Bull. Natn. Sc i. M us ., Tokyo, Ser. B, 27(2, 3). pp. 35-46. September 22, 200 I

Typification of Taxa of Hypogy mnia (Parmeliaceae)
Described from Japan and Sakhalin
Kozo Yoshida t and Hiroyuki Kashiwadani 2
1

Saitama Museum of Natura l History, 1065- 1 agatoro, Chichibu, Saitama 369- 1305, Japan
E-mail: kozoy@beige.ocn .ne.jp
2
Department of Botany, Nationa l Science Museum , 4- 1- 1
Amakubo , Tsukuba, lbaraki 305- 0005, Japa nE-ma il : hkashiwa@kahaku.go.j p

Abstract Ten taxa belonging to the genu s Hypogymnia were described under the
gen us Parmelia from Japan a nd Sakhalin before 1956 by vari ous authors(Zahlbruckne r 1927, Hillmann 1938, Asahina 1935, 1950, 195 1, 1952): P elongata
var. stricta, P enteromorpha f. inactiva, Pfragil/ima , Pfujisanensis, P hypotrypella,
P metaphysodes, P nikkoensis, P pseudophysodes , P pseudophysodes f. reagens andP submundata f. co lorans. They are typified by designatin g a ho loty pe, lectotype or
neotype for each taxon . As a result of studies o n their morpho log ica l a nd che mi ca lfeatures, a new comb ination, Hypogymnia stricta ( Hillma nn ) K. Yoshid a a nd a newname , H. asahinae K. Yos hida, a re proposed.
Key words: Hypogymnia , 1-fypogymn ia asahinae, Hypogymnia stricta, Japan ,Sakha li n, typification, lichen

During a study Japanese taxa of the genus Hypogymnia , an attempt has been
made to typ ify taxa described unde r the ge nus Parmelia from Japan and Sakhalin before 1956. Zahlbruckner ( 1927) described Parmelia nikkoensis and Hillmann ( 1938)
descri bed P fragillima and P elongata var. stricta. Through ftori stic studi es of Japanese Parmelia sens. lat., Asahina ( 1935 , 1950, 1951 , 1952) added seven new taxa.
These taxa were described under the genus Parmelia and Rassadina ( 1956, 1960,1967) a nd Kurokawa ( 1971) transferred them to Hypogy mnia. Some of them werecited in reg ional revision s of the genus Hypogymnia by Krog ( 1968), Ohlson ( 1973)
and Lai ( 1980). However, they were not typified clearly in these studies. In the prese nt study, typification has been made by des ignating a ho lotype, lectotype or neotype for each taxon and clarify ing their morphologica l features and chemical ingredients. Even though Hypogy mnia pseud(o)enteromorpha M. J. Lai (Lai, 1980) and H.
hokkaidensis Kurok . (Kurokawa & Nakanishi , 197 1) were described after 1956 on
the basis of material s collected in Japan, they are not menti oned in the present paper,
sin ce they have been well typifi ed.

36

Kozo Yoshida and Hiroyuki Kashiwadani

1.

Parmelia elongata var. stricta Hillmann
Parmelia elongata var. stricta was described by Hillmann ( 1938) on the basis ofa single specimen collected on Mt Fuji by Asahina, but unfortunately the holotype
was destroyed at Berlin as stated by Krog ( 1968), Hawksworth (197 4) and Laundon(1979). According to Asahina (1952), two duplicates of the type were preserved in
his private herbarium, which was later transferred to TNS. One of them is annotatedas "Parmelia elongata var. stricta Hillm. , Det. Hillm." by Asahina. This specimen isdesignated here as a lectotype of var. stricta (Fig. I a) following Article 9. 10 of theCode (Greuter et al. 2000).
The present taxon is characterized by I) loosely adnate thalli composed of linear
elongate pendulous lobes with very few minute lobules, 2) absence of soredia andisidia, 3) lobes with very narrow black rim, 4) negative color reaction with P in the
medulla (physodalic acid absent) . Thus it can be considered to be a distinct species
separated from the esorediate morph of Hypogy mnia vittata, in which black marginal
rim is very wide and often wider than glaucous grey zone as pointed out by Bitter(1901), Lai (1980) and Thomson (1984).
Hypogymnia stricta (Hillmann) K. Yoshida, comb. nov.
Parmelia elongata var. stricta Hillmann, Repert. Spec. Nov. Regni Veg. 45 : 171 .
1938-Parmelia vittata f. stricta (Hillmann) Asahiana, J. Jpn . Bot. 26 : 97 . 1951 Hypogymnia vittata f. stricta (Hillmann) Kurok., Mise. Bryol. Lichenol. 5: 130.
1971.
Type. Ohmiya-guchi 2-gome, Mt. Fuji , Prov. Suruga, 6 July 1925, Y. Asahina 49(lectotype in TNS).
Chemistry. Atranorin, 2 '-0-methylphysodic acid, physodic acid and 3-hydroxyphysodic acid.
Parmelia enteromorpha f. inactiva Asahina.
When Asahina (1952) described P enteromorpha f. inactiva , he cited three specimens: CD Mt Iide, Prov. Uzen (erroneously stated as Prov. 1washiro in Asahina 1952),@ Mt Morrison, Taiwan and @ Aikawa, Sakhalin. In specimen CD, the black marginal
rims of lobes are conspicuous and vividly visible from above; vittatolic acid is present along with atranorin and physodic acid as in Hypogymnia vittata (Ach.) Parrique. Specimen @ is similar to H. stricta (=Parmelia elongata var. stricta ) in morphology and chemistry. Thus, specimens CD and @ are identified with H. vittata andH. stricta, respectively. In contrast, specimen @ agrees well with the original description given by Asahina; vittatolic or 2' -0-methylphysodic acid as well as 3-hydroxyphysodic acid were not determined by TLC (Table I and Nuno, 1964). Thus, specimen @ is designated as the lectotype of the present taxon (Fig. 1b).
Asahina ( 1952) considered the present form to be closely related to the Japanese
population of Parmelia enteromorpha, which is currently named Hypogy mnia
pseud(o)enteromorpha M. J. Lai. However, it is quite different from the latter species2.

Typification of Taxa of Hypogymnia Described from Japan and Sakhalin.

·.. : ·b
Fig. I. a) Lectotype of Parmelia e/ongata var. stricta (Asahina 49 in TNS).
enreromorpha f. inactiva (Asa hin a s. n. in TNS). (Bar= I cm).

b) Lectotype of P.

37

Kozo Yoshida and Hiroyuki Kashiwadani

38

Table I . Chemical constituents of Hypogymni a taxa described from Japan and Sakhalin before
1956. Atra: Atranorin, Usni: Usnic acid, 2'0M: 2' -0-Methylphysodic ac id, Pdic : Physodic acid,
3-Hyd : 3-Hydroxyphysodic acid, Pdalic : Physodalic acid, Vitta: Vittatolic acid, Proto: Protocetraric ac id, +: chemical constituent present, blank: chemical constituent absent
Taxon

SpecimensAtra

P elongata
var. stricto
P enteromorpha
f. inactiva
Pfragillima
P fujisanensis
P hypotrypella
P metaphysodes
P nikkoensis
P pseudophysodes
P pseudophysodes
var. reagens
P submundata
f. co/orans

Lectotype+

Lectotype+

NeotypeHolotype
Lectotype
Lectotype
Holotype
Lectotype
Lectotype++

Holotype

Usn i

2'0M

Pdic

3-Hyd

+

+

+

Pdalic

Proto

+
+
+

+
+

+
+

+

+
+
+

+

+
+
+
+

+

+

+

+

+

+

+
+
+
+

Vitta

+

+
+

+

not only by the lack of physodalic acid but also by the adnate thalli which lack pendulous lobes. It also resembles H. stricta, from which it is clearly distinguished by
the lack of pendulose lobes and the lack of 3-hydroxyphysodic and 2'-0-methylphysodic acids. Thus, the present taxon can be regarded as a distinct species.
Since the epithet "inactiva" is pre-empted by Hypogymnia inactiva (Krog)
Ohlson (Ohlson, 1973), a species distributed from California to southern Alaska inwestern North America, a new name is proposed for this taxon as follows.
Hypogymnia asahinae K. Yoshida, stat. & nom. nov.
Parmelia enteromorpha f. inactiva Asahina, Lich. Japan 2: 37. 1952- Hypogymnia enteromorpha f. inactiva (Asahina) Kurok., Mise. Bryol. Lichenol. 5: 129. 1971.
Type. Aikawa, Sakhalin, 22 July 1932, Asahina s.n. (lectotype in TNS).
Chemistry. atranorin and physodic acid.

3.

Parmelia fragillima Hillmann
The present species was described on the basis of a single specimen collected at
Hakutyoko in Sakhalin by Sato (Hillmann 1938, Sato 1936). Unfortunately the halotype was destroyed at Berlin as in the case of Parmelia elongata var. stricta. No duplicate of the type has been located in Sato's collection preserved in TNS (his earlier
collection was tranferred from TI to TNS) nor in the herbarium of the lbaraki Nature
Museum where the rest of his collection was tranferred from lbaraki University.
According to Asahina ( 1951 ), he recognized this species as a morphologicalTypification of Taxa of Hypogymnia Described rrom Japan and Sakhalin

39

variant of Parmelia enteromorpha prior to Hillmann's report in 1938 and stated"Diese Art wurde zuerst in Sakaehama (Sakhalin) gefunden und bezogen auf die
Abart von Parmelia enteromorpha". In TNS, there are two specimens collected at
Sakaehama and annotated as Parmelia fragillima by Asahina. Both of them coincide
well with the description given by Hillmann. One of them is designated as a neotype(Fig. 2a) as shown below.
The present species resembles H. stricta, because they both have elongate lobes
without soredia. However, it is readily distinguished from the latter by perforations
often formed in a row on the lower surface of each lobe.
Hypogymnia fragillima (Hillmann) Rass., Novit. Syst. PI. non Vase. 8. 1956
Parmelia fragilima Hillmann, Repert. Spec. Nov. Regni Veg. 45 : 172. 1938.
Type. Sakaehama (loc. clas.), Sakhalin, 21 July 1932, Asahina s.n. (neotype in
TNS).
Chemistry. Atranorin, 2'-0-methylphysodic acid, physodic acid and 3-hydroxyphysodic acid.

4.

Parmelia fujisanensis Asahina
When Asahina (1935) described Parmelia fujisan ensis, he cited only one specimen "Ad corticem Laricis leptolepidis in monte Fuji , Prov. Suruga (1600 metr. s.
m.)" in his protologue and showed a photograph of the type. A specimen annotated as"Typus" by Asahina is preserved in TNS and agrees with the photograph which accompanies the original description. Thus the specimen is readily regarded as theholotype of the present taxon, although the elevation " 1600 metr." is not recorded on
the label. Four other specimens determined as Parmelia fujisanensis by Asahina arealso preserved in TNS. One of them is a duplicate of the holotype. Three other specimens were not cited by Asahina in the original description, even though they arecited in "Lichens of Japan" by Asahina ( 1952). Therefore, they cannot be used as
type material.
This taxon is characterized by small adnate thalli composed of rather narrow
lobes without asexual propagules, a medulla composed of loosely interwoven hyphae ,and the presence of physodalic acid.
Hypogymnia fujisanensis (Asahina) Kurok. , Mise. Bryol. Lichenol. 5: 129.
1971 .
Parmelia fujisanensis Asah. J. Jpn. Bot. 11: 310. 1935.
Type. Subashiri-guchi 1-gome, Mt Fuji, Prov. Suruga, on Larix kaempferi, 7April 1932, Asahina I 035 (holotype and isotype in TNS).
Chemistry. Atranorin, 2'-0-methylphysodic acid, physodic acid, 3-hydroxyphysodic acid and physodalic acid.
5.

Parmelia hypotrypella Asahina
In his original description, Asahina ( 1950) reported the present species from

40

Kozo Yoshida and Hiroyuki Kashiwadania
3 3 724

Fig. 2. a) Neotype of P .fragillima (Asahina s.n. in TNS).
(Asahina 33724 in TNS). (Bar = I cm).

b) Lectotype o f P hypo!ly pellaTypification of Taxa of Hypogymnia Described !Tom Japan and Sakhalin

41

Japan , Sakhalin and Korea. Actually I 0 specimens of Parmelia hypotrypella collected from these localities before 1950 are found in TNS. One specimen annotated as"Typus" by Asahina is designated here as the lectotype (Fig. 2b).
As pointed out by Asahina (1950, 1952), the present species closely resembles
Hypogymnia hypotrypa (Nyl.) Rass ., from which it is readily distinguished by thepresence of diffuse soredia. H. hypotrypella is the only species of Hypogymnia inJapan which produces usnic acid.
Hypogymnia hypotrypella (Asahina) Rass., Bot. Materialy Otd. Spor. Rast.
13:23. 1960.
Parmelia hypotrypella Asahina, Act. Phytotax. Geobot. 14: 34. 1950.
Type. Mt. Hafu, Prov. Musashi, 24 July 1933, Asahina 33 724 (lectotype in TNS)
Chemistry. Usnic, physodalic and protocetraric acids.

6.

Parmelia metaphysodes Asahina
In the original description, Asahina (1950) reported the present species fromJapan (Hondo and Kiushu) and Sakhalin. In TNS, there are seven packets of P metaphysodes collected at these localities before 1950 and one of them was annotated as"TYPUS" by Asahina. This packet contains three fragments. In the largest one, lobes
are scarcely perforated on the lower surface and the specimen agrees very well withthe original description given by Asahina (1950). In contrast, the other two do notagree with the original description, since the thalli composed of linear elongate lobes
with distinct perforations below. Thus, the largest fragment in the packet is designated here as the lectotype (Fig. 3a).
This species has thalli composed of contiguous esorediate lobes with very few
perforations below. It is also characterized by the negative color reaction with P in the
medulla.
Hypogymnia metaphysodes (Asahina) Rass., Novit. Syst. PI. non Vase. 291.
1967.
Parmelia metaphysodes Asahina, Act. Phytotax. Geobot. 14: 33. 1950.
Type. Subashiri, Mt Fuji, Prov. Suruga, July 1924, Asahina 14 pr. p. (lectotype
in TNS).
Chemistry. Atranorin, 2' -0 -methylphysodic acid, physodic acid and 3-hydroxyphysodic acid.
7.

Parmelia nikkoensis Zahlbr.
This species was described by Zahlbruckner (1927) on the basis of a single
specimen "Asahina 15" collected from Nikko in August 1923 by Asahina. The halotype is preserved in W.
In TNS, there is a specimen annotated by Asahina as "cotype of Parmelia
nikkoensis Zahlbr." This specimen was collected by E. Ochiai on Mt Shirane, Nikko
on 11 August 1923 . Since the species is very common locally on twigs of Larix

42

Kozo Yoshida and Hiroyuki KashiwadaniaFig. 3. a) Lectotype of P metaphysodes (Asahina 14 in TNS).
(Asahina 15 in W). (Bar= I cm).

-b) Ho lotype of P nikkoensis

Typifi cation of Taxa of Hypogymnia Described from Japan and Sakhalin

43

kaempferi in Nikko , E. Ochiai , who accompanied Asahina on his trip, might havecollected a specimen on the same day at the nearby type-locality on Mt Shirane. Even
though Asahina ( 1932, 1952) mentioned that he sent a specimen to Zahlbruckner collected by E. Ochiai (Fig. 3b ), based on which P nikkoensis was described, the specimen annotated as "cotype" by Asahina cannot be regarded as a type .
The present species is characterized by I) the small shiny thalli (2- 5 cm broad)composed of narrow lobes (about I mm wide), 2) distinctly wrinkled lower surface,rarely perforated at the apices, and 3) the lack ofphysodalic acid.
Hypogymnia nikkoensis (Zahlbr.) Rass., Novit. Syst. PI. non Vase. 294. 1967.
Parmelia nikkoensis Zahlbr. , Bot. Mag. , Tokyo, 41: 364. 1927.
Type. Nikko, Prov. Shimotsuke, on twig of Larix kaempjeri, August 1923 , Y.
Asahina 15 (holotype in W).
Chemi stry. Atranorin , physodic acid and 3-hydoroxyphysodic acid .
8.

Parmelia pseudophysodes Asahina
When he described the present species, Asahina ( 1951) reported it from Japan(Hondo, Shikoku), Formosa and Sakhalin. In 1952, he cited 13 localities in theseareas and 14 specimens collected at these localities are preserved in TNS ; althoughnone of these was indicated as " Typus" by Asahina, a description of apothecia wasgiven in a protologue . Thus one of fertile specimens collected on Mt Fuji is designated here as the lectotype (Fig. 4a) as shown below.
Thi s is a distinct species characterized by glaucous to pale brown thalli , fragile
upper cortex and diffuse soredia.
Hypogymnia pseudophysodes (Asahina) Rass ., Novit. Syst. PI. non Vase. 294.
1967.
Parmelia pseudophysodes Asahina, J. Jpn. Bot. 26:100 . 1951.
Type. Ohmiyaguchi 3-gome, Mt Fuji, Prov. Suruga, 21 August 1932, Asahina s.
n. (lectotype in TNS).
Chemistry. Atranorin , physodic acid and 3-hydroxyphysodic acid.
\

9.

Parmelia pseudophysodes var. reagens Asahina
Asahina ( 1951) described var. reagens, which was identical morphologicallywith but was different chemically from typical variety. In TNS, there are three specimens determined as Parmelia pseudophysodes var. reagens by Asahina. One of themcollected on Mt Shirane, Nikko is designated as the lectotype as shown below.
The present variety was separated from the typical variety of H. pseudophysodes
only by the production of physodalic acid. However, more studies on morphologicaland chemical variations are required before its final taxonomic status can be decided.
Parmelia pseudophysodes var. reagens Asahina, J. Jpn . Bot. 26: 100. 1951.
Hypogymnia pseudophysodes var. reagens (Asahina) Kurok., Misc.Bryol.
Lichenol. 5: 130. 1971.

44

Kozo Yoshida and Hiroyuki KashiwadaniabFig. 4. a) Lectotype of P pseudophysodes (Asahina s.n. in TNS). (Bar=2cm).
P submundata f. colorans (Asahina s.n. in TNS). (Bar = I cm).

b) Holotype ofTypification of Taxa of Hypogymnia Described from Japan and Sakhalin

45

Type. Mt Shirane, Nikko, Prov. Shimotsuke, Hashimoto s.n. in 1930 (lectotype
in TNS).

10.

Parmelia submundata f. co1orans Asahina
When he described Parmelia submundata f. colorans, Asahina (1951) gave a
Latin diagnosis but did not cite any specimen. In 1952, however, he cited one specimen collected on Mt Ashibetu in Hokkaido.
In TNS , there are four packets of Parmelia submundata f. colorans collected on
Mt Ashibetu. They are apparently duplicates of a single collection and one of them isconsidered as the holotype (F ig. 4b).
Parmelia submundata f. colorans is easily recognized by the positive reaction
with P. However, the taxonomic status of this taxon will be decided when more detailed morphological and chemical variations are studied.
Parmelia submundata f. colorans Asahina, J. Jpn. Bot. 26: 99. 1951.
Hypogymnia submundata f. colorans (Asahina) Rass ., Novit. Syst. PI. non
Vase.: 295. 1967.
Type. Mt Ashibetsu , Prov. lshikari , Hokkaido, Asahina, s.n. in 1955 (holotype in
TNS).
Acknowledgments
We are deeply grateful to Or Syo Kurokawa of the Botanic Gardens of Toyama
for his kind guidance and valuable suggestions. We also wish to express our sincere
thanks to Dr Orvo Yitikainen of the Botanical Museum, University of Helsinki forhis help in consulting literature, to the Curator of the Naturhistorisches Museum
Wien (W), for the loan of the type spec imen , and to Prof.. M. R. D. Seaward, University of Bradford, for correcting the English text.

References
Asa hina, Y. 1932. Raikendokugo (45). J Jpn. Bot. 8: 164- 170. (In Japanese).
Asahina , Y. 1935. Lichenologische Notizen (V II). J Jpn. Bot. 11 : 3 10- 313.
Asahina, Y. 1950. Lichenes Japoniae Novae ve l minus Cogn itae. Acta Phy totax. Geobot. 14: 33- 35.
Asahina , Y. 1951. Lichenes Japoniae novae ve l minus cognitae (2). J Jpn . Bot. 26 : 97- 102.
Asahina , Y. 1952. Lichen s of Ja pan. 11. Genus Parmelia. 162 pp., 23 pis. Research In st. Nat. Res ., Tokyo( In Japanese).
Bitter, G. 190 I. Zur Morphologie und Systematik von Parme/ia Untergattung Hypogymnia. Hedwigia
40: 171 - 274, PI. X-XI.
Greuter, W. et al. 2000. International Code of Botanical Nomenclature (Saint Luis Code). 474 pp. Koeltz
Sci., Konigstein.
Hawksworth, D. L. 1974. Mycologist Handbook. 23 1 pp. Commonwealth Mycol. Inst. , Kew, England.
Hillmann , J. 1938. Neue oder wenig bekannte Flechten aus a ll e r Welt. Repert. Spec. Nov. Regni Veg. 45 :

171 - 177.

46

Kozo Yoshida and Hiroyuki Kashiwadani

Krog, H. 1968. The macrolichens of Alaska. Norsk Polarinst. Skr. 144: 1- 180.
Kurokawa, S. 1971. Nomenc lature of Japanese taxa of Hypogymnia and Menegazzia. Mise. Bryol.
Lichenol. 5: 129- 130.
Kurokawa, S. and S. Nakanishi 1971. Lichens of the Hidaka Mountains, Hokkaido. Mem. Natn. Sci.
Mus., Tokyo (4): 59- 70.
Lai, M.-J. 1980. Notes on some Hypogymniae (Parmeliaceae) from east Asia. Quart. Journ . Taiwan
Mus. 33 : 209- 214.
Laundon, J. R. 1979. Deceased lichenologists: their abbreviations and herbaria. Lichenologist 11 : 1- 26.
uno , M. 1964. Chemism of Parmelia subgenus Hypogymnia Nyl. J Jpn . Bot. 39: 97- 103.
Ohlsson, K. E. 1973. New and interesting macrolichens of British Columbia. Th e Bryologist 76:

366- 387.
Rassadina, K. A. 1956. Species lichenum novae et curiosae. 11 . Novit. Syst. PI. non Vase . 5- 12. ( In Russian).
Rassadina, K. A. 1960. Species Parmeliae et Hypogymniae URSS novae et curiosae. Bot. Materia/yOtd. Spo1: Rast. 13 : 20- 25 . (In Russian) .
Rassadina, K. A. 1967. Species et formae Hypogymniae novae et curiosae. Novit. Syst. PI. non Vase.
289- 300. ( In Russian).
Sa to, M. 1936. Notes on the li chen flora of Minami-Karahuto, or the Japanese Saghalien. Bull. Biogeogr.
Soc. lap. 6:97- 121.
Thomson, J. W. 1984. American arctic lichens. I . Macrolichens. 504 pp. Columbia University Press,ew York.
Zahlbruckner, A. 1927. Additamenta ad Lichenographiam Japoniae. Bot. Mag. Tokyo 41 : 313- 364.