Joumal of the Siam Socieか"Natural

History Supplement 8: 99-101. POELT, J. 1966. Zur Kenntnis der Flechtenga即時 Physconia.Nova Hedwigia 12: 107-135. 町 RVIS,O.W. ET AL. 1992. The Lichen Flora of great Britain and Ireland百 eNatural 回 storyMuseum inぉ・

sωiation with The British Lichen Society, London. RIαfARDSON, D.H.S. 1975. The Vanishing Lichens David & Charles, Vancouver. SCNEIDER, A. 1897. A text-book of General Lichenology Willard N. Clute & Co. New York. SWINSCOW T.D.V. AND H. KR∞. 1988. Macrolichens of East Africa British Museum (Natura1 History) London.

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332 P.A. Wo凶 ELEYAND B. AOUlRRE-HunsON

Wm百 F.J.AND P.W. JA阻 S1985. A new guideωmicrochemical蜘 h凶quesfor identification of lichen sub-S旬ncesBrit.おhLichen Society Bulletin, No. 57.

YOSHIMURA, 1. (1978) Some lichens ofThailand collected by Danish Botanists, 1958-68. Bulletin ofthe Kochi Gakuen Junior College 9: 35-40 + 2pls.

APPENDIX. Glossary of Terms.

adnate: adjoining the surface (e.g. of the subs住ateor the白a11us).佃 astomose:net-forming. antic1加a1:of hyphae meeting the surface at right angles.

apo血ecioid:apo血ecia-like.apothecium: :加itbody of the funga1 component of a lichen, often cup-shaped, usua11y

open above, with disc exposed. 紅achnoid:web-like, of a tissue consist泊gof loosely crisscrossing funga1 hyph鵠.

ascoma: genera1 denomination for a 柑 uc旬開 containingasci and ascospores in the

ascomycetes. 出 cospore:spore of国 comycetes.

ascus: sac-like cell, in which after k紅 yogamyand meiosis the ascospores are produced.

basidiocarp: a basidium producing organ. basidiospore: spore of the basidiomycetes. basidium: cell or org阻 of血ebasidiomycetes企omwhich after karyogamy and meiosis

basidiospores are extema11y produced. byssoid: cotton-like; made up of delicate白reads.

c副 lag泊ous:面m 姐 dtough, but readily bent (refering to cen凶拡i山 Usnea).centrum: genera1 term to describe asci and hamathecium. cepha1odium: a body composed of blue-green photobiont cells and funga1 hyphae growing

the surface or within the tha11us of lichens with a green photobiont. cana1icu1ate: channelled. cilia: hair-like outgrow血sfrom the m紅g泊sor upper surface of the tha11us. chondroid (axis): the cartilagenous鼠 isoccupying the cen住a1portion of the medulla.

clavate: club-like; narrowing祖 thedirection of the base.

complanate: flattened dorsiventra11y, of a tha11us. conidiomata: hypha1 s凶 C旬rewhere conidia bearing cells and conidia are formed.

conidium: norトmotileasexua1 spore. cortex: a tissue of loose町 comp配 tedhyph即, which may appe紅 cellu1aror fi.brous,

forming the outer layer of the tha11us. corticolous: growing on the bark of trees or shrubs. corticate: having a cortex.

crustose: crust-like; i.e.血a11uss住'etchingover and firmly fi.xed to出esubs住atumby血ewhole lower surface and genera11y lacking of rhizines.

cyanobacteria: a blue-green a1ga1 cy組 obiont.dactyl: fi.nger-like protuberance. dichotomous: branching into two or more equa1 arms, usua11y repeatedly and successively. effigurate: having a defi.凶teform, not effuse. ellipsoida1: (of spore) elliptica1 in optica1 section.

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KEY TO LICHEN GENERA IN THAILAND 333

epiphyte: a plant living on another but not as a par描 ite.

epithecium: layer of tissue at the surface of an apothecium formed by the branching of the apical cells of血ep釘 aphyses.

exciple: a tissue or tissues containing白ehymenium of the apo也氏ium,or forming the walls of a peri由民ium.

自bril:a short simple branch perpendicular to the main stem (in Usnea). filamentous:血read-like,白ephotobiont forming a filament of cells which is surrounded by

出.emycobiont, or to fungal hyphae. filiform: thread-like (spore).

foliose: leaf-like, with a distict upper and lower surface. fruticose: shrubby, having an upright or hanging thallus with a radial s佐uc旬re.

fusiform: spindle-like, narrowing at both ends. gelatinous sheath: gelatinous membrane (spore); see perispore. glabrous: smooth, not hairy globose: spherical or almost so. hamathecium: neutral term used for all types of hyphae and other tissues between the asci.

hapter: an organ of attachment (of the thallus to the substrate).

heteromerous: (of a layered thallus) having出emycobiont and the photobiont in well-marked layers in the thallus.

holdfast: an expanded often disc-like, attachment of thallus to substrat怠.

homoiomerous: (of姐 unlayeredthallus) having白emycobiont and白ephotobiont evenly dis凶buted白roughthe thallus.

hyaline:住ansparent(colo町 Iess).

hymenium: the spore-be出 nglayer of a fru地 19body, contai凶ngasci, spores佃 dp館司.physes.

hypha: one of the filaments of a fungal mycelium. hypothallus: a血icklayer of hyphae, white to darkly coloured, on the under side of the

thallus and often projecting beyond its margins.

hypothecium: tissues below the hymenium. imbricate: overlapping.

isidium: a (pin-like) vegetative propagule bounded by a住ue cortex, comprising both photobiont cells and fungal hyphae; usually concolourous with the upper surface of

the thallus. leprose: having a loose powdery surface without a cortex.

lirella: long and n釘 TOWapothecium.

lirellate: lirella-shape. maculate: spotted or blotched, due to uneven distribution of photobiont cells. mazaedium: a powdery mass of spores together with disintegrating asci and p紅 aphyses.

medulla: (of lichen thallus) the loose layer of hyphae below the cortex and algallayer. muriform: brick wall-like, having transverse and longi旬dinalsepta. mycobiont: the fungal partner in the lichen symbiosis nasse: finger-like pro旬sionof the

innerp釘tof a bitunicate ascus into由eendotunica or endoascus; internal apical beak

or apparatus. oblong-ellipsoid: (of spores) having long sides parallel, and ends almost hernispherical. ostiole: cavity ending in a pore in the papilla or neck of a perithecium or pycnidium. p紅aphyses:hyphae originating from the base of the cavity, usually unbranched and not

anastomosed.

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334 P.A. WOLSELEY AND B. AGUIRRE-HUOSON

paraplectenchymatous: a tissue of::l: isodiame凶cthin-walled fungal cells_

periclinal hyphae: of hyphae, lying in the same plane as出esurface. perithecia: a subglose or flask-like ascoma.

perithecioid: peri出町ia-like.phorophyte:出e‘host'住民 ofan epiphyte.

photobiont: the algal p紅 tnerin the lichen symbiosis. phyllidium: a small corticate, scale司 like,dorsiventral s佐ucturedeveloped at the margins or

the upper surface of the thallus, narrowed or constricted at出epoint of attachment. phyllocladium: a small corticate thalline 蛇 uc旬re由atmay be granular, verrucose, coralloid,

squamiform, digitate or peltate, contai凶nga green photobiont in Stereocaulon.

placodioid: crustose at the centre, lobed and plicate at出emargins of the thallus. ple氾tenchyma:a tissue formed of fused and twisted hyphae.

podetium: lichenised stem-like portion bearing apothecia.

polarilocular: (of ascospores) bicellular, and the two cells separated by a central perforated sepωm.

proper exciple: (of an apothecium) non-lichenized excipular tissue.

prosoplechtenchyma: tissue with elongated longi同dinallyarranged hyphae (in L.Sふpruinose: having a frost-like or flour-like surface covering. pseudocyphellae: openings in the cortex of lichens where the medulla is exposed to the泊r,

but lacking specialised cells surrounding the cavity. pseudopodetia: a stalk be紅 ingone or more apo曲目ia,developmentally p紅 tof the thallus.

pseudostroma: a stroma formed of thalline tissue and remnants of host tissue. pubescent: having soft hairs. pyc凶dium:::1: flask-shape, ostiolar conidioma, consisting entirely of fungal tissue. reticulate: net-like.

rhizine: root-like hair or thread; the attachment organ of many foliose lichens Plate 1. saxicolous: growing on rock.

sessile: having no stem, attached direct1y to出esurface. sinuous: having rounded angles, wavy (of a m訂gin).soralia: soredia on larnina or margins of thallus in a variety of diffuse or delimited or

figurate pattems Plate 11. soredium: a nOIトcorticatecombination of phycobiont cells and fungal hyphae having the

appear姐 ceof a powdery granule and capable of vegetative reproduction.

spore: a uni-or multicellular reproductive structure (asexual or sexual) in fungi, bacteria and cryptog創ns.

squamule: a small separate thallus scale or lobe (less由加 5mmlong) with or without a lower cortex.

squ紅 rose:of rhizines br姐 chingat right angles e.g.brush-like. subhymenial: generative tissue below the hymenium.

stipe: a stalk supporting an apothecium. stipitate: of姐 apotheciumsupported by a stipe. stroma: a mass or ma仕ixof vegetative hyphae in or on which spores紅'eproduced often

covering several ascomata.

sublageniform: rod shaped with a swelling ne紅 butnot at one end (of pycnoconidia).

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KEY TO LICHEN GENERA IN THA且AND 335

subovate: nearly ovate, or egg-shape. sympodial: of a thallus with a main蹴 isand a succession of subsidiary lateral branches. terete: :t: circular in transverse section. terricolous: growing on the ground. 出allineexciple: (of an apothecium) lichenized excipular tissue, external to a prop町 exciple.白allus:the vegetative body of a lichen. tomentose: having a dense covering of soft matted hairs; downy. 佐田lcate:ending abruptly, as with the end cut off. urceolate: pitcher-like.

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