Table Of Content植物研究雑誌
J. Jpn. Bot.
75: 164-177(2000)
Usnea subfloridana Stirt. (Parmeliaceae) and Its Related Species
in Eastern Asia
Yo shihito OHMURA and Hiroyuki KASHIWA DANlb
a
aDepartment of Biological Sciences，Gr aduate School of Science，Un iversity of Tokyo，
Hongo 7・3・1，Bunkyo-ku，To kyo，1 13-0033 JAPAN;
bDepartment of Botany，Na tional Science Museum，
Amakubo 4-1-1，Ts ukuba，Ib araki，3 05・0005JAPAN
(Received on December 27，1 999)
Usnea subfloridana Stir .tand its related species in eastem Asia etaxonomically
訂
revised. Five species，U. florida，U. ルlvoreagens，U. glabrescens，U. wasmuthii as well
as U. subfloridana are recognized. While U. florida forms no asexual propagule，t he
other four esorediate. Soralia morphology is pticularlyuseful for distinguishing these
訂 訂
four species，s ince its origin and development are differen .tAmong the five species
treated here，U .ルlvoreagensand U. wasmuthii are newly recorded for eastem Asia.
Key words: eastem Asia，l ichenized ascomycetes，s oralia，t axonomy，U s nea
When Asahina (19 56， 1959a， 1959b) convex or concave tops echaracteristic of
紅
revised Usnea subsect. Comosae (sensu the species. However，w hen Clerc (1987a，
Motyka 1936-38) in eastern Asia， he 1987b，1 992) studied U. subfloridana and its
recognized U. comosa (Ach.) Vain.，n Offi. related species，fo llowing features of soralia
illeg. (= U. subfloridana Stirt.) and U. are also recognized as important characters;
glabrescens (Nyl.) Vain. According to him， shape and size，c onvex or concave tops，
these two species were separated mainly by presence or absence of isidiomorphs，a nd the
the soralia with convex or concave top. He degree of excavating. In addition，C lerc and
also recognized four subspecies under U. Herrera-Campos (19 97) emphasized the
comosa based on their chemical characters; importance of the origin where soralia begin
subsp. colorans Asahina，s ubsp. comosa， their development.
subsp. melanopoda Asahina and subsp. The aim of the present study is to revise
praetervisa Asahina. In addition， he U. subfloridana and its related species in
considered U. glabrescens to include three eastern Asia，b ased on the discussion of
subspecies also separated by their chemical soralia morphology.
characters; subsp. asiatica Asahina，s ubsp.
glabrescens， and subsp. pseudocolorans Materials and Methods
Asahina. In addition， Asahina (19 68) The study is primarily based on herbarium
reported U. florida (L.) F且 Wigg.from specimens housed in the Na tional Science
Taiwan，w hich is closely related to U. Museum，T okyo (TNS) as well as the
subfloridana and U. glabrescens. specimens collected by the present authors
Asahina (1956，1 959a，1 959b) recognized through field works in 1994-1997. Field
that the absence or presence of soralia with trips were made in Japan (Hokkaido，
-164-
June 2000 Journal of Japanese Botany Vo .l75 No. 3 165
Honshu，S hikoku，a nd Kyushu). The total and can be readily distinguished from the
number of specimens used for the study was others. In other four species， soralia
133. Although they all are preserved in TNS， morphology is of taxonomic importance .as
only representatives of them are cited in this discussed by Clerc (19 87b) . Although
paper. Type specimens preserved in BM，H ， soralia e variable in morphology as
紅
LINN，T NS，a nd TUR were also studied. mentioned below，t hey show rather stable
Morphological observation was made features as described under the description of
using ad issecting microscope. The diameter each species and in Table 1. Soralia are
of branches and the thickness of the cortex， elliptic or rounded in the vertical view or
medulla and axis were measured at the well- confluent each other to form irregular mass
developed thicker branches under X 50 of asexual propagules. They have convex or
magnification. The ratio of the thickness of concave tops and slightly to distinctly
the cortex，t he medulla and the axis，c an be elevated from the surface of branch which
given following Clerc (1984，1 987 a). The look like being stipitate. Cortical margins of
anatomical observation was carried out using soralia e'perpendicular to the surface of
紅
a bright field microscope. Sections for branch or are reflexed outside of soralia.
anatomical studies of apothecia were made Isidiomorphs and/or granular soredia are
by hand-razor. They were observed in GAW formed on soralia in certain species. When
solution. isidiomorphs e'formed，t hey 'every few
訂 紅
Lichen substances of specimens including or abundant.
type specimens were studied by means Soralia of this group are convex or
of thin layer chromatography (TLC) concave at the top when matured，a nd can be
(Culberson and Johnson 1982) or divided into two types; one with convex top
microchemical crystal tests (Asahina 1956) (type 1) and another with concave top (type
when necess y.Only the solvent B system 2). Convexity or concavity at the top of
紅
(hexane: methyl tert山崎11ether :f ormic soralia is correlated with the presence or
acid = 140:72: 18) was employed for TLC absence of granular soredia. When granul紅
tests. soredia eonly produced on soralium，th ey
紅
can be easily separated，re sulting the top of
Results and Discussion soralium to be concave. When isidiomorphs
Usnea subfloridana Stirt. and its related are generated on soralium， the top of
species are well delineated with the soralium becomes convex. The juvenile
following characters: 1) the fn肱 0民 erect stage of isidiomorphs sometimes looks like
to subpendent thallus with aj et black base; granul soredia， but they can be
紅
2) the terete，no t inflated，gr adually tapering distinguished by the presence of cortex on
branches with mat surface and usually with the surface of granules.
numerous lateral branches and fibrils; 3) the Soralia of the type 1a re developed from
presence of veucoseto cylindrical papillae the top of eroded papillae，a nd those of type
町
on thicker branches; 4) production of 2 are developed directly from cortex.
depsides or depsidones (Pigs. 1-2，Ta ble 3) . Although four origins of soralia in Us nea are
As a result of the present study，f ive portedby Clerc and his co-workers (Clerc
問
ιρ
species， U. florida， uvoreagens， U. 1987 a，C lerc and Heera-Campos 1997，
町
glabrescens， U. subfloridana and U. Herrera-Campos et al. 1998)，w hich e'1)
紅
wasmuthii are recognized in the present area. the top of eroded papillae，2 ) the scars of
Of the five species of the present group，U . detatched fibrils，3) the cracks，a nd 4) the
florida is the only species forming no soralia cortex，t he origins from sc sof detatched
紅
166 植物研究雑誌第75巻第3号 平成12年6月
Fig. 1. Diagnostic characters of Usnea subfloridana and its related species. A. Sterile thallus of
ιsubfloridana. (Yanagawa，M t. Yatsugatake. July 25，1 959，M . Togashi and S. Kurokawa
s.n.). B. Papi1lae ofι subfloridana on thicker branches. (y. Ohmura 2900a). C. Fertile
thallus ofι florida. (collector unknown 106). D. Jet black base of thallus ofι
glabrescens. (y. Oh立lUra3824b). Scales =1 c m (A and C)，a nd 1m m( B and D).
fibrils and cracks enot predominantly cortex，th eir cortical margins are slightly or
紅
involved in this group. When soralia e distinctly reflexed outside as they grow. This
紅
developed from the top of eroded papillae， type of soralia may be caused by the pressure
they are usually stipitate and their cortical of increasing of hyphae and algae under
margins are not reflexed outside of soralia. the cortex. This type is predominant in
When soralia are developed from the top of U. fulvoreagens， U. glabrescens and U.
very early stage of papillae，t hey look like wasmuthii. Thus，t he soralia morphology is
sessile even if they increase in mass. correlated with their origins and ontogeny，
However，in the case of development from and considered to be of taxonomic
mature stage of papillae，s oralia is stipitate importance to distinguish the species of this
even if they enlarge their masses. This type group.
ル
is cOrIlmonly found in U. subfloridana. In addition，so ralia of U. lvoreagens，U.
When soralia紅'edeveloped directly from subflol幼 naand U. wasmuthii are confluent
June 2000 Journal of Japanese Botany Vo .l75 No. 3 167
Fig. 2. Soralia of Usnea subfloridana and its related species. A. Convex soralia with numerous
isidiomorphs (ι subfloridana，Y . Ohmura 2900a). B. Concave soralia with slightly excavating
surface forming e11iptic masses of soredia. (ι wasmuthii，T. Matsuoka s.n.). C. Concave soralia
with slightly excavating surface forming rounded masses of soredia (U. glabrescens，Y . Ohmura
3824b). D. Concave soralia with deeply excavati昭 surface，and the cortical margin distinctly
reflexed (U.かlvoreagens，H. Kashiwadani 19907). Scales =1 m m.
with adjacent soralia as they grow，w hile papillae vydepending on their devel-
紅
those of U. glabrescens are more or less opmental stage and/or by environmental
discrete when they contact with an adjacent conditions. The larger projection， which
soralium as ported by Clerc (19 92) . contains medullary tissue，i s called a tu-
陀
Although the nature of confluence or bercle by Swinscow and Krog (1979). How-
discreteness of soralia has not been well ever，t his structure is treated here as a
studied， this is usually correlated with papilla，s ince presence or absence of me-
another feature (e.g.，c hemistry). Therefore， dullary tissue in papillae is variable among
whether soralia are confluent or discrete can sorediate species of the genus Us nea.
be accepted as one of taxonomic characters. Although the ratio of thickness of the
Papillae are formed on the surface of cortex，m edulla and axis (CMA) of each
thicker branches in all treated species. species was measured，n o notable difference
Although three types of papillae，h emi- was found among them (Table 2).
spherical，v errucose (or conical) and cylin- The chemistry of each species was
drical，a re known in Us neαas reported by examined by means of TLC methods，a nd
Swinscow and Krog (19 79) and Clerc the results are shown in Table 3. As in most
(1987 a)，pa pillae of U. subfloridana and its lichen species with pale yellow thalli，us nic
related species become cylindrical when acid is constantly produced in the cortex of
matured. However，h eight and amount of all species treated. Atranorin， another
】
小
∞
Table 1. Soralia morphology in Usnea subfloridana and its related species in eastem Asia
Origin of Granular
Species Shape and Size Stipes Cortical margin Tops Isidiomorphs
dominated soralia soredia
U.floridα
U.fulvoreαgens cortex confluent，ir regular，la rger slightly stipitate distinctly reflexed concave，d eeply present absent
than branch diameter excavating into nearly
central axis
U. glα~brescens cortex :t discrete，ro unded，la rger slightly stipitate slightly reflexed concave，slightly present rarely present
than branch diameter excavating and never
茜
reached to central axis 書
u U. co wu a'bL p smo ur thJu iα in a pcaopirltleaxe ctchooannnff llbuuerennattn，，ciher1 lrideipgtauimlcea rttoe， rs m aller OS剖『れ引引・・吐いMUbwnrσbn-ubw由民us p1tmp ・ 、，，且•・・・・也64.，，‘ •IEyh ル nsloitgrhetlfyl erxeefdl exed ccoonncvaevxe ，sl ightly parbesseenntt cporcoecmsaemsnoiton nlayl1 y 出司判間帯砕
i汀egularlyrounded，larger excavating and never present
than branch diameter reached to central axis 瀧
斗
凶
拙
附
鵜
凶
山
叩
Table 2. The ratio of thickness of the co巾 x，medulla and axis (CMA) in Usnea subfloridana and its related species. Mean (italic numeral)，s tandard deviation
(no omament numeral) and extreme values (in the parenthesis) eshown，n = to tal number of samples examined
訂
Species Cortex (% of radius) Medulla (% of radius) Axis (% of diameter) Diameter of branch (mm) n
U.florida (8到-11-(13) (13)-16-(17) (42)-46-(48) (1.1)- 1.2-( 1.3) 3
U. fulvoreagens (7.4) -9.2-10-12-(13) (14 )-18-22-26ー(28) (24)-28-36-44ー(51) (0.7)-0.9-1.1-1.3-(i.4) -1
E
U. glabrescens (8.7)-11-(14) (18)-21-(25) (31)-37-(46) (1.0)-1.2-(1.3) O3
U. subfloridanα (7.3)-9-11-13-(16) (9.1)-14-18-22-(27) (27)-34-41-48-(58) (0.5) -0.8-1.0-1.2ー(1.6) OA0
U.wαsmuthii (6.5)-8.7-11-13ー(17) (7.3)-1牛18-22ー(29) (29)-37-43-49-(57) (0.7)ー0.8-1.0-1.2ー(1.8) 斗2
川
市
回
開
-
M
明h
。
泊
June 2000 Journal of Japanese Botany Vol. 75 No. 3 169
common cortical substance in lichens，i s is only presence or absence of a
demonstrated in some specimens of U. hydroxy 1r adical. Therefore，t hey every
紅
fulvorωgens， though it is a very re closely related substance each other
， 紅
substance in the genus Us nea. It should chemically. The occuenceof zeorin in U.
町
be noted that atranorin is produced in some ルlvoreagensis p ticularlyinteresting from
紅
species of Ramalina (Stevens 1987)，i n a chemotaxonomic viewpoint. Zeorin is
which usnic acid is a constant cortical synthesized through the mevalonic acid
substance，a nd is considered to be accessory. pathway，w hile depsides and depsidones are
As regard to depsides， following three produced through the acetylpolymalonyl
聞
biosynthetically different groups are consid- pathway. Thus，th e presence of zeorin can be
ered: 1) diffractaic acid and its related sub- considered to have rather important
stances group， 2) barbatic and 4-0- taxonomic value，e specially in separating
demethylbarbatic acids group， and specles.
ル
3) squam副icand/or thamnolic acids group. Although U. lvoreagens， U. sub-
In the case of presence of diffractaic acid，it floridana and U. wasmuthii produce several
always follows barbatic，4 -0-demethylbar- chemical races as shown in Table 3，n o
batic，b aeomycesic，a nd squamatic acids. morphological or ecological difference was
According to Yo shimura and Kurokawa recognized among them，a nd it is considered
(19 80) and Yo shimura et al. (1992)，th e 0- that each chemical race has no necessity to
methyl of diffractaic acid is partially substi- be given any taxonomic ranks.
tuted by H0 to form barbatic acid (= 2-0-
2
demethyldiffractaic acid). Similarly，t he 0- Taxonomic treatment
methyl of barbatic acid may be substituted Usnea florida (L.) Weber ex F且 Wigg.，
by H0 to form 4-0demethylbarbaticacid. Primit. Flor. Holsat. 91 (1780).
2 ・
In addition，o xidation of the methyl radical Lichen floridus L.， Spec. Pl. 1: 1156
of barbatic acid forms baeomycesic acid，an d (17 53) .Le ctotype (selected by Clerc 1984) :
further oxidation of its radical of Sweden?， Linne? (LINN!). Chemistry:
baeomycesic acid form squamatic acid. alectorialic (in apothecia) ， bourgeanic，
Therefore， these substances are closely hypothamnolic，t hamnolic and usnic acids
related to each other biosynthetically as well (by Jφrgensen et al. 1994).
as chemically. The presence or absence of Thallus fruticose，er ect，u p to 6.5 cm long，
one or af ew of them can be considered jet black at the base; branching anisotomic-
to have certain taxonomic value， even dichotomous; branches mat on the surface，
though the concentration of each substance lacking pseudocyphella，t erete，n ot inflated，
is variable as mentioned in the description gradually tapering，wi th numerous fibrils and
of U. fulvoreagens in the present study. lateral branches， 1.1-1.3 mm in diameter;
The biosynthetic and/or chemical relation- lateral branches not constricted at the base;
ship between squamatic and thamnolic papillae common on thicker branches，
acids have not been studied extensively cylindrical; soralia absent. Cortex moderate
so far. However， they are considered in thickness，8 .9-13% of the radius. Medulla
as biosynthetically closely related substances dense，mo derate in thickness，1 3-17 % of the
as discussed by Culberson (19 69)， and radius. Axis solid，m oderate in thickness，
they often replaces each other in paired 42-48% of the diameter. Apothecia
races in lichens (Fiscus 1972). Norstictic common，su bterminal on terminal and lateral
and salazinic acids belong to depsidone. branches，up to 6.5 mmin diameter，fl at; disc
The chemical difference between them pruinose; epihymenium 4-15μm high;
170 植物研究雑誌 第75巻 第3号 平成12年6月
Table 3. Lichen substances in Usnea subfloridana and its related species. +， present.一，absent. tr，tr ace. ::I:， ac-
cessory substance. Atr，a tranorin; Usn，u snic acid; Bar，b arbatic acid; Dif，d iffractaic acid; 40B，4 -0
ω
demethylbarbatic acid; Bae，ba eomycesic acid; Squ，sq uamatic acid; Tha，th amnolic acid; Nor，no rstictic acid;
Sal，sa lazinic acid; Zeo，ze orin
Medullary substance
Cortical
substance
Species Race Depside D'epsidone Terpenoid
Atr Usn Bar Dif 40B Bae Squ Tha Nor Sal Zeo
U.florida + +
U.ルlvoreagens Race 1 + 十 +
Race2 + + tr + tr tr tr + +
U. glα:brescens + + +
U. subfloridαm Race 1 + +
，
Race2 + +
Race3 + + +
Race4 + +
U. wasmuthii Race 1 + +
Race2 + + + +
Race3 + +
hymenium 56-70μm high; hypothecium 30- record for the species in the present area，
60μm high; spores 8-10 Xι6μm. though it is widely distributed in Europe
Chemistry: usnic and thamnolic acids. (Clerc 1984).
From the lectotype of the present species， Specimen examined. Taiwan，P rov. Taitung，M ~.
Jφrgensen et al. (19 94) reported the Dai-wu，o n bark，el evation about 2000 m，S eptember，
occuenceof alectorialic (in apothecia)， 1958，co llector unknown 106.
町
bourgeanic and hypothamnolic acids as well
Usnea fulvoreagens (RasaneωRasanen，
as usnic and thamnolic acids. The former
Lich. Fenn. Exs. .13( 1935).
three substances can be considered accessory
Usnea glabrescens (Nyl.) Vain. v
as pointed out by Clerc (1984). A specimen 紅.
βtlvoreagens Rasanen，Fl echt. Estlands 1，34 :
collected in Taiwan well agrees with the type
20 (1931) . Holotype: Russia， Karelian
and coincides very well with the description
given by Clerc (1984). Republic，K l. Kurkijoki，L apinlahti，A ugust
22，1 923，R asanen s.n. (H 1). Chemistry:
Us nea florida is frequently confused with
U. masudana Asahina，U. ogatai Asahina，U. usnic， norstictic， menegaZZIalC， stIctIc，
constictic acids，a nd zeorin 佐(ace).
orientalis Motyka，U . pseudogatai Asahina，
Thallus fruticose，er ect to subpendent，u p
U. shimadai Asahina and U. sinensis
to 16 cm long， jet black at the
Motyka，si nce they all occur in Japan and/or
base; branching isotomic-or anisotomic-
Taiwan and form similar thalli lacking
dichotomous; branches mat on the surface，
soralium with numerous apothecia.
However，U . floridlαis easily distinguished lacking pseudocyphella，t erete，n ot inflated，
gradually tapering，wi th numerous fibrils and
from the other six species by the jet black
lateral branches，0 .7-1.4 mm in diameter;
base of thallus and by the production of
lateral branches not constricted at the base;
thamnolic acid.
papillae common to sp on thicker
Asahina (19 68) reported the occuence 町田
町
branches，c ylindrical to veucose;soralia
of U. florida from Taiwan. This is the only 町
June 2000 Joumal of Japanese Botany Vol. 75 No. 3 171
common，f ormed mainly on terminal and 2ぱ0fU.ルρuルvoreωαge附n1臼s加inthe pr陀es詑e凶ntp叩apeぽr紅e
lateral branches， developed from cortex， also known from Europe and North America
concave at the top，d eeply excavating often (Cαler'c1992，H alo
民
into nearly central axis， without a .l1992).
ル
isidiomorphs， sessile or often slightly In Japan，U . lvoreagensis distributed in
stipitate，o ften conf1uenting each other to Hokkaido to central Honshu，w here it grows
form irregular masses of soredia larger than on tree barks such as Abies，L arix and Salix.
the branch diameter， cortical margin It is found at elevations between 50 and 1800
distinctly ref1exed; soredia granular. Cortex m (at lower elevation in Hokkaido and at
moderate in thickness，7 .4-13 % of the higher elevations in central Honshu) In
，
radius. Medulla moderate to dense，1 4-28 Taiwan，i t grows on barks at elevations
% of the radius. Axis solid，m oderate in between 1500 and 2600 m. Although U.
thickness， 24-51 % of the diameter. βρtlルνlJor陀e，αgenshas been reporte吋dfrom Europe
Apothecia not seen. and North America (Cαler'c1992，Ha lω10
，氏
Chemistry: Race 1，u snic，n orstictic acids， al. 1998，P urvis et al. 1992)，th is is the first
and zeorin; Race 2， usnic， norstictic， record from eastern Asia.
diffractaic acids and zeorin， and trace Representative specimens examined. Race 1.
amount of barbatic，4 O-demethylbarbatic， JAPAN. Hokkaido. Prov. Nemuro: Ochiishi，o n bark
・
baeomycesic，s quamatic acids，a nd atranorin of Abies sachalinensis， elevation about 50 m，
September 1，1 965，S . Kurokawa 65808. Honshu.
(土).
Prov. Kai: Tokusa Pass，A ugust 7，1 953，S. Kurokawa
Usneaかlvoreagensis distinguished from 521244. Prov. Shinano: en route from Azusayama to
other related species by having deeply Jumonji Pass，K awakami-mura，M inamisaku-gun，el e-
excavating SOI叫iawithout isidiomo中h，and vation about 1600 m，J une 13，1 983，H. Kashiwadani
by the presence of norstictic acid and zeorin. 19907. TAIWAN. En route from Ssu-yuan to To-chia-
tun Shanm Mt. Nanhuta Shan，H oping，o n bark of
Two chemical races were recognized
ル Salix sp.，el evation 1900-2250 m，N ovember 9，1 989，
within U. lvoreagensfrom the present area
H. Kashiwadani 35767.
as shown above. No morphological or Race 2. JAPAN. Honshu. Prov. Shinano: ca. 3k m
ecological difference was found between ESE of Azusayama，K awakami-mura，M inamisaku
・
Race 1 and Race 2. Therefore， it is gun，on bark of Larix kaem/).作ri，elevation 1460-1500
considered that the chemical differences of m，D ecember 9，1 996，Y . Ohmura 2906. TAIWAN.
Mt. Nan-FuTa-San，elevation 1500-2400 m，J anuary
this species have no taxonomic value. ・
19，1 964，S. Kurokawa 963.
Although diffractaic acid is always present in
the thalli of Race 2，it s concentration is not
Usnea glabrescens (Nyl.) Vain.，M eddel.
uniform even within at hallus. When two or
Soc. Fa. Fl. Fenn. 48: 173 (1925).
three different p tsof the thallus were
訂 Usnea barbata (L.) F. H. Wigg. var.
examined，t he concentration of the acid is
glabrescens Ny .lin Vainio Meddel. SocFa.
，
app entlyvariable， being detected as a
紅 Fl. Fenn. 2: 46 (1878). Holotype: Finland，
distinct or ap ale ambiguous spot on TLC
Tavastia australis，K o中ilahti，Tianea，1 873，
plate.
E. Lang s.n. (TUR 1). Chemistry: usnic，
Although menegazziaic， stictic and
norstictic， protocetraric (trace)， salazinic
constictic acids were detected in the holotype
acids，a nd atranorin.
specimen of U. fulvoreagens ，th ey were not
Thallus fruticose，e rect to subpendent，u p
detected in specimens in the investigated
to 16 cm long，j et black at the base;
area. However，n o morphological difference
branching isotomic-dichotomous; branches
was observed between the type and other
mat on the surface，l acking pseudocyphella，
specimens. It is noted that Race 1a nd Race
172 植物研究雑誌第75巻第3号 平成12年6月
terete，n ot inflated，g radually tapering，w ith fulvoreagens.
sparse fibrils and lateral branches，1 .0-1.3 In Japan， U. glabrescens is found in
mm in diameter; lateral branches not central Honshu，w here it grows on tree bark
constricted at the base; papillae common on (e.g.，Sa lix) at elevations between 1440 and
thicker branches，ve ucose;soralia common， 1800 m. In Taiwan，th is species grows on
町
formed mainly on terminal and lateral twigs (e.g.， Osmanthus) at elevations
branches，d eveloped from cortex，c oncave at between 2400 and 2600 m. Us nea
the top， slightly excavating， rarely with glabrescens is widely distributed in boreal to
isidiomorphs， sessile or often slightly temperate regions in Europe，e おast旬ernAsia
stipitate，m ore or less discrete，r ounded in and North America (As鈎a副h廿由ina 1959a丸，
shape， sometimes becoming larger than Halo
branch diameter， cortical margin usually Specimens examined. JAPAN. Honshu. Prov.
slightly reflexed; soredia granular. Cortex Shinano: en route from Gyozya-goya to Minoto-sanso，
moderate in thickness， 8.7-14% of the Yatsugatake Mts.，Chino-city，on bark of Salix sp.，el e-
vation 1740-1800 m，A ugust 29，1 997，Y . Ohmura
radius. Medulla moderate to dense，m oderate
3824b. TAIWAN. Mt. Nan-Fu-Ta-San，o n twigs of
in thickness，1 8-25% of the radius. Axis
Osmanthus bioritsuensis， elevation 2400-2600 m，
solid，m oderate in thickness，3 1-46% of the January 20，19 64，S. Kurokawa 1088.
diameter. Apothecia not seen.
Chemistry: usnic，n orstictic and salazinic Usnea subfloridana Stirt.，S co .tNatur. 6:
acids. 294 (1882).
Although atranorin and a汀aceamount of Holotype: Scotland， Perthshire， Killin，
protoce住aric acid were detected in the July 19， 1881， Stirton s.n. (BM!).
holotype specimen of U. glabrescens，t hey Chemistry: usnic and thamnolic acids.
were not detected in specimens in the Lichen comosus Ach.， Kgl. Vetensk.
investigated 'ea.However，n o morpholo- Acad. Nya Handling. 209，t ab. 8，f ig. 1
訂
gical difference was observed between the (1795) -Usnea plicata (L.) F.H.Wigg. var.
type and other specimens. The presence or comosa (Ach.) Ach.，M ethod. Lich. 311
absence of them seems to have no taxonomic (1803); Roh ，.l Deutsch. Fl. 3(2): 144
value. (1813) -U. florida v comosa(Ach.)
紅・
Asahina (19 59a，1 959b) reported three Biroli，Fl ora Asconiens. 2: 199 (1808) -u.
subspecies under U. glabrescens which were comosa (Ach.) Vain.，i n Norrl. and Ny ，.l
separated by the chemistry: 1) subsp. Herb. Lich. Fenn. exs. No. 457，4 58 (1921);
glabrescens characterized by norstictic acid， Motyka，L ich. Gen. Usnea Stud. Monog ，.r
2) subsp.ωiatica Asahina by salazinic acid， pars. sys ，.t264 (1936); non U. comosa Pers.
and 3) subsp. pseudocolorans Asahina by (1826). Holotype: Sweden， Acharius?
thamnolic acid. In the present paper， (H !). Chemistry: lic，thamnolic and
凶
however， subsp. asiatica and subsp. hypothamnolic acids (annotated by Clerc).
pseudocolorans are treated as synonyms of Us nea comosa subsp. colorans Asahina，
U. wasmuthii (see note under U. waSl thii). Lich. Jap. 3: 94，1 956，s yn. nov. Lectotype
仰
Usnea glabrescens resembles U. (selected here): Jap an，H onshu，P rov. Kai，
ル
lvoreagensin having concave soralia and Yo shida-guchi l-gome，M t. F吋i，August 10，
in producing norstictic acid. However，it can 1952，M . Togashi s.n.，pr . m司.p. (TNS 1).
be distinguished from U.かlvoreagensby the Chemistry: usnic and thamnolic acids.
discrete soralia with slightly excavating Us nea comosa subsp. praetervisa
surface. It also differs in the absence of Asahina，L ich. Jap. 3: 95，1 956，s yn. nov.
zeorin which is constantly produced in U. Lectotype (selected here): Japan，H onshu，
June 2000 Journal of Japanese Botany Vol. 75 No. 3 173
Prov. Kai， Asahigaoka， Lakeside of The type specimen of U. comosa subsp.
Yamanaka， Nakano-mura， Minami-Tsuru- colorans includes 95 individuals in ap acket.
gun，J uly 30，1 954，S . Kurokawa 540328 Seventy-nine of 95 contain thamnolic acid.
(TNS 1). Chemistry: usnic and norstictic The thalli containing thamnolic acid well
acids. coincide with the protologue given by
Thallus fruticose，er ect to subpendent，u p Asahina (1956). The rest is am ixture of U.
to 13.5 cm long，j et black at the base; subfloridana containing squamatic acid，U .
branching anisotomic- or isotomic- hakonensis，U . pangiana and U. wasmuthii.
dichotomous; branches mat on the surface， Specimens containing thamnolic acid were
lacking pseudocyphella，t erete，n ot inflated， designated here as the lectotype of U.
gradually tapering，u sually with numerous comosa subsp. colorans， which is well
fibrils and lateral branches，0 .5-1.6 mmi n identical with the type specimen of U.
diameter; lateral branches not constricted at subfloridana.
the base; papillae common on thicker When Asahina (19 56) described U.
branches，v eucoseto cylindrical; soralia comosa subsp. praetervisa，h e cited three
町
common，f ormed mainly on terminal and specimens: two specimens collected in Japan
lateral branches，d eveloped from the top of (Nakasaroma， Kitami， Hokkaido; and
eroded papillae，c onvex at the top，u sually Asahigaoka，Y amanaka，M t. Fuji) and one
with numerous isidiomorphs，se ssile or often in China (Kanto，K inso，K onshun). How-
distinctly stipitate，s ometimes confluenting ever，t he specimen collected from Naka-
each other forming irregular masses of S紅omawas not located in TNS. Speci
四
soredia， usually smaller than branch mens collected from Asahigaoka and China
diameter， cortical margin not reflexed; agree well with the protologue given by
soredia farinose. Cortex moderate in Asahina (1956). The collection from Asa-
thickness，7 .3-16% of the radius. Medulla higaoka (S. Kurokawa 540328，J uly 30，
moderate to dense，9 .1-27% of the radius. 1954) which is marked as‘'Typus" on the
Axis solid，m oderate in thickness，2 7-58 % herbarium packet by Asahina is selected here
of the diameter. Apothecia very rare，la teral as the lectotype. The lectotype of U. comosa
on terminal and lateral branches，u p to 3.0 subsp. praeterνisαis well identical with
mmi n diameter，c up-shaped; disc pruinose; Race 4o f U. subfloridana.
epihymenium 6-18μm high; hymenium 60 Therefore，U . comosa subsp. colorans and
-80μm high; hypothecium 60-90μm high; subsp. praetervisa are reduced to synonyms
spores 8-10 X4 -6μm. of U. subfloridana in the present paper.
Chemistη: Race 1，u snic and thamnolic Asahina (1956) described one more
acids; Race 2，u snic and squamatic acids; subspecies，s ubsp. melanopoda，u nder U.
Race 3，u snic，t hamnolic and squamatic comosα. For subsp. melanopoda，s ee note
acids; Race 4，us nic and norstictic acid. under U. wasmuthii.
Although this species has been well Us nea subfloridana is easily distinguished
known as U. comosa (Ach.) Vain. (1921) mo中hologicallyfrom the related species by
by Japanese lichenologists， the name is having convex soralia. Four chemical races
illegitimate because it is al ater homonym of were recognized in U. subfloridana from the
U. comosa Pers. (1826) [ICBN 53.1 present area as shown above. The most
(Greuter et al. 1994)]. It is replaced by the common race was the Race 1 (thamnolic
name U. subfloridana Stirt. which has acid containing race， 74%) . Race 2
priority [ICBN 58.1 (Greuter et al. 1994)] (squamatic acid containing race，1 6 %) and
(Laundon 1965). Race 4( norstictic acid containing race，9 %)
