A CONSPECTUS OF THE LICHEN GENUS STEREOCAULON ...

Journ. Hattori Bot. Lab. No. 43: 191-355 (Nov. 1977) 

A CONSPECTUS OF THE LICHEN GENUS 

STEREOCAULON (SCHREB.) HOFFM. 

Page 

INTRODUCTION ................................... .I91 

TAXONOMY.. ....................................... .I93 

SYSTEM ......................................... .I93 

CONSPECTUS SPECIERUM .................... .200 

APPENDIX 1. TYPIFICATION OF CERTAIN 

APPENDIX 2. DOUBTFUL, EXCLUDED 

AND REJECTED TAXA .................... .321 

APPENDIX 3. HERBARIA CONSULTED ..... .33l 

LITERATURE CITED ............................. .334 

SUMMARY ......................................... .346 

TAXA 

-307 INDEX 

.347 

......................................... 

............................................... 

This synoptic account is a highly condensed version of an originally planned full 

scale monograph of the genus Stereocaulon, the publication of which was not possible 

for economic reasons. It presents in abridged form the results of over 30 years' study, 

both in the field and in many herbaria, and aims at presenting the most essential data 

on the taxonomy of the species and their subsidiary taxa. The data here given include: 

the original literature references and those to extant useful descriptions of later date; 

typification of all taxa; synonymy, both obligate and facultative; exsiccata; references 

to useful illustrations; chemical constituents; general geographical distribution; 

and pertinent remarks. All type materials cited have been studied by the author 

unless otherwise stated. 

The information on chemical constituents is derived mainly from the examination 

of selected specimens by thin layer chromatography (CULBERSON and KRISTINSSON, 

1970), in some cases also by microcrystallization tests, and is to be evaluated with 

the critical caution imposed by the inherent present-day limitations of these techniques 

(cfr. J. SANTESSON, 1970), also by the limited number of samples examined. We are 

heavily indebted to several colleagues, mentioned further on, for valuable assistance 

in this respect. More detailed information on the taxonomically less important subsidiary 

constituents is to be found in the book of C. CULBERSON (1969), with its two 

supplements (C. F. CULBERSON, 1970; C. F. CULBERSON, W. L. CULBERSON and A. 

JOHNSON, 1977). 

Careful discrimination is required in using specimens in the listed exsiccata for 

comparison, as the same number in different sets may sometimes be heterogeneous, 

C/o Dr. Elke Mackenzie, Organization for Tropical Studies, Ciudad Universitaria, San JosB, 

Costa Rica.

192 Journ. Hattori Bot. Lab. No. 43 1977 

or intermixtures of other species may be present. We have mentioned such cases in 

our listing when they were detected in the limited number of sets examined. 

Stereocaulon belongs to the class Ascomycetes, subclass Ascomycetidae, order 

Lecanorales, family Stereocaulaceae, which also includes the genera Argopsis, Compsocladium 

and Pilophorus (HENSSEN and JAHNS, 1974; POELT in AHMADJIAN and HALE, 

1973). According to COOKE and HAWKSWORTH (1970) the first valid publication of the 

family Stereocaulaceae was by CHEVALLIER, 1826-27, as an "order", with the spelling 

"Stereocauleae". The characters of the family are: fruit body an apothecium; ontogeny 

gymnocarpic; developmental type ascohymenial; asci unitunicate; true paraphyses 

present; erect structure a pseudopodetium; phycobiont of thallus Chlorophyceae; 

cephalodia present (HENSSEN and JAHNS, 1974). In Stereocaulon the spores are elongateellipsoid 

to cylindrical-fusiform or vermiform, transversely multiseptate (very rarely with 

1 or 2 longitudinal septa); in Argopsis muriform or submuriform; in Compsocladium 

acicular or filiform, transversely multiseptate; and in Pilophorus simple (unicellular). 

The generic epithet Stereocaulon (SCHREB.) HOFFM. (1796) is a conserved name 

(LANJOUW et al., Int. Bot. Code, 1966, p. 247)'. It is conserved against Stereocaulon 

SCHRAD. (1794), which was based on Lichen corallinus L.,=Pertusaria corallina (L.) 

ARN., and also against Coralloides HOFFM. (1790, non WOLF, 17763), which was based on 

Lichen paschalis L. (also the type species of Stereocaulon HOFFM.). Leprocat~lon NYL., 

which has commonly been included in Stereocaulon, is a distinct and unrelated genus; 

see LAMB and WARD, 1974. 

The first published monographic treatment of the genus was by TH. FRIES (1857, 

1858). Prior to that date at least two monographic studies appear to have existed in 

manuscript, by DELISE and by LAURER; references are found in the early literature and 

on old herbarium labels to "Delise Monogr. ined." and "Laur. Monogr. ined." Later 

published studies, with the exception of those of DODGE (1929) and LAMB (1951), have 

been mostly regional, the most important being by MAGNUSSON (1926), FREY (1932), 

KERSHAW (1960), POELT (1969), DAHL and KROG (1973) for the European species, 

RIDDLE (1910) for the North American species, and YOSHIMURA (1974) for the Japanese 

species. A study of the pseudopodetial and apothecial ontogeny in several species of 

Stereocaulon was made by JAHNS (1970), who confirmed the validity of the criteria 

on which the subgenera Enteropodium (= Stereocaulon) and Holostelidium LAMB (1951) 

The first usage of the epithet Stereocaulon was by SCHREBER (1791), as Lichen sect. Stereocaulon. 

He did not mention any species, but it is probable that he included in the section species of both Stereocaulon 

and Pertusaria. On account of the fact that SCHREBER did not mention any type species, the 

question of subsequent typification of the genus arises. The first typification was that of SCHRADER, 

1794, on Lichen corallinus L. (=Pertusaria corallina (L.) ARN.), but this is now rejected. HOFFMANN, 

1796, included several species, including St. paschale (L.) HOFFM., in Stereocaulon, without citing any 

particular one of them as the type species. As far as we have been able to ascertain, the first lectotypification 

of the genus on St. paschale (L.) HOFFM. was by CLEMENTS and SHEAR on p. 320 of "The 

Genera of Fungi" (1931), where the genus is written "Stereocaulurn", and this lectotypification is approved 

by the entry in the International Code (LANJOUW et al., 1966). 

The lectotype of Coralloides WOLF (1776) is Lichen subulatus L. (=CIadonia subulata (L.) WIGG.); 

see Ross, 1966.

I. MACKENZIE LAMB: A Conspectus of Stereocaulon 193 

were established. 

The system of subgeneric classification here presented (p. 195 et sqq.) is based on 

that originally proposed by the present author (LAMB, 1951) with alterations derived 

from subsequent further study, and also incorporating certain features of the scheme 

elaborated by DUVIGNEAUD in his excellent treatment of the Central African species 

(1956). Previous subgeneric classifications are discussed in LAMB (1951). 

The number of species of Stereocaulon here recognized is 123 (of which 19 new), 

with 47 varieties (of which 14 new) and 40 forms (of which 11 new). For further 

details, see the Summary (p. 346). 

We have here used the traditional terms excipulum, central cone, and hypothecium 

in describing apothecial structure; it may however be preferable in future to use the 

alternative termspseudoexcipulum, medullary tissue, and subhymenial layer (A. HENSSEN, 

personal commun.). 

Reagents mentioned in the text are potassium hydroxide (KHO, abbreviated K), 

a 5-25 % aqueous solution, and para-phenylenediamine (H,NCeH,NH,, abbreviated 

PD), a freshly prepared 0.1 % alcoholic solution. They are applied to the phyllocladia 

or to young pseudopodetial tips; in some cases it may be necessary to scrape away 

the cortex and apply the reagent to the exposed medullary tissue. 

Our work was financially supported by two grants from the National Science 

Foundation (GB-4503, GA-119) and by smaller grants from the American Academy 

of Arts and Sciences, the Harvard Foundation for Advanced Study and Research 

(including the Joseph H. Clark Bequest Fund) and the William F. Milton Fund of 

Harvard University. Grateful thanks are also due to many colleagues who assisted us 

in other ways; in particular we would like to acknowledge the help received from Drs. 

C. CULBERSON, C. FOX, M. HALE and S. HUNECK in the elucidation of chemical constituents, 

Dr. G. SAYRE in matters of bibliography, Dr. M. ZIMMERMANN in preparing 

microtome sections and photomicrographs, and Dr. V. AHMADJIAN for contributing 

an instructive series of specimens collected by him in Swedish Lapland in 1959. 

It is planned to publish a supplementary key to all the recognized species at a 

later date. 

TAXONOMY 

SYSTEM^ 

The main departures from the scheme presented in LAMB, 1951, are the following: 

Subgen. Enteropodium sect. Chondrocaulon TH. FR., comprising the species of 

the imperfect genus Leprocaulon NYL., has been eliminated, since Leprocaulon is a 

distinct and unrelated genus (LAMB and WARD, 1974). 

Subgen. Enteropodium LAMB is now called subgen. Stereocaulon, as it contains 

the type species of the genus, and for the same reason sect. Eustereocaulon KORB. 

is now called sect. Stereocaulon. 

Supplementary notes on pp.198-199.


Sect. Denudata JATTA now replaces sect. Leucocheilon LAMB, being an earlier 

epithet in the same rank, and sect. Leucocheilon subsect. Peltophyllum (Du RIETZ) 

LAMB is now called subsect. Denudata, as it contains the type species of the section. 

Sect. Botryoideum G. T. JOHNS. em. LAMB is transferred, as a subsection, from 

subgen. Holostelidium to subgen. Stereocaulon, as the development has proven to be 

enteropodious; a connection with the vesuvianum-group (sect. Denudata JATTA) is 

plainly demonstrated by two of its species, St. saviczii and St. hokkaidense. These 

two species seem to be the most primitive of the subsect. Botryoideum, which comprises 

the majority of the Japanese species, and they seem to indicate that this assemblage 

originated in the subarctic zone of the northern Pacific region and spread southwards 

into Japan, there differentiating into a number of highly specialized species. The same 

origin is postulated by KUROKAWA (1972) for Parmelia galbina (L.) AcH., which is 

believed to have migrated southwards not only into Japan, but also into the southeastern 

United States. Additional evidence for the new taxonomic disposition is 

the fact that in St. japonicum TH. FR. the youngest phyllocladia in the emergent stage 

have a button-like form, suggesting that the coralloid phyllocladia of the advanced 

species of subsect. Botryoideum are modifications of a basically squamulose or peltate 

type. 

DUVIGNEAUD'S subgenus Lobophoron is incorporated into the system, as a section 

of subgen. Stereocaulon. It shows that the development of protosacculate cephalodia 

is not restricted to the subgen. Holostelidium. 

Sect. Pilophoropsis LAMB (non DUVIGN.) is now given the rank of a third subgenus 

of Stereocaulon, representing apparently a connecting link between the subgenera 

Stereocaulon and Holostelidium. Its development appears to be of the holostelideous 

type, but its cephalodia are truly botryose, corresponding to those produced by the 

species of subgen. Stereocaulon and not found in subgen. Holostelidium. 

In subgen. Holostelidium, sect. Sacculata SAT6 is replaced by sect. Holostelidium, 

as it contains the type species of the subgenus, and for the same reason subsect. Ascaridisporae 

DU RIETZ ex LAMB is now called subsect. Holostelidium. Subsect. Holostelidium 

comprises species with protosacculate as well as those with sacculate cephalodia. 

St. piluliferum TH. FR. and St. pomiferum DUVIGN., for instance, are closely related 

species, differing mainly in the cephalodial anatomy (sacculate and loose-cored in the 

former, protosacculate and solid-cored in the latter). 

Subgen. Holostelidium sect. Ecephalodiata LAMB is now placed in synonymy with 

subgen. Stereocaulon sect. Denudata subsect. Denudata, as the two species involved, 

St. scutelligerum TH. FR. and St. virgatum AcH., are clearly referable here, being 

connected with species of the more typical vesuvianum-type by intermediates such as 

St. anomalum LAMB, St. jibrillosum LAMB and St. furfuraceum DUVIGN. Cephalodia 

have since been found in one of the species (St. virgatum), and their presence or absence 

is not a very important taxonomic criterion.


STEREOCAULON (SCHREB.) HOFFM. 

Deutschlands Flora oder Botanisches Taschenbuch, Zweyter Theil: 128 (1796) (nom. conserv.). - 

Coralloides HOFFM. Descriptio et Adumbratio Plantarum, 1 : 23, 25 (1790) (non Coralloides WOLF ex 

anno 1776). 

Subgen. I. Stereocaulon 

Syn. Stereocaulon subgen. Eustereocaulon JATTA, Bull. SOC. Bot. Ital. 1892: 436 (1892). - Stereocaulon 

subgen. Enteropodium LAMB, Canad. J. Bot. 29: 566 (1951). - Stereocaulon subgen. Platycladium 

DUVIGN. Lejeunia, MCm. no. 14: 41, 43 (see Note I). - Stereocaulon subgen. Pilophoropsis DUVIGN. 

Lejeunia, MCm. no. 14: 42, 44 (non LAMB; see Note 2). 

Pseudopodetia produced by the basal medullary stratum only of the primary thallus; 

true phyllocladia present, similar to those forming the primary thallus (except in sect. Stereocladium). 

Type species: St. paschale (L.) HOFFM. 

Sect. 1. Stereocaulon 

Syn. Stereocaulon sect. Eustereocaulon K&B. Systema Lichenum Germaniae: 13 (1855); LAMB, Canad. 

J. Bot. 29: 566 (1951). - Stereocaulon subgen. Eustereocaulon sect. Paschalia JATTA, Flora Italica 

Cryptogama, Pars 111, Fasc. IV: 463 (1911) (see Note 3). - Stereocaulon subgen. Platycladium ser. 

Granulata DWIGN. Lejeunia, MCm. no. 14: 43 (1956); Stereocaulon subgen. Enteropodium sect. Eustereocaulon 

ser. Granulata DUVIGN. Lejeunia, Mem. no. 14: 49, 54, 56 (see Note 4). 

Primary thallus persistent or evanescent, composed of squamules or verrucae similar to 

the phyllocladia of the pseudopodetia; phyllocladia without darker centers. Cephalodia 

simple to botryose, with poorly developed cortical layer. Type species, St. paschale (L.) 

HOFFM. Other species: St. alpinum LAUR., St. antarcticum VAIN., St. austroindicum LAMB, 

St. azoreum (SCHAER.) NYL., St. botryosum ACH. em. FREY, St. capitellatum MAGN., St. 

condensatum HOFFM., St. cymosum CROMB., St. dactylophyllum FLORKE (see Note 5), St. depreaultii 

DEL. ex NYL., St. depressum (FREY) LAMB, St. esterhuysenae LAMB (see Note 6), 

St. evolutum GRAEWE, St. glabrum (MULL. ARG.) VAIN., St. glareosum (SAV.) MAGN., St. 

glaucescens TUCK., St. grande (MAGN.) MAGN., St. groenlandicum (DAHL) LAMB, St. incrustatum 

FLORKE, St. intermedium (SAV.) MAGN., St. leprocauloides LAMB, St. myriocarpum 

TH. FR., St. papuanum LAMB, St. pileaturn AcH., St. pseudodepreaultii ASAH., St. rivulorum 

MAGN., St. sasakii ZAHLBR., St. saxatile MAGN., St. sibiricum LAMB, St. spathuliferum VAIN., 

St. sterile (SAV.) LAMB, St. subcoralloides (NYL.) NYL., St. supervestiens MAGN. (see Note 7), 

St. tennesseense MAGN., St. tomentosum FR., St. uliginosum LAMB. 

Sect. 2. Stereocladium (NYL.) DODGE 

Ann. Cryptog. Exot. 2 (2): 95 (1929); emend. LAMB, Canad. J. Bot. 29: 566 (1951). - Stereocladium 

NYL. Flora, 58: 302 (1875). 

Primary thallus persistent, of erect, flabellate-spathulate squamules; phyllocladia of 

pseudopodetia granulose to verrucose, without darker centers. Cephalodia simple, scabridtuberculate, 

with poorly developed cortical layer. Type and only known species, St. nanodes 

TUCK. (syn. St. tyroliense (NYL.) LETT.). 

Sect. 3. Denudata JATTA 

Flora Italica Cryptogama, Pars 111, Fasc. IV: 463 (1911) (see Note 8). - Stereocaulon subgen. Enteropodium 

sect. Leucocheilon LAMB, Canad. J. Bot. 29: 566 (1951).


Primary thallus persistent or evanescent, of squamules similar to the phyllocladia of the 

pseudopodetia; phyllocladia squamulose, peltate, verrucose, dactylaeform to cylindric- 

coralloid, flabellate or flattened-foliose, usually with darker centers and paler margins. 

Cephalodia simple to botryose, with poorly developed cortical layer; in some species absent. 

Type species, St. vesuvianum PERS. (lectotypus, lectotypif. nov.; see Note 8). 

Subsect. A. Denudata 

Syn. Stereocaulon subgen. Enteropodium sect. Leucocheilon subsect. Peltophyllum @U RIETZ) LAMB, 

Canad. J. Bot. 29: 567 (1951). - Stereocaulon subgen. Platycladium ser. Umbilicata DUVIGN. Lejeunia, 

MBm. no. 14: 44 (1956). - Stereocaulon subgen. Enteropodium sect. Eustereocaulon ser. Umbilicata 

DUVIGN. Lejeunia, Mem. no. 14: 49, 55, 79 (1956) (see Note 9). - Lecanocaulon NYL. Synopsis Methodica 

Lichenum, 1 (2): 233 (1860) (as "fere genus proprium"), emend. LAMB (emend. nov.; see Note 

10). - Stereocaulon subgen. Lecanocaulon (NYL.) VAIN. Etude sur la Classification et la Morphologie 

des Lichens du Bresil, Pars prima: 61 (1890) (see Note 10). - Stereocaulon subgen. Cylindrocladium 

sect. Lecanocaulon (NYL.) DUVIGN. Lejeunia, Mem. no. 14: 44 (1956) (sensu erroneo; see Note 11). 

- Stereocaulon subgen. Holostelidium sect. Ecephalodiata LAMB, Canad. J. Bot. 29: 565 (1951) (see 

Note 12). - Chlorocaulum CLEMENTS, The Genera of Fungi: 175 (1909) (see Note 24). 

Phyllocladia lateral, sessile, verrucose to peltate, coralloid, or foliose. Apothecia 

usually lateral and small. Type species, St. vesuvianum PERS. Other species: St. anomalum 

LAMB, St. arcticum LYNGE, St. arenarium (SAV.) LAMB, St. cornutum MULL. ARG., St. fibrillosum 

LAMB, St. furfuraceurn DUVIGN., St. graminosum SCHAER., St. lavicola MAGN. (see Note 

13), St. leprocephalum VAIN., St. nigromaculatum DUVIGN., St. obesum TH. FR., St. pityrizans 

NYL., St. scutelligerum TH. FR., St. subdenudatum HAV., St. symphycheilum LAMB, St. vanoyei 

DUVIGN., St. verruciferum NYL. (see Note 14), St. virgatum AcH., St. vulcani (BORY) AcH., 

St. weberi LAMB. 

Subsect. B. Phyllocaulon (TUCK.) LAMB 

Canad. J. Bot. 29: 567 (1951). - Stereocaulon subgen. Phyllocaulon TUCK. Genera Lichenum: 144 

(1872). - Stereocaulon sect. Phyllocaulon (TUCK.) RAS. Acta Bot. Fenn. 33: 10 (1943). - Phyllocaulon 

(TUCK.) VAIN. Ark. Bot. 8 (4): 36 (1909). 

Phyllocladia apical, flattened, spathulate. Apothecia lateral. Type species, St. wrightii 

TUCK. Other species: St. apocalypticum NYL. 

Subsect. C. Botryoideum (G. T. JOHNS. emend. LAMB) LAMB (n. stat.) 

Syn. Stereocaulon sect. Botryoideum G. T. JOHNS. Bryologist, 44: 7 (1941); emend. LAMB, Canad. J. 

Bot. 29: 565 (1951). - Stereocaulon subgen. Lecidocaulon sect. Verruculata SAT^, Cladoniales (I), in 

NAKAI et HONDA, Nova Flora Japonica: 57, footnote (1941) (see Note 15). 

Phyllocladia lateral, usually cylindric-coralloid and without darker spots or centers, 

rarely verrucose with darker spots or centers, never flattened-foliose, perhaps rarely deliquescent 

into soredia (cfr. St. exile ASAH.). Apothecia terminal. Type species, St. exutum 

NYL. Other species: St. commixtum (ASAH.) ASAH., St. curtatoides ASAH., St. curtatum 

NYL., St. etigoense (ASAH.) LAMB, St. exile ASAH. (see Note 16), St. hokkaidense ASAH. et 

LAMB, St. japonicum TH. FR., St. nigrum HUE, St. octomerellum MULL. ARG., St. octomerum 

MULL. ARG., St. pendulum ASAH., St. prostratum ZAHLBR., St. saviczii Du RIETZ, St. verruculigerum 

HUE.

I. MACKENZIE LAMB: A Conspectus of Stereocaulon 

Sect. 4. Lobophoron (DUVIGN.) LAMB (n. stat.) 

Syn. Stereocaulon subgen. Lobophoron DwrGN. Lejeunia, Mern. no. 14: 44,49, 131 (1956). 

Phyllocladia lateral, verrucose, coralloid, or foliose, or completely dissolved into soredia. 

Cephalodia protosacculate (solid-cored, with well developed gelatinized cortical layer). 

Type species, St. foliolosum NYL. Other species: St. coniophyllum LAMB, St. himalayense 

AWAS. et LAMB, St. humbertii DUVIGN. 

Subgen. 11. Pilophoropsis (LAMB) LAMB (n. stat.) 

Syn. Stereocaulon subgen. Holostelidiurn sect. Pilophoropsis LAMB, Canad. J. Bot. 29: 565 (1951) (non 

Stereocaulon subgen. Pilophoropsis DUVIGN. ex anno 1956, norn. inval. ; see Note 2). 

Development (probably) holostelideous (see Note 17). Both phyllocladial branchlets 

(harmonic) and true phyllocladia (disharmonic) present, the latter derived from the pseudopodetial 

thallus-mantle, verrucose to digitate-coralloid. Cephalodia botryose, with pseudoparenchymatous, 

non-gelatinized cortical layer. Type and only known species, St. curtum 

(RKs.) LAMB. 

Subgen. 111. Holostelidiurn LAMB. 

Canad. J. Bot. 29: 564 (1951). - Stereocaulon subgen. Cylindrocladium DUVIGN. Lejeunia, Mern. no. 

14: 44 (see Note 18). 

Pseudopodetia produced by all layers of the primary thallus; phyllocladial branchlets 

usually present; no true phyllocladia. Cephalodia sacculate or protosacculate, with well 

developed gelatinized cortical layer. Type species, St. sorediiferum HUE. 

Sect. 1. Holostelidiurn 

Syn. Stereocaulon sect. Sacculata SAT^, Cladoniales (I), in NAKAI et HONDA, Nova Flora Japonica: 

82, footnote (1941); LAMB, Canad. J. Bot. 29: 565 (1951). 

Pseudopodetia usually well branched and with numerous and well developed phyllocladioid 

branchlets. Cephalodia sacculate or protosacculate. Type species, St. sorediiferum 

HUE. 

Subsect. A. Holostelidiurn 

Syn. Stereocaulon subgen. Holostelidiurn sect. Sacculata subsect. Ascaridisporae Du RIETZ ex LAMB, 

Canad. J. Bot. 29: 565 (1951). - Stereocaulon subgen. Cylindrocladium sect. Rhopalophoron DUVIGN. 

Lejeunia, MBrn. no. 14: 44 (1956). - Stereocaulon subgen. Rhopalophoron Duvrc~. op. cit. 49, 106, 

with series Protoclavulifera DWIGN. and Euclavulifera DUVIGN. (see Note 19). 

Spores vermiform, (6) 8-30-septate. Apothecia hypophysate in the young condition. 

Cephalodia sacculate or protosacculate, smoothly globose to commonly indented-scrobiculate, 

with uniform cortex. Hypothecium colorless. Type species, St. sorediiferum HUE. 

Other species: St. brassii LAMB, St. claviceps TH. FR., St. macrocephalum MULL. ARG., St. 

massartianum HUE, St. montagneanum LAMB, St. piluliferum TH. FR., St. pomiferum DUVIGN., 

St. procerum GROENH., St. paradoxum LAMB (see Note 20), St. togashii LAMB. 

Subsect. B. Aciculisporae Du RIETZ 

ex LAMB, Canad. J. Bot. 29: 565 (1951). - Stereocaulon subgen. Lecanocaulon sensu DWIGN. Lejeunia, 

Mern. no. 14: 49 (1956) (non Nn., nec VAIN.; see Note 11). - Corynophoron NYL. Synopsis Methodica


Lichenum, 1 (2) : 232 (1 860), as "fere genus proprium" ; Lichenes Novae Zelandiae : 15 (1 888). - Stereocaulon 

subgen. Cylindrocladium sect. Corynophoron (NYL.) DUVIGN. Lejeunia, Mem. no. 14: 44 (1956) 

(see Note 21). 

Spores fusiform-cylindric, 3-7-septate (rarely up to 16-septate). Apothecia not hypophysate. 

Cephalodia sacculate or protosacculate, smoothly globose to commonly indentedscrobiculate 

or rarely cristate, with uniform or dimorphous cortex. Type species, St. ramulosum 

(Sw.) RAUSCH. Other species: St. argus HOOK. F. et TAYL. em. TH. FR., St. atlanticum 

(LAMB) LAMB, St. colensoi BAB., St. corticatulum NYL., St. crambidiocephalum LAMB, St. 

delisei BORY, St. didymicum LAMB, St. dusenii LAMB, St. fronduliferum LAMB, St. halei LAMB, 

St. implexum TH. FR., St. melanopotamicum LAMB, St. meyeri B. STEIN, St. microcarpurn 

M~~LL. ARG., St. novogranatense LAMB, St. pachycephalum VAIN., St. philippinense RAS., 

St. pseudomassartianum LAMB, St. rubiginosum PERS., St. rugulosum LAMB, St. salazinum 

(BORY) F ~E, St. staufferi LAMB, St. stenospermum (LAMB) LAMB, St. trachyphloeum LAMB, 

St. wadei LAMB. 

Subsect. C. Dactyloideum (LAMB) LAMB (n. stat.) 

Syn. Stereocaulon subgen. Holostelidium sect. Dactyloideum LAMB, Canad. J. Bot. 29: 565 (1951). 

Spores vermiform, (7-) 8-13-septate. Apothecia not hypophysate. Cephalodia protosacculate, 

finally digitately divided (dactylae form), with uniform cortex. Hypothecium 

pigmented (dark brown). Type and only known species, St. strictum TH. FR. (see Note 22). 

Sect. 2. Redingeria LAMB 

Canad. J. Bot. 29: 565 (1951). - Gymnocaulon DUVIGN. Lejeunia, Mem. no. 14: 14, 41, 46, 50 (1956) 

(see Note 23). 

Pseudopodetia simple or sparingly branched; no phyllocladioid branchlets developed; 

cephalodia sacculate (sometimes indistinctly so). Type species, St. caespitosum RED. Other 

species: St. gregarium RED., St. loricatum LAMB. 

Supplementary Notes 

1. Stereocaulon subgen. Platycladium DUVIGN. Nom. inval., no Latin diagnosis given. 

No type species designated by DUVIGNEAUD. 

2. Stereocaulon subgen. Pilophoropsis DUVIGN. (non Stereocaulon subgen. Pilophoropsis 

(LAMB) LAMB). Nom. inval., no Latin diagnosis given. St. sphaerophoroides TUCK. 

(in the sense of St. azoreum (SCHAER.) NYL.) designated by DUVIGNEAUD as type species 

of his subgen. Pilophoropsis. 

3. Stereocaulon subgen. Eustereocaulon sect. Paschalia JATTA. Sectional name spelt 

with small initial letter by JATTA. NO type species designated, but St. paschaale (L.) 

HOFFM. listed among other species. 

4. Stereocaulon subgen. Enteropodium sect. Eustereocaulon ser. Granulata DUVIGN. St. 

paschale (L.) HOFFM. designated by DUVIGNEAUD as type species of ser. Granulata. 

5. St. dactylophyllum FLORKE. Position in this section somewhat doubtful; it may possibly 

belong to sect. Denudata subsect. Botryoideum, although this would seem unlikely 

for phytogeographical reasons. 

6. St. esterhuysenae LAMB. Position in this section somewhat doubtful. 

7. St. supervestiens MAGN. Position in this section doubtful.


Stereocaulon subgen. Eustereocaulon sect. Denudata JATTA. Sectional name spelt with 

small initial letter by JATTA. NO type species designated by JATTA, but St. vesuvianum 

PERS. listed among other species. We here lectotypify it on St. vesuvianum (lectotypif. 

nov.). 

Stereocaulon subgen. Enteropodium sect. Eustereocaulon ser. Umbilicata DUVIGN. St. 

denudatum FLORKE (syn. St. vesuvianum var. nodulosum (WALLR.) LAMB) designated by 

DUVIGNEAUD as type species. 

Lecanocaulon NYL. Now lectotypified by us on St. scutelligerum TH. FR. ("Stereocaulon 

salazinum" sensu NYL.), lectotypif. nov. NYLANDER, loc. cit., cited the following 

3 species under Lecanocaulon, without designating a type: St. Iecanoreum NYL. (=St. 

strictum TH. FR.), St. salazinum (BORY) (in the sense of St. scutelligerum TH. FR., as 

plainly shown by NYLANDER'S figure), and St. pityrizans NYL. VAINIO, in making the 

combination Stereocaulon subgen. Lecanocaulon (NYL.), did not mention any species. 

Stereocaulon subgen. Cylindrocladium sect. Lecanocaulon (NYL.) DUVIGN. Comb. inval., 

no basionym indicated. Wrongly typified by DUVIGNEAUD on St. ramulosum (SW.) 

RAUSCH., a species not included in the protologue of Lecanocaulon. 

Stereocaulon subgen. Holostelidium sect. Ecephalodiata LAMB. St. virgatum ACH. 

designated by LAMB as type species of sect. Ecephalodiata. 

St. lavicola MAGN. Position in this section doubtful. 

St. verruciferum NYL. Syn. St. speciosum LAMB. 

Stereocaulon subgen. Lecidocaulon sect. Verruculata SAT6 St. exutum NYL. designated 

by SAT^ as type species of sect. Verruculata. Published 3 months later than sect. 

Botryoideum G. T. JOHNS. 

St. exile ASAH. Position in this subsection doubtful. 

Subgen. Pilophoropsis (LAMB) LAMB. None of the material of the only species, St. 

curtum (Rxs.) LAMB, seen to date shows the earliest stages of development necessary 

to determine conclusively whether the origin of the pseudopodetia is holostelideous or 

enteropodious, but the former seems more probable. 

Stereocaulon subgen. Cylindrocladium DUVIGN. Nom. inval., no Latin diagnosis 

given. No type species designated by DUVIGNEAUD. 

Stereocaulon subgen. Rhopalophoron DUVIGN. St. sorediiferum HUE designated by 

DUVIGNEAUD as type species of subgen. Rhopalophoron. St. nesaeum NYL. (in the sense 

of St. massartianum HUE) designated as type species of ser. Protoclavulifera, and St. 

piluliferum TH. FR. as type species of ser. Euclavulifera. There is an error here, because 

one of the two series would have to be based on St. sorediiferum HUE and called 

ser. Rhopalophoron. 

St. paradoxum LAMB is doubtfully placed in subsect. Holostelidium pending discovery 

of fertile specimens. 

Stereocaulon subgen. Cylindrocladium sect. Corynophoron (NYL.) DUVIGN. Comb. 

inval., no basionym indicated. 

St. strictum TH. FR. Syn. St. explanaturn LAMB, St. lecanoreum NYL., St. peladense 

VAIN., St. substrictum HUE. 

Gymnocaulon DUVIGN. Nom. inval., no Latin diagnosis supplied. 

Chlorocaulum CLEMENTS. Based on Lichen salazinus BORY (in the sense of St. scutelligerum 

TH. FR.).

Journ. Hattori Bot. Lab. No. 43 

CONSPECTUS SPECIERUM 

Stereocaulon (SCHREB.) HOFFM. 

Deutschlands Flora oder Botanisches Taschenbuch, Zweyter Theil, Cryptogamie: 128 (1796) (nom. 

conserv.; vide LANJOUW et al.: 247; 1966). -Lichen sect. Stereocaulon SCHREB. 

Genera Plantarum. . . Editio octava, 2: 768 (1791). 

Subgen. I. Stereocaulon 

Sect. 1. Stereocaulon 

1. Stereocaulon paschale (L.) HOFFM. 

Caroli a Linne . . . 

Deutschlands Flora oder Botanisches Taschenbuch, Zweyter Theil, Cryptogamie: 130 (1796); MAGN. 

Goteborgs K. Vetensk.- o. Vitterh.-Samh. Handl. ser. 4, 30 (7): 45 (1926); FREY, Rabenhorsts Krypt.- 

FI. 9, Abt. IV, 1. Halfte: 147 (1932); DAHL et KROG, Macrolichens of Denmark, Finland, Norway and 

Sweden: 140 (1973). - Lichen paschalis L. Species Plantarum 2: 1153 (1753) (basionym). - Cladonia 

paschalis (L.) WIGG. Primitiae Florae Holsaticae: 90 (1780). - Coralloides paschale (L.) HOFFM. 

Descriptio et Adumbratio Plantarum 1 : 23 (1789). - Verrucaria paschalis (L.) HUMB, Florae Fribergensis 

Specimen: 43 (1793). - Baeomyces paschalis (L.) WAHLENB. Flora Lapponica: 450 (1812). - 

Patellaria paschalis (L.) WALLR. Flora Cryptogamica Germaniae, Pars prior: 441 (1831). -Stereocaulomyces 

paschalis (L.) THOMAS, Beitr. Kryptogamenfl. Schweiz, 9 (1): 180 (1939) (nom. nud.), ex 

CIF. et TOMAS. Atti 1st. Bot. Lab. Crittog. Univ. Pavia, ser. 5,lO: 272 (1954) (cum des~ript.)~. -Stereocaulon 

paschale [f.] a. paschale (L.) MUDD, A Manual of British Lichens: 65 (1861). 

Typus: country of origin not indicated ("Helvetiae, Italiae, Cambriae, Lapponiae, Scaniae, Groenlandiae, 

Pensylvaniae alpestribus"), most probably from Swedish Lapland; in herb. L. (LINN), sheet 

1273-259, lowermost specimen (lectotype, lectotypit. nov. ; see p. 308). 

Facultative synonyms: Stereocaulon paschale f. typicum Du RIETZ, Svensk Bot. Tidskr. 20: 96 

(1926). - Stereocaulonpaschale [var.] a. vulgare TH. FR. De Stereoc. et Pilophor. Comment.: 33 (1957). 

- Stereocaulon paschale [f.] a. vulgare (TH. FR.) TH. FR. Lichenographia Scandinavica, Pars prima: 

47 (1871). - Stereocaulon paschale [var.] 6. serpens TH. FR. De Stereoc. et Pilophor. Comment.: 

33 (1857). - Stereocaulon paschale [f.] 6. serpens (TH. FR.) TH. FR. Lichenographia Scandinavica, 

Pars prima: 47 (1871). - Stereocaulon paschale f. colligatum LAMB, Ann. Rep. Nat. Mus. Canada, 

1952 - 53, Bull. no. 132: 273 (1954). - Stereocaulonpaschale f. congesturn LAMB ex LEPAGE, Naturaliste 

Canad. 85: 184 (1958) (nom. nud.). - (?) Lichen crustaceus GUIB. Exercitia Phytologica, 2: 604 (1792) 

(ut synon. Stereocaulipaschalis; dubium). 

Exsicc.: ARN. Lich. Exs. no. 1363b (1888), as St. alpinum (c, FH, L, PC. The example in FH Contains 

also St. alpinum LAUR.). - BARTLING et HAMPE, Veget. Cellul. German. Septentr., Lich., Decas II 

(date ?) no. 17 pr. p. (M. This example is partly St. incrustaturn FLK. The numbers on the labels of 

this exsiccat do not agree with those cited by LYNGE, 1915 - 20). - BREUT. Flora German. Exs. ser. II, 

Cent. I1 (1843) no. 107, as St. paschale var. thyrsoideum (FH, PC, UPS. An example in s is St. alpinum 

LAUR., and one in L is a mixture of St. paschale and St. alpinum). - BROCKMWLLER, Mecklenburg. 

Kryptog. Fasc. II(1862) no. 67 (MB). - CUMM., WILLIAMS et SEYMOUR, Decades N. Amer. Lich. Decade 

3 (1892 ?) no. 25, pr. min. p. (c, CAN. The example in c is mixed with St. tomentosum FR. Those in 

DUKE, OC, US, WELC and wIs are St. saxatile MAGN.). - EHRH. PI. Cryptog. Linn. Decas XI1 (1789) no. 

118, as Lichen paschalis (ups). - ELENK. Lich. Flor. Ross. Fasc. II (1907) no. 72a, b (FH). - FLORKE, 

Deutsch. Lich. Fasc. X (1821) no. 199 (MSC. Acc. to FREY, 1932, p. 149, an example seen by him is partly 

St. tomentosum FR.). - FLOT. Deutsche Lich. Exs. no. 12A (UPS; lectotype specimen of St. paschale 

[var.] a. vulgare TH, FR.~). - FR. Lich. Suec. Exs. Fasc. III (1824) no. 89 pr. p., as St. paschale B. conglo- 

Refers to the mycobiont artificially isolated from the alga. 

Lectotypif. nov.; see p. 308.

I. MACKENZIE LAMB: A Conspectus of Stereocaulon 20 1 

meratum (s. Examples of this number seen in FH, LD and ups are St. evolutum GRAEWE or mixtures of 

St. paschale and St. evolutum). - HALE, Lich. Amer. Exs. Fasc. IV (1970) no. 98, as St. dactylophyllum 

(FH, LD). - HAV. Lich. EXS. Norveg. Fasc. VII (1905) no. 262 (BG); Lich. Norveg. Occid. Exs. Fasc. 

VII (1938) no. 153 (PR. This example has some intermixture of St. grande (MAGN.) MAGN.). - HEPP, 

Flechten Europas Band VZ (1857) no. 304, as St. paschale B. conglomeratum (FH, LD, PC, s.). - Krypt. 

Exs. Vindob. Cent. XL (1957) no. 3918 pr. p. (co~o, FH, L, LD. The examples in FH and L contain also 

St. alpinum LAUR. var. erectum FREY). - KUROK. Lich. Rarior. et Crit. Exs. Fasc. I11 (1971) no. 145, 

as St. sasakii (FH, LD). - Lich. Canad. Exs. no. 66 (1970), no. 67 (1970) (CAN). - Lichenotheca Fenn. 

Fasc. V (1947) no. 103, as St. squamescens, pr. p. (CAN, UPS. The example in LD is St. saxatile MAGN.); 

Fasc. VII (1947) no. 171 pr. p. (FH. This example is partly St. dactylophyllum FLK.); Fasc. X (1947) 

no. 248 (FH, HAK); Fasc. XXXI (1953) no. 760 (FH); Fasc. XXXIX (1956) no. 961 (FH); Fasc. XLIII 

(1957) no. 1056 (FH, with parasymbiontic Scutula); Fasc. XLIV (1958) no. 1081 (FH, with parasymbiontic 

Scutula); Fasc. L (1959) no. 1230, as St. subcoralloides (FH). - MAC. Canad. Lich. ser. I (1893) no. 

44, as St. tomentosum (WELC); ser. I1 (1909 - 12) no. 133 (FH, MSC, US. An example in WELC is St. saxatile 

MAGN., and one in DUKE is St. tomentosum FR.); no. 134, as St. tomentosum, pr. p. (us. The material 

distributed in this number was collected from four different Canadian localities. An example in 

DUKE is St. tomentosum FR., and one in WELC is St. saxatile MAGN.); no. 258 (us); no. 259, as St. tomentosum 

(us). - MAGN. Lich. Sel. Scand. Exs. Fasc. VII (1933) no. 157 (DEGEL, PR). - MALME, Lich. 

Suec. Exs. Fasc. XXI (1915) no. 509A, B (PR, WIS). - MERESCH. Lich. ROSS. EXS. Fasc. I1 (1913) no. 39 

(FH). - MIG. Cryptog. German. Austr. et Helvet. Exs. Fasc. XXV (1904) no. 123 (FH, MSC, OC, PR, u). - 

NORRL. et NYL. Herb. Lich. Fenn. Continuatio (1921) no. 501 ("pascale") (ups). - RABENH. Lich. 

Europ. Exs. Fasc. V (1856) no. 134 (PR). - RAS. Lich. Fenn. Exs. Fasc. I1 (1935) no. 57, as St. paschale 

var. squamescens (GB); Fasc. IV (1936) no. 197 (ups); no. 198, as St.paschale var. grandis (ups). - SAV. 

Lichenotheca Ross. Decas VIII (1956) no. 74, as St. alpinum (LD). - STENHAM. Lich. Suec. Exs. Fasc. 

I11 (1860) no. 80 (FH, LD, us); no. 81 pr. p., as St. paschale b. gracilentum (LD. In this example it occurs 

intermixed with St. alpinum LAUR. f. gracilentum (TH. FR.) LAMB; in that in FH it is entirely the latter). - 

TOBOL. Lichenotheca Polon. Fasc. VI (1955) no. 113 (FH. An example seen in LD is St. saxatile MAGN.). 

- TUCK. Lich. Amer. Septentr. Exs. Fasc. V (1 854) no. 112 (FH, UPS. An example seen in TENN, and one 

of two in FH, copiously fertile and without cephalodia, are tentatively referred to St. subintricans NYL., 

a doubtful species; see p. 324). - V~ZDA, Lich. Sel. Exs. Fasc. XLIX (1974) no. 1211 (FH). 

Icon.: GALLBE 1972, P1. 74, figs. 566 - 572; PI. 75, figs. 573 - 578; P1. 76, figs. 579 - 583; P1. 77, 

figs. 584 - 587. - FREY 1932, fig. 23 ; PI. 111, figs. 1, 2. - RAs. 1951, PI. XVI, fig. 1. - YOSHIM. 1974, 

figs. 74a, 77d; PI. 33, fig. 331. 

Mat. chim.: atranorin and lobaric acid; occasionally occurring in a deficient phase with 

only atranorin demonstrable by present chromatographic techniques. 

Distrib. Sweden, Norway, Denmark, Finland, Netherlands (probably now extinct), Scotland, 

W. Germany, E. Germany, Poland, U. S. S. R. (including Estonia and Karelia), Mongolian People's 

Republic, Iceland, Greenland, Canada (N. W. Temt., Yukon Territ., British Columbia, Alberta, 

Saskatchewan, Manitoba, Ontario, Quebec, Labrador, Newfoundland, Nova Scotia), Miquelon 

Island, U. S. A. (Maine, New Hampshire, Massachusetts, where probably now extinct, Virginia, Michigan, 

Montana, Alaska), Manchuria, Korea, Japan (Hokkaido, Honshu), Chile (Valdivia, Magallanes), 

Argentina (Chubut). Unconfirmed literature records also from Belgium (DE WILDEMAN et 

DURAND 1898 - 1907 (1898)), N. W. France (DES ABBAYES 1934), Italy (JATTA 1900), Rumania (CRE- 

TZOIU 1939; MORUZI, PETRIA et MANTU 1967), Bulgaria (NIKOLOFF 1931), U. S. A., California (TUCKE- 

RMAN 1866), Sri Lanka (Ceylon) (BISWAS 1947, almost certainly erroneous, referring to St. austroindicum 

LAMB), Central Africa (LINDAU 191 I), China (MULLER ARC. 1893, erroneous, = St. myriocarpum 

TH. FR.), Ecuador (KREMPELHUBER 1861, erroneous, = St. myriocarpum TH. FR.). 

Remarks: the type species of the genus. See p. 307 for discussion of typification. The described 

infraspecific taxa which really belong to St. paschale (var. serpens TH. FR. and f. colligatum LAMB) are 

obviously environmental modifications of no taxonomic significance.


2. Stereocaulon alpinum LAUR. 

ex FUNCK, Cryptogamische Gewachse des Fichtelgebirg's. Ed. I1 (1806 - 1838), Heft 33: 6 (1827); 

MAGN. Goteborgs K. Vetensk.- o. Vitterh.-Samh. Handl. ser. 4, 30 (7): 52 (1926); FREY, Rabenhorsts 

Krypt.-F1. 9, Abt. IV, 1. Halfte: 158 (1932); DAHL et KROG, Macrolichens of Denmark, Finland, Norway 

and Sweden: 141 (1973). - Stereocaulon tomentosum [var.] y. alpinum (LAuR.) TH.FR. De Stereoc. 

et Pilophor. Comment.: 30 (1857). -Stereocaulon tomentosum subsp.* alpinum (LAuR.) TH. FR. 

Monogr. Stereoc. et Pilophor.: 357 (1858). - Stereocaulon paschale [f.] y. alpinum (LAuR.) MUDD, A 

Manual of British Lichens: 66 (1861). - Stereocaulonpaschale var. alpinum (LAuR.) DU RIETZ, Svensk 

Bot. Tidskr. 20: 96 (1926). 

Typus: Austria, Alps of Tirol and Karnten, "besonders schon am Pasterzen Gletscher," but no 

holotype designated; collector and date of collection not known, but the material sent to FUNCK by 

LAURER, and distributed in the exsiccat Cryptog. Gewachs. d. Fichtelgeb. no. 684, accompanied by 

diagnosis in the schedae, may be regarded as isotype. 

Facultative synonyms: Stereocaulon tomentosum [var.] y. thyrsoideum SCHAER. Lichenum Helveticorum 

Spicilegium, Pars prima, sect. VI: 276 (1833). - Stereocaulon paschale [f.] b. thyrsoideum 

(SCHAER.) RABENH. Deutschlands Kryptogamen-Flora, 2, Abt. 1 : 112 (1845). - Stereocaulon tomentosum 

[subsp.]* alpinum [f.] a. verrucosum TH. FR. Monogr. Stereoc. et Pilophor.: 357 (1858). - Sterocaulon 

alpinum f. adpressurn MAGN. Goteborgs K. Vetensk. - o. Vitterh.-Samh. Handl. ser. 4, 30 (7): 55 

(1926). - Stereocaulon botryosum var. subincrustatum FREY, Rabenhorsts Krypt.-F1. 9, Abt. IV, 1. 

Halfte: 127 (1932). 

Exsicc.: ANZI, Lich. Exs. Ztal. Super. Fasc. I (1865) no. 26 (FH-TUCK); no. 27, as St. alpinum var. 

botryosum (BM, FH-TUCK). - ANZI, Lich. Prov. Sondr. et Novo-Comensi Exs. Fasc. I (date ?) no. 7, as 

St. tomentosum (UPS); no. 9A (ups); no. 9B, as St. alpinum f. botryosum (ups). - ARN. Lich. EXS. 

(Lich. Jurae) no. 65la (date ?) (PC, UPS. Note that FREY, 1932, p. 155, states that in M and W this number 

is St. grande (MAGN.) MAGN.); no. 1363a (1888) (PC. In FH and MICH it is St. rivulorum MAGN.). - 

BILLOT, Flora Exs., continue' par B~voux, GUICHARD et PAILLOT, no. 3797 (date ?), as St. tomentosum 

(us. The example in PC, however, is a mixture of St. alpinum and St. dactylophyllum FLK.). - BREUT. 

Flora German. Exs., Ser. 11, Cent. I1 (1843) no. 107, pr. p., as St. paschale var. thyrsoideum (s. Examples 

of this number in FH, PC and ups are however St. paschale (L.) HOFFM.). - DESM. Plantes Cryptogames 

de France, ed. 2, Ser. 11, Fasc. X (1857) no. 495, as St. tomentosum b. majus (FH). - ELENK. Lich. 

Flor. Ross. Fasc. I1 (1907 ?) no. 73a (BM, FH, LE). - Erb. Crittog. Ztal., ed. DE NOTARIS, CESATI, BAGLI- 

ETTO etc. Fasc. XV (1862) no. 736 (FH). - Erb. Crittog. Ztal., publ. Soc. Crittogamolog. Itat., Ser. IT, 

Fasc. VII (1870) no. 320, pr. p., as St. alpinum, alpestre (FH. Intermixed with St. incrustaturn FLK.); 

no. 320 bis, as St. alpinum, botryosum (FH, LD). - FOLLM. Lich. Exs. Sel. Mus. Hist. Nut. Casselensi 

Fasc. VII (1975) no. 134 (KASSEL). - FUNCK, Cryptog. Gewachs. des Fichtelgebirg's, Ed. 11, Heft 33 

(1827) no. 684 (isotype material) (FH, LE). -HARM. Lich. Gall. Rar. Exs. Fasc. I (1908) no. 25 (FH. 

With some intermixture of St. saxatile MAGN.). - HEPP, Flechten Europas, Band VI (1857) no. 303 

(FH-TUCK, PC). - Krypt. Exs. Vindob. Cent. I1 (1896) no. 152b (from Norway) (BM, COLO, FH, PR); 

same number (from Switzerland) (co~o, FH); Cent. XLIII (1961) no. 4237 (FH, LD, LE); Cent. XLVI 

(date ?) no. 4566 (FH). - Lich. Groenlandici Exs. (ed. Mus. Bot. Hauniense) Fasc. I (1972) no. 8 (LD). 

- MALME, Lich. Suec. Exs. Fasc. XXVII (date ?) no. 652 (ws. Seems to be a transitional state to the 

var. erectum FREY). - RABENH. Lich. Europ. Exs. Fasc. XXXII (1870) no. 859 (FH-TUCK, GB, MICH, 

WIS). - ROUM. Lich. Gall. Exs. Cent. I1 (1880) no. 164, as St. vesuvianum (UPS). - SCHAER. Lich. 

Helvet. Exs. Fasc. XI (1836) no 262, pr. p., as St. tomentosum B. majus (PC. The example in FH-TUCK is 

transitional to thevar. erectum FREY. FREY, 1932, p. 186, states that material of this number seen by him 

is a mixture of St. alpinum, St. tomentosum FR., and St. tomentosum var. alpestre FLOT. This number 

appears to be type material of Stereocaulon tomentosum B. majus SCHAER. Lichenum Helveticorum 

Spicilegium, Pars prima, Sect. VI: 276 (1833) and St. tomentosum a. granulosum SCHAER. Enumeratio 

Critica Lichenum Europaeorum: 181 (1850). In view of the heterogeneous nature of this material, 

both these epithets of SCHAERER should be discarded as nomina confusa); no. 263 (FH-TUCK, PC);


no. 264, as St. alpinum B. botryosum (FH-TUCK, PC). - THOMS. Lich. Arct. no. 37 (1960) (WIS). - 

TREV. Lichenotheca Veneta, Ser. I, Fasc. IV (1869) no. 144 (ups), - V~ZDA, Lich. Sel. Exs. Fasc. XXXIX 

(1971) no. 964 (FH, LD). - WARTMANN et SCHENK, Schweizer. Kryptog. Cent. I, Fasc. I1 (1862) no. 51 

(FH). 

Icon. : ANDERS, 1928, PI. XVIII, fig. 10. - FREY, 1932, fig. 25; PI. 111, fig. 5. - MIG. 1927 - 31, 

Lief. 268 - 277 (1928), PI. 74, fig. 1. - CERN., NADV. et SERV. 1956, PI. VI, fig. 5. - SCHENCK, 1914, 

PI. 29 a, b (field photograph). - POELT, 1969, PI. 3, fig. 13. - OZENDA et CLAUZADE, 1970, fig. 394 A, 

B, C. - DAHL et KRoG, 1973, fig. 49. - HENSS. et JAHNS, 1974, fig. 13.14~. 

Mat. chim.: atranorin, lobaric acid, B-sitosterin. Occasionally lobaric acid apparently 

absent or in too low a concentration to be demonstrated by present chromatographic techniques. 

H. KRISTINSSON (in litt.) reports an Icelandic specimen allegedly containing atranorin, 

lobaric acid and stictic acid7. A polyol compound, "Stereocaulit", described by 

ZOPF (1907); this shown to be identical with D-arabitol ("arabit") by HUNECK, FOLLMANN 

and REDON (1973). 

Distrib.: More extensive than that of St. paschale, and penetrating further polewards than that 

species. Sweden, Norway, Finland, France, W. and E. Germany, Switzerland, Austria, northern 

Italy, Czechoslovakia, Hungary, Poland, Bulgaria, U. S. S. R. (Siberia, Murmansk region, Ural Mts., 

Caucasus, Transbaikalia, Novaya Zemlya, Franz Josef Archipelago, Extreme Orient, Kamtchatka), 

Mongolian People's Republic, Iceland, Spitsbergen, Greenland, Jan Mayen, Canada (N. W. Territ., 

Yukon Territ., British Columbia, Alberta, Saskatchewan, Manitoba, Ontario, Quebec, Newfoundland, 

Labrador), Miquelon Island, U. S. A. (New Hampshire, Vermont, Colorado, Wyoming, Montana, 

Oregon, Washington, Alaska), Korea, northern India, Pakistan, Peru, Chile (Cautin, Aiskn, Magallanes), 

Argentina (Rio Negro, Santa Cruz, Tierra del Fuego), South Georgia, Antarctic Peninsula. 

Unconfirmed literature records also from Belgium (DE WILDEMAN and DURAND 1898 - 1907 (1898)), 

Rumania (MORUZI, PETRIA and MANTU 1967), China (ZAHLBRUCKNER 1934), Hawaii (MULLER ARG. 

1889), and Central Africa (ZAHLBRUCKNER and HAUMAN 1936, erroneous, referring to St. nigromaculatum 

DUVIGN.). 

Remarks: DAHL 1956, p. 359, gives an instructive analysis of the morphological, 

physiological and ecological differences between St. alpinum and St. paschale (L.) HOFFM. 

KLEMENT (1959) distinguishes in northern Lapland an association named the Stereocauletum 

alpinum, dominated by St. alpinum and including also St. paschale (L.) HOFFM. and St. tomentosum 

FR. 

Fo. platycladum FREY 

ex POELT, Bestimmungsschliis~e1 Europaischer Flechten: 647 (1969). 

Typus: France, Dauphine, La Bkrarde, Vallon du Veneon, "graviers silicieux, alluvions", leg. 

E. FREY and A. SCHMIDT, 1932 (no. 2513) (FREY, holotypus; FH, isotypus). 

Remarks: material referable to this form has been seen also from Finland, Switzerland, 

U.S.A. (Colorado), Canada (Alberta, Saskatchewan) and Greenland. 

Var. erectum FREY 

Rabenhorsts Krypt.-F1.9, Abt. IV, 1. Halfte: 164 (1932); Ber. Schweiz. Bot. Ges. 69: 196 (1959); MAGN. 

Bot. Not.: 469 (1934). - Stereocaulon paschale var. erectum (FREY) LAMB ex LLANO, J. Wash. Acad. 

Sci. 41: 197 (1951). 

Typus: Switzerland, Canton Graubiinden, Sarnaden, altit. 1725 m s. m., "Kies- und Sandalluvi- 

Dr. KRISTINSSON 

later (in litt. 1976) informed us that this result is not quite certain, since it is 

possible that the material extracted may have been contaminated by St. tomentosum FR.


onen des Inn nahe beim Bahnhof", leg. E. FREY, 1931 (no. 1636) (FREY, CAN. Lectotypus, lectotypit. 

mv.). 

Exsicc.: ARN. Lich. EXS. (Lich. Jurae) no. 1604 (1894), as St. alpinum (BM, FH, PC. But FREY, 1932, 

p. 155, states this number to be St. grande (MAGN.) MAGN. in M and W, and St. tomentosum FR. in z). 

- Krypt. Exs. Vidob. Cent. I1 (1896) no. 152, as St. alpinum (the specimen from Tirol) (FH, GB, LE, 

PC, PR, US, WELC); Cent. XL (1957) no. 3918, pr. p., as St. paschale var. paschale & var. erectum (FH, L, 

Lich. Sel. Scad. Exs. 

LD. All of these contain an intermixture of St. paschale (L.) HOFFM.). - MAGN. 

Fasc. IX (1935) no. 210 (PR). - MASS. Lich. Ztal EXS. Fasc. I (1855) no. 11, as St. alpinum (FH-TUCK, 

LE. But FREY, 1932, p. 160, states that an example seen by him is a mixture of St. tomentosum FR. and 

St. incrustatum FLK.).- MERR. Lich. Exs. Ser. 11, Fasc. IV (1926) no. 86, as St. tomentosum (c, co~o, 

DUKE, US, WIS). - VEZDA, Lich. Sel. Exs. Fasc. XL (1971) no. 989 (FH). 

Zcon.: FREY 1932, PI. I, fig. 8; PI. 111, fig. 6; PI. 111, fig. 4 pr. p., as St. grade; Fig. 24 a, b, as St. 

grande. 

Remarks: As widely distributed as the typical species. Difficult to distinguish from 

St. grande (MAGN.) MAGN. 

Var. gracilentum (TH. FR.) MAGN. 


F1.9, Abt. IV, 1. Halfte: 162 (1932). - Stereocaulon paschale [var.] P. gracilentum TH. FR. De Stereoc. 

et Pilophor. Comment.: 33 (1857). - Stereocaulon paschale [f.] P. gracilentum (TH. FR.) TH. FR. 

Lichenographia Scandinavica, Pars prima: 47 (1871). - Stereocaulon alpinum f. gracilentum (TH. FR.) 

LAMB ex POELT, Bestimmungsschliisse1 Europaischer Flechten: 646 (1969). 

Typus: Sweden, Gotland, Fare, "in collibus arenae volatilis", leg. C. STENHAMMAR (date ?) '(ups- 

TH FR, holotypus; UPS-MAGN, isotypus). 

Exsicc.: RAs. Lich. Fenn. Exs. Fasc. XVIII (1946) no. 879, as St. paschale (UPS). STENHAM. 

Lich. Suec. Exs. Fasc. 111 (1860) no. 81, as St. paschale b. gracilentum (FH-TUCK LD. In the latter there 

is some intermixture of St. paschale (L.) HOFFM., as is stated by FREY, 1932, p. 162). - FREY, 1932, p. 

162, states ROUM. Lich. Gall. Exs. Cent. I1 (1880) no. 162, as St. incrustatum, to be St. alpinum var. 

gracilentum in M-ARN; in UPS, however, we found this number to be St. incrustatum FLK. as stated on the 

label. 

Remarks: characteristic of movable dunes or alluvial sands, in either maritime or inland 

localities. In addition to material from the type locality, we have seen var. gracilentum from 

Sweden, Vasterbotten, "f oppna flackar i rished pi sand nara havet"; Finland, Nylandia, 

Tvarminne, "in arena volubilis" (Lich. Fenn. Exs. no. 879); Canada, Saskatchewan, Lake 

Athabasca, on sand dunes; and U.S.A., Alaska, "recent flood plain of the Matanuska River, 

on friable dusty alluvial soil". 

3. Stereocaulon antarcticum VAIN. 

Lichens, in: Rks. Voy. Belgica, Botanique: 16 (1903). 

Typus: Antarctic Peninsula, Gerlache Strait, Brabant Island, altit. 300 m s. m., leg. ? (Exp6d. 

Antarct. Belge, 1897 - 99), no, 247 (no. 3932 in TUR-VAIN). 

(EXS~CC.~) 

Zcon. : VAIN. 1903, PI. 11, fig. 7. 

Mat. chim. : atranorin and stictic acid, sometimes with norstictic acid in addition. 

Distrib.: Antarctic Peninsula and adjacent islands; Tierra del Fuego. 

No. 55 of the exsiccat FOLLM. 

Lich. Exs. Sel. Mus. Bot. Berol. (later titled Lich. Exs. Sel. Mus. 

Hist. Nat. Casselensi), called Stereocaulon antarcticum, consists of Catillaria corymbosa (HUE) LAMB. 

A corrected label was issued with Fasc. VIII (1975).


Remarks: closely related to St. glabrum (MULL. ARG.) VAIN. Widely misinterpreted in 

the literature. HUE'S records and descriptions (1908, 1915) refer to St. alpinum LAUR., 

according to his material seen by us in PC-HUE. The specimen recorded by HUNECK and 

FOLLMANN (1966), in which atranorin, lobaric acid, and /3-sitosterin were found, is likewise 

St. alpinum. RASANEN'S record (1932) of "Stereocaulon antarcticum" from Fuegia is also 

based on a misidentification; the specimen in question, from Cerro Millaldeo, leg. ROIVAINEN, 

1929 (H-RAS) is scanty and depauperate, not determinable with certainty, but obviously not 

the present species. The reported occurrence of St. antarcticum in Tristan da Cunha, determined 

by MAGNUSSON (LYNGE, 1937) could not be verified, as no specimen could be found 

either in UPS-MAGN or in o. We collected St. antarcticum in Argentina, Tierra del Fuego, 

near Ushuaia, in 1965, the first extra-Antarctic find. 

4. Stereocaulon austroindicum LAMB (n. sp.) 

Diagn.: Thallus primarius (ad saxum decompositum) persistens, granulosus; pseudopodetia 

erecta, vulgo sat brevia, usque 2 cm longa, aggregata, substrato arcte affixa, simplicia 

aut ramosa, ecorticata et etomentosa, phyllocladiis confertim vestita, apicibus vulgo alboefflorescentia, 

sed haud vere sorediata. Phyllocladia primum granulosa, deinque coralloidea 

vel coralloideo-squamulosa, ca. 1 mm longa. Cephalodia verruculosa vel botryosa, ca. 

1 mm lata, pallida, fere ecorticata. Apothecia terminalia, fere ab initio pileato-convexa, 

usque 2 rnrn lata, saepe tandem composita. Hypothecium vulgo leviter pigmentiferum. 

Hymenium 50-75 ,u altum. Sporae fusiformes, 3(-7)-septatae, (23-)3M2 p longae, 2.5- 

3.5 ,u latae. Atranorinum et acidum lobaricum continet (K+flavescens, PD+pallide flavescens). 

- Antea cum St. condensato HOFFM. confusum fuit, a quo praesertim cephalodiis 

pallidis differt; habitu St. octomerellum M~~LL. ARG. vage simulans, sed magis robustum et 

phyllocladiis saepe subsquamulosis differens. 

Typus: India, Madras State, Madurai Distr., Shembaganur, Palni Hills, altit. ca. 1950 m s. m., 

on decomposed earthy rock by roadside, leg. G. FOREAU, 1958 (FH, holotypus; AWAS, isotypus). 

Mat. chim. : atranorin and lobaric acid, or in some cases apparently atranorin only. 

Distrib.: southern India (Madras, Travancore) and Sri Lanka (Ceylon). 

Remarks: material of this species from Sri Lanka collected by ALMQUIST in 1879 was 

named "Stereocaulon condyloideum" and "Stereocaulon condensatum" by NYLANDER (1900), 

and was referred to St. leprocephalum VAIN. by VAINIO in his description of the latter species 

(1909). St. austroindicum can be easily distinguished from St. condensatum by its larger 

size and pale cephalodia, and from St. leprocephalum by the absence of true capitate soredia. 

The occasional accidental presence of blackish tufts of free-living Stigonema on the primary 

thallus probably led to the misidentification as St. condensatum. The cephalodial algae in 

the type material were examined by Dr. FRANCIS DROUET and identified as Anacystis montana 

(LIGHTF.) DR. et DAILY, Anacystis being according to him the correct name for Gloeocapsa 

KijTz. (see DROUET and DAILY, 1956). Other specimens examined had Stigonema in the 

cephalodia. Some specimens from Sri Lanka tend to be more slender, with the phyllocladia 

partly more distinctly coralloid, as in a specimen collected by THWAITES (BM, PC, UPS), but 

others, e.g. material collected by BECCARI (M), are quite typical in habitus and morphology. 

5. Stereocaulon azoreum (SCHAER.) NYL. 

Actes Soc. Linn. Bordeaux, 21: 287 (1856). - Stereocaulon tomentosum [var.] 6. azoreum SCHAER. 

Enumeratio Critica Lichenum Europaeorum: 182 (1850).


Typus: Azores, leg. GUTHN~K (date ?) (G-SCHAER, holotypus. not seen by us). 

Facultative synonyms: Stereocaulon Ieporinum TH. FR. De Stereocaulis et Pilophoris Commentatio: 

25 (1857). - Stereocaulon granulosum LAUR. ex HARTUNG, Neue Denkschr. Allg. Schweiz. Ges. Gesammten 

Naturwiss. 15 (4): 147, 151 (1857). - Stereocaulon tomentosum var. granulosum (LAuR.) OLN. 

Mem. Real. Acad. Ci. Barcelona, ser. 3,16: 47 (1921).O - Stereocaulon maderense TUCK. Lichenes, in: 

United States Exploring Expedition, 17, Botany I, Lower Cryptogamia: 122 (1874). - Stereocaulon 

Ieporinum [var.] a. elatum TH. FR. De Stereoc. et Pilophor. Comment.: 25 (1857). - Stereocaulon 

sphaerophoroides [f.] a. elatum (TH. FR.) TH. FR. Monogr. Stereoc. et Pilophor.: 349 (sep. 45) (1858). 

- Stereocaulon Ieporinum [var.] j3. pumilum TH. FR. De Stereoc. et Pilophor. Comment.: 25 (1857). 

- Stereocaulon sphaerophoroides [f.] j3. pumilum (TH. FR.) TH. FR. Monogr. Stereoc. et Pilophor.: 

349 (sep. 45) (1858). 

Exsicc.: ALMB. Lich. Afric. Fasc. IV (1974) no. 86, as St. sphaerophoroides (FH). - BORN~LLER, 

Plant. Exs. Mader. (1900) no. 124 (LD, PR), no. 125 (LD), no. 126 (GB, LD, PR), as St. sphaerophoroides; 

no. 127, as St. denudatum var. pulvinatum (LD). - BORNMULLER, Plant. Exs. Canar. (1901) no. 3238 

(LD, PR), no. 3244 (LD), as St. sphaerophoroides. - BOURGEAU, Plant. Canar. no. 37 (1846-47), as St. 

botryosum (PC-HUE). - BOURGEAU, Plant. Canar., iter secund. no. 1594 (1856), as St. ramulosum (c, 

PC-HUE). - HEPP, Flechten Europas Band VI, Heft 1 1 (1857) no. 305, as St. granulosum (FH, FH-TUCK, 

PC-HUE, s). - HUSNOT Plant. Canar. no. 213 (1866 ?), as St. sphaerophoroides (PC-HUE). - Krypt. Exs. 

Vindob. Cent. VIII (1902) no. 767, as St. sphaerophoroides (c, FH, LD, PR, S, US, WELC); Cent. XLIII 

(1960) no. 4239, as St. sphaerophoroides (FH, s). - MANDON, Lich. Mad2re no. 49 (date ?), as St. 

sphaerophoroides (BM, FH-TUCK, PC-HUE, s). - TAV. Lich. Lusit. Sel. Exs. Fasc. I11 (1955) no. 65, as St. 

sphaerophoroides (FH); Fasc. X (1968) no. 238, as St. sphaerophoroides (FH). - VEZDA, Lich. Sel. Exs. 

Fasc. V (1961) no. 113, as St. sphaerophoroides (FH). 

Icon.: TH. FR. 1858, P1. IX, fig. 4, as St. sphaerophoroides. - HEPP 1853-67 (1857) no. 305, as 

St. sphaerophoroides (spores). - NYL. 1860, P1. VII, fig. 9, as St. sphaerophoroides (conidia and conidiophores). 

- DUVIGN. 1956, fig. 9, as St. sphaerophoroides (sections of pseudopod. and apoth.). 

Mat. chim.: atranorin, lobaric acid, stictic acid, norstictic acid; possibly also constictic 

acid and consalazinic acid. This is the only species of Stereocaulon in which lobaric acid and 

stictic acid are known with certainty to coexist. 

Distrib.: Azores, Madeira, Canary Islands. Unverified record (almost certainly incorrect): 

U. S. A., N. Carolina (NYLANDER, 1860; CURTIS, 1867). Erroneous record: Ecuador (KREMPELHUBER 

1861), = St. myriocarpum TH. FR. 

Remarks: related to St. tomentosum FR. An interesting species not only for its peculiar 

chemistry but also on account of lacking cephalodia. DUVIGNEAUD (1956) states that its 

phyllocladia contain Scytonema-filaments in addition to the normal phycobiont. Since 

1858 the name St. sphaerophoroides TUCK. has been consistently used for this species. This 

is however incorrect for the following reasons. The protologue of St. sphaerophoroides 

TUCK., An Enumeration of North American Lichenes, p. 52, footnote (1845) mentions no 

locality for the species, but states that it was collected by L. GOULDING (ex herb. LAMBERT). 

TH. FRIES (1858) adopted the name St. sphaerophoroides TUCK. for the present species, placing 

his own St. leporinum TH. FR. in synonymy, and stating that he had received the type material 

of TUCKERMAN'S species and found it to be the same as St. leporinum. This original specimen 

was labelled "In monte ignivomo Ind. Occid.? Rev. L. GOULDING", and we have seen it in 

UPS-TH FR; it is not the present species, but St. virgatum AcH., a West Indian endemic. TH. 

FRIES was therefore mistaken in identifying it with his St. leporinum. The protologue of 

Acc. to ZAHLBRUCKNER, Cat. Lich. Univ.; not seen by us. If it is based on LAURER'S epithet it 

is an illegitimate homonym; cfr. St. tomentosum [var.] a. granulosum SCHAER. Enum. Crit. Lich. Europ. : 

181 (1850), a nomen confusum (see p. 331).

I. MACKENZIE LAMB: A Conspectus of Stereocaz~lon 207 

St. sphaerophoroides cites only the GOULDING specimen, and therefore St. sphaerophoroides 

must be considered a synonym of St. virgatum ACH. The specimen in FH-TUCK from the 

Azores, leg. HEWETT C. WATSON, 1842, said to be the type of St. sphaerophoroides by DODGE 

(1929, p. 143), is not type material. The first epithet legitimately applicable to the present 

species seems to be St. azoreum (SCHAER.) NYL. (1856), antedating St. leporinum TH. FR. by 

one year. NYLANDER'S Prodromus Lichenographiae Galliae et Algeriae was reissued as a 

reprint with separate pagination in 1857, but originally appeared in volume 21 of Actes Soc. 

Linn. Bordeaux, occupying pages 249-467 of that volume, starting with part 4 ("4e Livraison"). 

The dates of issue of the various parts of this volume, which extends to 1858, are not 

clearly stated, but NYLANDER'S paper appears to have been published in parts 4 and 5 before 

the end of 1856. The priority over St. leporinum TH. FR. is further confirmed by NYLANDER'S 

remark in Bot. Not. 1857 (October), p. 167' "S. leporinum Fr. jam a me azoreum dictum fuit 

in Prodr. Lich. Gall. Alger. p. 41." 

6. Stereocaulon botryosum ACH. 

Lichenographia Universalis: 581 (1810). Emend. FREY, Rabenhorsts Krypt.-FI. 9, Abt. IV, 1. Halfte: 

120 (1932); FREY, 1. c.; Ber. Schweiz. Bot. Ges. 69: 192 (1959); MAGN. Gijteborgs K. Vetensk.- o. 

Vitterh.-Samh. Handl. ser. 4, 30 (7): 33 (1926), as St. fastigiatum; POELT, Bestimmung~~chliis~el Europaischer 

Flechten : 643 (1969); DAHL et KROG, Macrolichens of Denmark, Finland, Norway and Sweden: 

134 (1973). - Stereocaulon alpinum var. botryosum (AcH.) SCHAER. Lichenum Helveticorum 

Spicilegium, Pars prima (1823421, sect. VI: 277 (1833). - Stereocaulon alpinum f. botryosum (AcH). 

FR. Lichenographia Europaea Reformata: 204 (1831). - Stereocaulon tomentosum var. botryosum 

(AcH.) NYL. Lichenes Scandinaviae: 64 (1861). 

Typus: Switzerland ("ad montes Helvetiae"), leg. SCHLEICHER (date ?) (H-ACH, lectotypus). 

Facultative synonyms: Stereocaulon fastigiatum ANZI, Catalogus Lichenum quos in Provincia 

Sondriensi: 11 (1860). - Stereocaulon evolutum var. fastigiatum (ANZI) TH. FR. Lichenographia Scandinavica, 

Pars prima: 45 (1871). - Stereocaulon evolutum f. fastigiatum (ANZI) MIG. Kryptogamen- 

Flora von Deutschland, ~eutsch-bsterreich und der Schweiz, 4 (2), Lief. 268-277: 67 (1928). 

Exsicc.: ANZI, Lich. Rar. Langob. Exs. Fasc. I(1861) no. 16, as St. fastigiatum (authentic,probably 

isotype material) (BM, FH, s). - ARN. Lich. EXS. (Lich. Jurae), no. 1575 (1893), as St. alpinum f. botryosum 

(BM, FH, PC, S, UPS). - Lichenotheca Fenn. Fasc. XLII (1957) no. 1034, as St. paschale var. grande 

(TUR). - MALME, Lich. Suec. Exs. Fasc. XL (1926) no. 979a, as St. fastigiatum (s. In LD, however, this 

number is St. depressum (FREY) LAMB). - ROUM. Lich. GUN. Exs. Cent. I1 (1880) no. 163, as St. alpinum 

(UPS); no. 165, as St, denudatum (ups, mixed with Sphaerophorus sp.). 

Icon.: LYNGE 1928, PI. VI, figs. 3,4. - FREY 1932, fig. 19 a - c, PI. I, fig. 4 (lectotype specimen in 

H-ACH), P1. 11, fig. 1; 1959, fig. 8. - OZENDA et CLAUZADE 1970, fig. 390 A, B, C. - DAHL et KROG 

1973, fig. 41. 

Mat. chim.: atranorin and porphyrilic acid. 

Distrib.: boreal-arctic circumpolar and in the Alps of central Europe. Sweden, Norway, Finland, 

France, Switzerland, Austria, northern Italy, U.S.S.R. (Ural Mts., Novaya Zemlya, Franz Josef 

Archipelago, Extreme Orient), Iceland, Spitsbergen, Greenland, Canada (N. W. Territ., Alberta), 

U. S. A. (Alaska). Unconfirmed literature records from the British Isles (LEIGHTON, 1871, 1879; 

LAMB, 1941 ; KERSHAW, 1960, all probably erroneous), U. S. A., Oregon (MAGNUSSON, 1939), Canada, 

Labrador and Qdbec (DIx, 1956, as St. fastigiatum), and Corsica (WERNER and DESCHATRES, 1970). 

A record from Ecuador (KUNTH, 1822, 1824 - 25) almost certainly erroneous. Definitely erroneous 

are records from South Georgia (Du RIETZ, 1926a) and Argentine Patagonia (LAMB, 1959), both referring 

to St. glabrum (MULL. ARG.) VAIN.; St. botryosum apparently does not occur in the southern 

hemisphere.


Remarks: of the described infraspecific taxa, most belong to other species. A subspecies 

islandicum described by DUVIGNEAUD (1941) from Iceland is doubtful, the type having 

been lost, and should be discarded as a nomen dubium (see p. 321). 

Fo. congestum (MAGN.) FREY 

Rabenhorsts Krypt.-F1. 9, Abt. IV, 1. Halfte: 125 (1932); POELT, Be~timmung~~chlii~~el Europaischer 

Flechten: 643 (1969). - Stereocaulon fastigiatum f. congestum MAGN. Goteborgs K. Vetensk.- o. 

Vitterh.-Samh. Handl. ser. 4, 30 (7): 38 (1926). 

Typus: Sweden, Torne Lappmark, Kopparlsen, altit. 600 m s. m., leg. A. H. MAGNUSSON, 1921 

(no. 6074) (UPS-MAGN, lectotypus, lectotypif. nov.; seep. 310). 

Exsicc.: ARN. Lich. Exs. (Lich. Jurae) no. 651b (date ?), as St. alpinum (var.) (PC, s. The example 

in S has some intermixture of St. vesuvianum PERS. var. nodulosum (WALLR.) LAMB, and another in 

UPS is St. rivulorum MAGN.). - Krypt. Exs. Vindob. Cent. XXXVII (1948) no. 3664, as St. fastigiatum 

var. dissolutum (co~o, FH, LD, LIL, S, UPS, US). 

Remarks: widely distributed and just as common as the typical species. 

Fo. finmarkicum (MAGN.) FREY 

Rabenhorsts Krypt.-F1. 9, Abt. IV, 1. Halfte: 126 (1932); POELT, Be~timmung~~chlii~~e1 Europaischer 

Flechten: 644 (1969). - Stereocaulon fastigiatum f. finmarkicum MAGN. Goteborgs K. '~etensk.- 0. 


Typus: Norway, Finmark, Mikkelskog-Sjuosjaure, leg. B. LYNGE, 1924 (UPS-MAGN, holotypus). 

Remarks: rare; seen only from the type locality and from one locality in Sweden (Torne 

Lappmark). 

Fo. irregulare (MAGN.) FREY 

Rabenhorsts Krypt.-F1. 9, Abt. IV, 1. Halfte: 126 (1932); POELT, Be~timmung~~~hlii~~el Europaischer 

Flechten: 643 (1969). - Stereocaulon fastigiatum f. irregulare MAGN. Goteborgs K. Vetensk.- o. Vitterh. 

-Sarnh. Handl. ser. 4, 30 (7): 35 (1926). 

Typus: Sweden, Lycksele Lappmark, Brandsfjallet, altit. 750 m s. m., "near a waterfall", leg. A. 

H. MAGNUSSON, 1924 (no. 8959) (UPS-MAGN, lectotypus; lectotypif. FREY, 1932,l. c.). 

Icon.: FREY 1932, PI. 11, fig. 2 (lectotype specimen). 

Remarks: rare; seen only from the type locality and from one other locality in Sweden 

(Torne Lappmark). 

Fo. simplicior (MAGN.) FREY 

Rabenhorsts Krypt.-F1. 9, Abt. IV, 1. Halfte: 125 (1932); POELT, Bestimmungsschliissel Europaischer 

Flechten: 643 (1969). - Stereocaulon fastigiatum f. simplicior MAGN. Goteborgs K. Vetensk.- o. 


Typus: Sweden, Lycksele Lappmark, par. Tarna, Bjorkfors, Syterbacken, altit. 450 m s. m., leg. 

A. H. MAGNUSSON, 1924 (no. 8375) (UPS-MAGN, holotypus). 

Remarks: known only from the type specimen. 

Fo. subglobuliferum LAMB (n. f.) 

Diagn.: Statura et aspectu ut in f. congesto, sed sorediis genuinis capitiformibus globosis 

praeditum. 

Typus: Norway, Sor-Trondelag, Knutsho, altit. ca. 1400 m s. m., leg. I. M. LAMB, 1954 (no. 7039)


(FH, holotypus). 

Remarks: the type material was of scarce occurrence on a rock in the lichen-heath, firmly 

attached, forming compact pulvinate clumps ca. 4 cm diarn. with the typical cauliflower-like 

habitus and woody, fastigiate, glabrous pseudopodetia. Soredia PDffaint yellow; atranorin 

and porphyrilic acid demonstrated by chromatography. Resembles St. spathuliferum VAIN. 

f. globuliferum (MAGN.) LAMB and St. capitellaturn MAGN., but has the chemical constitution 

of St. botryosum. We have seen also one specimen from W. Greenland referable to this form. 

7. Stereocaulon capitellaturn MAGN. 


F1. 9, Abt. IV, 1. Halfte: 118 (1932); POELT, Be~timmung~~chlu~~el Europaischer Flechten: 636 (1969). 

opus: Sweden, Lycksele Lappmark, par. Tarna, Bjorkfors, Syterbacken, leg. A. H. MAGNUSSON, 

1924 (no. 8380) (UPS-MAGN, holotypus; s, isotypus). 

Facultative synonym: Stereocaulon farinaceurn MAGN. Goteborgs K. Vetensk.- o. Vitterh.-Samh. 

Handl. ser. 4, 30 (7): 72 (1926). 

Exsicc. : HAV. Lich. Exs. Norveg. no. 609 (1939) (BG). (Note: the material from Switzerland distributed 

as "Stereocaulon farinaceurn" in POELT et al. Lich. Alpiurn Fasc. VI (1961) no. 116 (FH, GZU, 

KLEM, LD) is a different species, containing lobaric acid; we have not been able to determine it with 

certainty.) 

Icon. : DAHL et KROG 1973, fig. 42. - FREY 1932, fig. 18; fig. 17 (as St. farinaceurn); 1959, fig. 6 

(as St. farinaceurn). 

Mat. chirn.: presumed full chemical spectrum: atranorin, perlatolic acid, anziaic acid, 

miriquidic acid (the type specimen belongs here). 

Deficient phase I: atranorin, perlatolic acid, anziaic acid (the type specimen of St. farinaceum 

belongs here). 

Deficient phase 11: atranorin only (one specimen from Greenland). 

Distrib.: Sweden, Norway, Finland, Austria, Switzerland, Iceland, Greenland. 

Remarks: examination of fairly copious material has convinced us that the differences 

supposed to separate St. farinaceurn from St. capitellatum are quite variable and taxonomically 

insignificant. 

8. Stereocaulon condensaturn HOFFM. 

Deutschlands Flora oder Botanisches Taschenbuch, Zweyter Theil, Cryptogamie: 130 (1796); FREY, 

Rabenhorsts Krypt.-Fl. 9, Abt. IV, I. Halfte: 92 (1932); POELT, Be~timmung~~chIu~~e1 Europaischer 

Flechten: 641 (1969); DAHL et KROG, Macrolichens of Denmark, Finland, Norway and Sweden: 135 

(1973). - Lichen condensatus (HOFFM.) ACH. Lichenographiae Suecicae Prodromus: 209 (1798). - Stereocaulon 

paschale [var.] j3. condensaturn (HOFFM.) SCHAER. Lichenurn Helveticorum Spicilegium, Pars 

prima, Sect. VI: 273 (1833). - Cereolus condensatus (HOFFM.) BOIST. Nouvelle Flore des Lichens, 2e. 

Partie: 33 (1903). 

Typus: Germany, no exact locality indicated, "in ericetis, montosis". Not seen by us. 

Facultative synonyms: Patellariapileata [var.] a. crustacea WALLR. Flora Cryptogamica Germaniae, 

Pars prior: 441 (1831). - Stereocaulon condensaturn f. crustaceurn (WALLR.) RABENH. Lich. Europ. Exs. 

Fasc. XIIT (1858) no. 370. - Patellaria pileata [var.] j3. tharnnodes WALLR. Flora Cryptogamica Germaniae, 

Pars prior: 441 (1831). - Stereocaulon acaulon NYL. Flora, 59: 232 (1876). - Stereocaulon 

condensaturn var. acaulon (NYL.) OLIV. Expose Systkmatique des Lichens de I'Ouest et du Nord-Ouest 

de la France, 1: 101 (1897). - Stereocaulon condensaturn f. acaulon (NYL.) MAGN. Goteborgs K. 

Vetensk.- o. Vitterh.-Samh. Handl. ser. 4, 30 (7) 68 (1926). - Cereolus condensatus subvar. acaulon 

("acaulis") (NYL.) BOIST. Nouvelle Flore des Lichens, 2e. Partie: 33 (1903). - Stereocaulon condensaturn


[var.] a. minor EGELING, Ber. Ver. Naturkunde in Cassel, 28: 93 (1881). - Stereocaulon condyloideurn 

ACH. Methodus qua Omnes Detectos Lichenes, Suppl.: 51 (1803). - Stereocaulon condensaturn subsp. 

condyloideurn (AcH.) NYL. Synopsis Methodica Lichenum, 1 (2): 251 (1 860). - Stereocaulon condensaturn 

var. condyloideurn (AcH.) NYL. Lichenes Scandinaviae: 66 (1 861). - Stereocaulon condensaturn f. 

condyloideurn (AcH.) MAGN. Goteborgs K. Vetensk.- o. Vitterh.-Samh. Handl. ser. 4, 30 (7): 68 (1926). 

- Stereocaulon condensaturn f. athallinurn SANDST. ex ERICHS. Ann. Mycol. 36, 143 (1938). - Stereocaulon 

condensaturn f. septentrionale MAGN. Goteborgs K. Vetensk.- o. Vitterh.-Samh. Handl. ser. 4, 

30 (7): 67 (1926). 

Exsicc.: (Note: ANZI, Lich. EXS. Ztal. Super. Fasc. I (1865) no. 29 in FH is not St. condensaturn as 

named; it may be St. glareosurn (SAV.) MAGN., but is too poor for certain identification.) ASAH. Lich. 

Japon. Exs. Fasc. VI (1959) no. 293 (UPS). - BARTLING et HAMPE, Veget. Cellul. German. Septentr., 

Lich. Decas I1 (1832) no. 18 (M. The numbers on the labels of this exsiccat do not agree with those 

cited by LYNGE, 1915-20). - BREUT. Flora German. Exs. Ser. 11, Cent. I1 (1843) no. 106 (FH. An example 

in S is however St. vesuvianurn PERS. var. nodulosurn (WALLR.) LAMB). - CUMM., WILLIAMS et SEYMOUR, 

Decades N. Arner. Lich. Decas I11 (1892) no. 26 (DUKE, MSC, US, WELC, WIS). - CUMM. et al. Lich. Bor.- 

Arner. Decas XI (1894) no. 159 (FH, PR, S, WELC). - FR. Lich. Suec. Exs. Fasc. I11 (1824) no. 88, as St. 

pileaturn (FH. An example in LD is a mixture of St. condensaturn and St. pileaturn AcH.). - TH. FR. 

Lich. Scandin. Rar. et Crit. Exs. Fasc. 111 (1865) no. 64 (BM, CAN, LD, PC, S, US). - FUNCK, Cryptog. 

Gewachse des Fichtelgebirg's Ed. 11, Heft 16 (1810) no. 343, as St. condyloideurn (FH, PR). -HARM. 

Lich. Gall. Rar. Exs. Fasc. I (1908) no. 26 (FH). - HARM. Lich. in Lotharingia (1 887-99) no. 156 (FH). - 

HAV. Lich. EXS. Norveg. Fasc. XXI (1947) no. 657 (BG). - HEPP, FIechten Europas Band VI (1857) no. 

300 (FH, PC, s). - KRAWIEC, Lichenotheca Polonica Fasc. I1 (1935) no. 71 (s). - KUT~K, Lich. Bohern. 

Fasc. I1 (1913) no. 75 (PR, u). - LEIGHT. Lich. Brit. Exs. Fasc. X (1858) no. 295 (EM, FH, LD, S, US). - 

Lichenotheca Fenn. Fasc. XLIV (1958) no. 1082 (FH). (Note: Lichenotheca Fenn. Fasc. XXXVI (1956) 

no. 883 in FH is not St. condensaturn as named, but a species of Lepraria.) - MAGN. Lich. Sel. Scand. 

Lich. Suec. 

Exs. Fasc. VII (1933) no. 155, as St. condensaturn var. condyloideurn (GB, PR). - MALME, 

Exs. Fasc. XX (1915) no. 486 (s, wls). - MIG. Cryptog. German., Austr. et Helvet. Exs. Fasc. XXV 

(1904) no. 122 (FH, MSC, PR, s). - MUDD, Lich. Brit. Exs. Fasc. I (1861) no. 33 (EM, FH). - NORRL. et 

NYL. Herb. Lich. Fenn. Fasc. I1 (1875) no. 87, as St. condensaturn var. condyloideurn (FH). - OLIV. 

Herb. Lich. Orne et Calvados (Lich. Exs.) Fasc. VII (1882) no. 315, as St. acaulon (DUKE). - PIantae 

Varsav. Exs. Fasc. I1 (1964) no. 40 (LD, s). - RABENH. Lich. Europ. Exs. Fasc. V (1856) no. 138 (FH, 

PR, S. The example in PR contains also St. paschale (L.) HOFFM.); Fasc. XI11 (1858) no. 370, as St. condensaturn 

B. crustaceurn (FH, s). - REICHENB. et SCHUB. Lich. Exs. Fasc. I11 (1823) no. 68, as St. pileaturn 

(M). - ROUM. Lich. GUN. Exs. Cent. V (1883) no. 424 (ups. Acc. to FREY, 1932, p. 99, an example 

of this number seen by him is St. pileaturn AcH.); Cent. VI (date ?) no. 511, as St. acaulon (UPS). - 

SCHADE, STOLLE et RIEHMER, Lkh. Sax. EXS. Decas L (1932) no. 496 (BM, KLEM). - SCHAER. Li~h. 

Helvet. Exs. Fasc. XXI (1847) no. 509, as St. paschale B. condensaturn (FH, PC). - STENHAM. Lich. 

Suec. Exs. Fasc. I11 (1860) no. 84 (BM, FH, LD, US). - SUZA, Lich. Bohernoslovak. Exs. Fasc. I (1926) 

no. 16 (BM, FH, LD). - VEZDA, Lich. Bohernoslovak. Exs. Fasc. I (1957) no. 42 (LD). - VBZDA, Lich. 

Sel. Exs. Fasc. VI (1962) no. 143 (LD); Fasc. XVII (1965) no. 414 (LD, s). - WESTEND. et WALLAYS 

Herb. Cryptog. Belge Fasc. XXVII (1859) no. 1315 (FH). -ZAHLBR. Lich. Rarior. Exs. no. 58 (1904) 

(PC, ups). 

Icon.: GALL~E 1972, PI. 1. fig. 6; PI. 51, figs. 44146; PI. 52, figs. 447452; PI. 53, figs. 453456; 

P1. 54, figs. 457462; P1. 55, figs. 463469; PI. 56, figs. 470-473. - FREY 1932, fig. 11; 1959, fig. 2. - 

HALE 1969, fig. 384. - OZENDA et CLAUZADE 1970, fig. 392 A, B. - YOSHIM. 1974, PI. 33, fig. 328. 

Mat. chim.: atranorin and lobaric acid, or sometimes apparently atranorin only. 

Distrib.: northern hemisphere, temperate to boreal-arctic, amphiatlantic and possibly circumpolar. 

Sweden, Norway, Denmark, Finland, Netherlands, Belgium, England, Scotland, France, 

Germany (W. and E.), Czechoslovakia, Poland, U. S. S. R. (Byelorussian SSR, Karelo-Finnish SSR, 

Kamtchatka), Mongolian People's Republic, Greenland, Canada (Quebec, Labrador, Ontario, Manitoba, 

Saskatchewan, N. W. Territ.), U. S. A. (Connecticut, Massachusetts, New Hampshire, Wisconsin,


Alaska, Hawaii), Japan, doubtfully from Indonesia (Java) (see below). Unconfirmed literature records 

from Ireland (PORTER 1948), Italy (JATTA 1900), Rumania (MORUZI, PETRIA and MANTU 1967), U.S.S.R. 

Lithuania (MINCEVIEIUS 1963), Kola Peninsula (TRASS, MAGI and PARN 1963; DOMBROVSKAYA 1970), 

Ural Mts. (SOROKIN 1877), Canada, British Columbia (OTTO 1968), U. S. A., Washington State (How- 

ARD 1955), Taiwan (Formosa) (ZAHLBRUCKNER 1933), and Ecuador (ZAHLBRUCKNER 1905, highly improbable). 

Erroneous records are from Sri Lanka (Ceylon) (NYLANDER 1900),=St. austroindicum 

LAMB, and northern India, Kashrnir (AWASTHI and SINGH 1970), revised by us and found to be St. 

glareosum (SAV.) MAGN. 

Remarks: all the facultative synonyms represent mere growth-stages or non-taxonomic 

modifications of the species. St. condensatum is most closely related to St. glareosum (SAV.) 

MAGN. and St. incrustaturn FLK. It was recorded from Indonesia (Java) by MONTAGNE and 

VAN DEN BOSCH (1856); ZAHLBRUCKNER (1956) considered this record probably to refer to 

St. graminosum SCHAER. However, our own study of the specimen in question, from Mt. 

Sindoro, leg. JUNGHUHN (L) showed it to have the diagnostic characters of St. condensatum, 

but the material was poorly preserved and hence its identity somewhat doubtful. St. condensatum 

certainly occurs in Hawaii and Japan, but is a rare species there. 

9. Stereocaulon cyrnosurn CROMB. 

J. Linn. Soc., Bot. 15: 182 (1876); J. Bot. 15: 103 (1877); DODGE, Ann. Cryptog. Exot. 2 (2): 116 (1929). 

- Argopsis cymosa (CROMB.) STIZENB. Ber. Thatigk. St. Gallischen Naturwiss. Ges. 1889-90: 237 (1891). 

Typus: Kerguelen Island, Observatory Bay, "on the ground", leg. A. E. EATON, 1874-75 (K, holotypus; 

BM, isotypus). 

Facultative synonym: Stereocaulon depreaultii f. corallinum HUE, Nouv. Arch. Mus. Hist. Nat. 

ser. 3, 2: 248 (1890) (pr. p.; vide infra). 

Mat. chim.: atranorin and lobaric acid, plus traces of some unidentified substances. 

Distrib.: Kerguelen and Marion Islands (southern Indian Ocean). 

Remarks: identical in vegetative morphology with Argopsis friesiana M~~LL. ARC., which 

differs in having submuriform spores and a different chemistry (see LAMB, 1974). Both species 

occur in Kerguelen, apparently growing intermingled. The material to which HUE gave the 

name Stereocaulon depreaultii f. corallinum was collected on Kerguelen by J. D. HOOKER 

and named first St.paschale by HOOKER and TAYLOR (1844), then St. corallinum by BABINGTON 

ex HOOKER (184547); NYLANDER (1860) referred it to St. depreaultii DEL. as "fere forma", 

and HUE, following this suggestion, made the combination St. depreaultii f. corallinum, attributing 

it to BABINGTON as original author. The name is of erroneous application, because 

BABINGTON did not intend the epithet corallinum as a new taxon, but used it with reference to 

FRIES, i.e. St. dactylophyllum FLORKE; see LAMB, 1974, p. 457, for further details. 

10. Stereocaulon dactylophyllurn FLORKE 

Deutsche Lichenen, Lief. IV: 13 (1819); RIDD. Bot. Gaz. 50: 291 (1910), as St. coralloides; MAGN. 

Goteborgs K. Vetensk.- o. Vitterh.-Samh. Handl. ser. 4, 30 (7): 25 (1926), as St. coralloides; MIG. 

Kryptogamen-Flora von Deutschland, Deutsch-bsterreich und der Schweiz, 4, Lief. 268-277: 66 (1 928), 

as St. coralloides; FREY, Rabenhorsts Krypt.-F1. 9, Abt. IV, 1. Halfte: 189 (1932), as St. coralloides; 

Oz. et CLAUZ. Les Lichens: 474 (1970). - Stereocaulon coralloides [var.] a. dactylophyllum (FLORKE) 

TH. FR. De Stereocaulis et Pilophoris Comrnentatio: 16 (1857). - Stereocaulon coralloides [f.] a. 

dactylophyllum (FLORKE) TH. FR. Lichenographia Scandinavica, Pars prima: 44 (1871). - Stereocaulon 

paschale [subsp.] B. dactylophyllum (FLORKE) BRANTH et ROSTR. Bot. Tidsskr. 3: 162 (1869). - Stereocaulon 

coralloides subvar. dactylophyllum (FLORKE) BOIST. Nouvelle Flore des Lichens, 2e. Partie: 36


(1903). 

Typus: Germany, 3 localities mentioned (Harz, Rehberger Graben and Andreasberg; Fichtelgebirge), 

leg. H. G. FLORKE, 1797, no holotype designated. The authentic material distributed by 

FL~~RKE in his exsiccat Deutsch. Lich. Fasc. IV (1819) no. 78, although without mention of locality or 

date, probably came from one of these localities and should be regarded as isolectotype (DODGE, 1929, 

p. 114). 

Facultative synonyms: Stereocaulon coralloides FR. Schedulae Criticae de Lichenibus Exsiccatis 

Sueciae, Part 111: 24 (1825). - Stereocaulon paschale [subsp.] a. coralloides (FR.) BRANTH et ROSTR. 

Bot. Tidsskr. 3: 162 (1869). - Stereocaulon dactylophyllum [var.] P. major SOMM. Supplementum Florae 

Lapponicae: 125 (1826). - Stereocaulon coralloides var. orientalis RAS. Ann Bot. Soc. Zool. Bot. Fenn. 

Vanamo, 12 (1): 55 (1939). (Note: the name Stereocaulon corallinum (L.) SCHRAD. was frequently used 

by earlier authors, from FRIES onwards, to designate the present species, but wrongly, because it is based 

on Lichen corallinus L. = Pertusaria corallina (L.) ARN. ; see LANJOUW et al., 1966, p. 247.) 

Exsicc.: ABB. Lich. Armor. Spect. Exs. Fasc. V (1932) no. 83, as St. coralloides (LD, ups). - ANZI, 

Lich. Exs. Ztal. Super. Fasc. 1 (1865) no. 28, as St. coralloides (BM, FH, s). - ANZI, Lich. Prov. Sondr. 

et Novo-Comensi Exs. Fasc. I (date ?) no. 8, as St. corallinum (ups). - ARN. Lich. Exs. (Lich. Jurae) 

no. 1483a, b (1889), as St. coralloides (BM, FH, PC, s). - BARTLING et HAMPE, Veget. Cellul. German. 

Septentr., Lich. Decas V (date ?) no. 46, as St. corallinum (M). - BREUT. Flora German. Exs., ser. 11, 

Cent. I1 (1843) no. 108, as St. corallinum (FH, s), - BRITZ. Lich. EXS. FI. Augsburgs, Suppl. Fasc. 111 

(1905) no. 542, as St. coralloides (BM, FH, PR). - CLAUD. et HARM. Lich. Gall. praecip. Exs. Fasc. 111 

(1904) no. 107, as St. coralloides (FH). - CROMB. Lich. Brit. Exs. Cent. I1 (1877) no. 119, as St. coral- 

Ioides (ups); no. 120 pr. p., as St. evolutum (St. dactylophyllum in BM, St. evolutum GRAEWE in ups). - 

DESM. Plantes Cryptogames de France, ed. 2, Ser. 11, Fasc. I (1853) no. 46, as St. corallinum (FH). - 

ELENK. Lich. Flor. Ross. Fasc. I1 (1907) no. 71, as St. coralloides var. dactylophyllum (FH). - Erb. 

Crittog. Ztal., ed, DE NOTARIS, CESATI, BAGLIETTO etc. Fasc. I (1858) no. 39, as St. corallinum (FH). - 

FLAG. Lich. Franche-Comptk Fasc. VII (1884) no. 306, as St. corallinum (m). - FLORKE, Deutsch. Lich. 

Fasc. IV (1819) no. 78 (PR, UPS, isolectotype material). -Flora Exs. Austro-Hungar. Fasc. VIII (1885) 

no. 1538, as St. coralloides (FH, PC, PR, s). - FR. Lich. Suec. Exs. Fasc. IV (1825) no. 118, as St. coral- 

Ioides (FH-TUCK, LD, M, s). - FUNCK, Cryptog. Gewachse des Fichtelgebirg's, Ed. 1, Heft 1 (1801) no. 

20, as St. paschale (PR. Mixed with St. tomentosum FR.). - Ed. 2, Heft 5 (1806) no. 117, as St. paschale 

(FH). -HARM. Guide Ekm. Lich. Vol. I (1904) no. 28, as St. coralloides (m). -HARM. Lich. in Lotharingia 

(1887-99) no. 158, as St. coralloides (FH). - HAV. Lich. Exs. Norveg. Fasc. VI (1904) no. 242, 

as St. coralloides (BG). - HAV. Lich. Norveg. Occid. Exs. Fasc. V (1935) no. 103, as St. coralloides 

(PR). - HEPP, Flechten Europas Band 111, Heft 5 (1853) no. 114, as St. corallinum (FH-TUCK, PC, s). - 

W. JOHNS. 

et 

North of England Lich.-Herb. Fasc. VI (1896) no. 210, as St. coralloides (BM). - KAVINA 

HILITZ. Crypt. eech. Exs. Fasc. I1 (1933) no. 77, as St. coralloides (BM, PR). - Krypt. Exs. Vindob. 

Cent. IV (1898) no. 355, 355b, as St. coralloides (FH, PC, PR, S, WELC). - KUROK. Lich. Rar. et Crit. Exs. 

Fasc. I1 (1969) no. 96 (lapsu typogr. "doctylophyllum") (FH). - LEIGHT. Lich. Brit. Exs. Fasc. V (1854) 

no. 148, as St. paschale (BM, FH. In us it is a mixture of St. vesuvianum PERs. var. nodulosum (WALLR.) 

LAMB and probably St. evolutum GRAEWE). - MALBR. Lich. Normand. Fasc. V (1869) no. 213, as St. 

coralloides (LD, s). - MALME, Lich. Suec. Exs. Fasc. XXI (1915) no. 508, as St. coralloides, pr. p. (PR, 

wrs. Both mixed with St. saxatile MAGN. An example in s is a mixture of St. evolutum GRAEWE and 

St. saxatile MAGN.). - MASS. Lich. Ztal. Exs. Fasc. I1 (1856) no. 44, as St. corallinum (BM, FH-TUCK, s). 

- MIG. Cryptog. German., Austr. et Helvet. Exs. Fasc. XVIII (1904) no. 98, as St. coralloides (FH, 

MSC, PC, PR); Fasc. XLV (1928) no. 174, as St. coralloides (FH, PR). - MOUG. et NESTL. Stirpes Cryptog. 

Vogeso-Rhenanae Fasc. I (1810) no. 73, as St. paschale (FH-TUCK. Acc. to FREY, 1932, p. 149, in G and M 

it is unmixed, but in w contains also St. tomentosum FR.). - OLIV. Herb. Lich. Orne et Calvados (Lich. 

Exs.) Fasc. VII (1882) no. 313, as St. coralloides (DUKE. With some intermixture of another species, 

probably St. condensatum HOFFM.). - RABENH. Lich. Europ. Exs. Fasc. V (1856) no. 137, as St. coral- 

Iinum (s. In PR this number is referable to the var. flabellatum (FREY) GRUMM.); Fasc. VIII (1857) no. 

210, as St. corallinum (lapsu typogr. "cor Ilinum") (PR, s); Fasc. XXXV (1874) no. 939, as St. corafloides


B. dactylophyllum (PR, s). - RAs. Lich. Fenn. Exs. Fasc. I1 (1935) no. 55, as St. coralloides (ups). - REI- 

CHENB. et SCHUB. Lich. Exs. Fasc. VI (1826) no. 142, as St. paschale (PC). - ROUM. Lich. GUN. Exs. 

Cent. I(1879) no. 23, as St. corallinum (ups); Cent. VI (date ?) no. 560, as St. corallinum var.pulvinatum 

(UPS). - SCHADE, STOLLE et RIEHMER, Lich. Sax. Exs. Decas VIII (1925) no. 78, as St. coralloides 

(WELC. In BM this number is referable to var. occidentale (MAGN.) GRUMM.). - SCHAER. Lich. Helvet. 

Exs. Fasc. XI (1836) no. 261, as St. paschale y. corallinum (FH-TUCK, PC). - STENHAM. Lich. Suec. Exs. 

Fasc. I11 (1860) no. 82, as St. coralloides (BM, FH-TUCK, LD). - TREV. Lichenotheca Veneta, ser. I, Fasc. 

IV (1869) no. 142, as St. corallinum (FH); no. 143, as St. paschale (ups. Acc. to FREY, 1932, p. 149, this 

number in w is St. tomentosum FR.). -TUCK. Lich. Amer. Septentr. Exs. Fasc. IV (1854) no. 94, as St. 

corallinum (FH, L, TENN). -V&DA, Lich. Sel. Exs. Fasc. I1 (1960) no. 44 (LD). - WARTMANN et 

SCHENK, Schweizer. Kryptog. Cent. VI, Fasc. XI1 (1866) no. 552, as St. coralloides a. dactylophyllum 

(FH). - ZW. Lich. EXS. Fasc. XVIII (date ?) no 910, as St. coralFdes (PC, s, ups). 

Icon.: ANDERS 1928, PI. XIX, fig. 1, as St. coralloides. - CERN., NADv. et SERV. 1956, P1. VI, 

fig. 7, as St. coralloides. - FREY 1932, fig. 30a, b; P1.11, fig. 11, as St. coralloides. - GALLQE 1972, PI. 

72, figs. 555-561 ; P1. 73, figs. 562-565. - MAGN. 1929, PI. VI, fig. 2, as St. coralloides. - MIG. 1927- 

31, Lief. 268-277 (1928), P1. 72, figs. 4-9; PI. 73, fig. 1, as St. coralloides. - OKSN. 1968, fig. 148a. 

Mat. chim.: atranorin and stictic acid (the "Pseudopsoromsaure" of ZOPF, 1907), usually 

with smaller amounts of norstictic acid. 

Distrib. : an amphiatlantic species, occurring in Europe and eastwards into Russia at least as far 

as the Urals; Greenland; eastern Canada and eastern and central U. S. A. In Scandinavia it is distinctly 

southern in its distribution (ALMBORN, 1948), but has been found as far north as Angermanland in 

Sweden (DEGELIUS, 1932). Sweden, Norway, Finland, Denmark, Belgium, England, Scotland, France, 

Germany (W. and E.), Switzerland, Austria, Italy, Czechoslovakia, Poland, U. S. S. R. (Estonian SSR, 

Karelian SSR, Ukraine, Ural Mts., W. Siberia), S. W. Greenland, Canada (Quebec, Ontario, New 

Brunswick, Nova Scotia, Newfoundland), Miquelon Island, U. S. A. (New York State, Vermont, 

Massachusetts, New Hampshire, Maine, N. Carolina, Tennessee, Michigan). Unverified records from 

Ireland (PORTER, 1948), Rumania (CRETZOIU, 1933, 1939), U. S. S. R., Kola Peninsula, Murmansk 

Distr. (DOMBROVSKAYA, 1970), U. S. A., Pennsylvania (MOZINGO, 1948), in addition to records from 

Chile (JATTA, 1906) and Japan SAT^, 1965) which are almost certainly erroneous. The following 

records were checked by us and found to be erroneous: U. S. A., Alaska (EYERDAM, 1949,=St. intermedium 

(SAV.) MAGN.; HEUSSER, 1954,=St. glareosum (SAV.) MAGN.); China (MULLER ARC., 1893, 

HUE, 1898-1901 (1898), =St. myriocarpum TH. FR.) ; and Java (MERRILL, 191 3, =St. verruculigerum 

HUE). 

Remarks: St. dactylophyllum occupies a somewhat isolated position among its European 

congeners. It has some characteristics in common with sect. Denudata subsect. Botryoideum, 

with which it however probably has no close relationship, as it is geographically completely 

isolated from that E. Asiatic assemblage. 

Fo. expansum (MAGN.) GRUMM. 

Catalogus Lichenum Germaniae: 23 (1963). - Stereocaulon coralloides f. expansum MAGN. Goteborgs 

K. Vetensk.- o. Vitterh.-Samh. Handl. ser. 4,30 (7): 27/28 (1926); FREY, Rabenhorsts Krypt.-Fl. 9, Abt. 

IV, 1. Halfte: 194 (1932). 

Typus: Sweden, Vastergotland, par. Landvetter, Gokskulla, leg. C. STENHOLM, 1925 (UPS-MAGN, 

holotypus). 

Remarks: a minute, stunted, non-caespitose condition, with primary thallus absent or 

evanescent; perhaps merely a juvenile state of no taxonomic value. In any case rare; we 

know it from only two localities, both in Sweden.


Fo. subbotryosum (FREY) GRUMM. 

Catalogus Lichenum Germaniae: 24 (1963). - Stereocaulon coralloides f. subbotryosum FREY, Rabenhorsts 

Krypt.-F1. 9, Abt. IV, 1. Halfte: 194 (1932). 

Typus: Germany, Fichtelgebirge, leg. LAURER, 1871 (formerly in B; not seen by us; probably destroyed 

in World War 11). 

Icon.: FREY 1932, PI. 11, fig. 10 (the holotype), as St. coralloides f. subbotryosum. 

Remarks: an unusual form with massive woody pseudopodetia. Rare. Material agreeing 

very well with FREY'S description and illustration has been seen by us from Austria (Karnten). 

Fo. sessile LAMB (n. f.) 

Diagn.: Thallus primarius bene evolutus; pseudopodetia brevissima, usque 2.5 mm longa, apotheciis 

pileatis coronata. Apothecia partim quoque thallo primario sessilia. - St. pileato ACH. 

habitu persimile. 

Typus: Sweden, Vastergotland, par. Floby, leg. C. STENHOLM, 1896 (GB, holotypus). 

Remarks: the holotype was labelled by MAGNUSSON "Stereocaulon pileatum f. sessile H. 

MAGN.", and is one of the two specimens on which he founded the latter taxon; the other 

specimen we have selected as lectotype of St. pileatum f. sessile MAGN.,=S~. vesuvianum var. 

nodulosum f. sessile (MAGN.) LAMB; see p. 247. Known only from the type specimen. 

Var. flabellatum (FREY) GRUMM. 

Catalogus Lichenum Germaniae: 24 (1963). - Stereocaulon coralloides var. flabellatum FREY, Rabenhorsts 

Krypt.-FI. 9, Abt. IV, 1. Halfte: 196 (1932). 

Typus: Germany, Bohmer Wald, Bayrisch-Eisenstein, "am Aufstieg zur Hochbergplatte und zum 

Brennesattel, auf schattigen Felsen im Wald". More than one specimen mentioned, no holotype 

designated. Most of the type material was in herb. J. HILLMANN, which was destroyed in World War 

11. However, FREY retained one of the specimens (syntype), which we have seen; it should be designated 

as lectotype (lectotypif. nov.), and is from the Hochbergplatte, leg. J. HILLMANN, 1930 (FREY, 

herb. no. 1688). 

Exsicc.: FOLLM. Lich. EXS. Sel. Mus. Hist. Nat. Casselensi Fasc. VII (1975) no. 135, as St. dactylophyllum 

(KASSEL). - RABENH. Lich. Europ. EXS. Fasc. V (1856) no. 137, as St. corallinum (PR. In s 

this number represents the typical species). - SCHADE, STOLLE et RIEHMER, Lich. Sax. Exs. Decas XXVI 

(1927) no. 257, as St. coralloides (EM, PR. In BM it closely approaches var. occidentale (MAGN.) GRUMM.). 

- VENDRELY, Flora Sequania Exs. no. 251 (date ?), as St. coralloides (PC). 

Zcon.: FREY 1932, fig. 31, as St. coralloides var. flabellatum. 

Remarks: rather common and as widely distributed as the typical species. 

Var. occidentale (MAGN.) GRUMM. 

Catalogus Lichenum Germaniae: 24 (1963).1° - Stereocaulon coralloides var. occidentale MAGN. 


F1. 9, Abt. IV, 1. Halfte: 195 (1932). 

Typus: Sweden, 4 localities mentioned (in Bohuslan and Vastergotland), no holotype designated. 

As lectotype we select the one from Bohuslan, par. Dragsmark, leg. A. H. MAGNUSSON, 1915, and 

marked as "Type" in his herbarium (UPS-MAGN) (lectotypif. nov.). 

Facultative synonyms: Stereocaulon spissum NYL. ex HUE, Rev. Bot. Bull. Mens. 6: 192" (1888); 

emend. LAMB, Ann. Rept. Nat. Mus. Canada, 1952-53, Bull. no. 132: 272 (1954). - Stereocaulon coral- 

lo The combination was first made by LAMPE and KLEM. in Zeitschr. Mus. Hildesheim, n. F., Heft 

12: 71 (1958), but invalidly (basionym not designated).

I. MACKENZ~E LAMB: A Conspectus of Sfereocaulon 215 

loides var. spissum (NYL.) LAMB, loc. cit. - Sfereocaulon dacfylophyllum var. spissum (NYL.) LAMB ex 

GRUMM. Catalogus Lichenum Germaniae: 24 (1963). - Sfereocaulon spissum f. dactylinum FREY, 

Rabenhorsts Krypt.-F1. 9, Abt. IV, 1. Halfte: 141 (1932). 

Exsicc.: AND. Lich. EXS. Bohem. Bor. Fasc. VI (1933) no. 305, as Sf. fomenfosum (PR. Other 

examples of this number in PR contain also Sf. tomentosum FR. and another species, perhaps Sf. saxafile 

MAGN.). - BOHL. Lich. Brit. Fasc. I1 (1835) no. 14 (XIV), as Sf. paschale (FH). - MALME, Lich. Suec. 

Exs. Fasc. XXIII (1916) no. 559, as Sf. coralloides (PR, S, WIS. The example in s with slight intermixture 

of Sf. saxafile MAGN.). - MIG. Crypfog. German., Ausfr. ef Helvef. Exs. Fasc. LXII (1933) no. 342, as 

Sf. coralloides (FH, OC, PR, U. In FH there is an intermixture of Sf. saxafile MAGN.). - SCHADE, STOLLE 

et RIEHMER, Lich. Sax. EXS. Decas VIII (1925) no. 78, as Sf. coralloides (BM. In WELC this number is 

referable to the typical species); Decas XV (1925) no. 148, as Sf. coralloides (BM); Decas XXVI (1 927) 

no. 258, as Sf. coralloides (BM); no. 260 pr. p., as Sf. fomenfosum (BM. Mixed with Sf. saxafile MAGN.); 

Decas XXXVII (1927) no. 363, as Sf. coralloides (BM), - V~ZDA, Lich. Sel. Exs. Fasc. XXXV (1970) 

no. 866, as Sf. dacfylophyllum (LD). - Zw. Lich. Exs. Fasc. XIX (date ?) no. 997, as Sf. pileafum, 

spissum (H-NYL, lecfofypus; PC, isolectotypus). 

Icon.: FREY 1932, fig. 21, as Sf. spissum; fig. 30 c, d, as Sf. coralloides var. occidentale; PI. 11, fig. 

9, as Sf. spissum. - ULLRICH et KLEM. 1960, fig. 1112.9 (field photograph). 

Remarks: very common and as widespread as the typical species. 

Var. pygmaeum (FREY) GRUMM. 

Catalogus Lichenum Germaniae: 24 (1963). - Sfereocaulon coralloides var. pygmaeum FREY, Rabenhorsts 

Krypt.-FI. 9, Abt. IV, 1. Halfte: 197 (1932). 

Typus: Germany, Harz, Ilsenburg, between Eckerkrug and Rabenklippen, "auf stark verwittertem 

Granit", leg. J. HILLMANN, no date given (herb. HILLMANN, holotypus; destroyed in World War I1 and 

not seen by us). 

Icon.: FREY 1932, fig. 32, as Sf. coralloides var.pygmaeum. 

Remarks: characterized by very dwarf pseudopodetia forming a sward over the stone; 

phyllocladia as in var. flabellatum. FREY, loc. cit., admits the possibility of it being merely 

a juvenile condition of var. flabellatum or var. occidentale. We have seen only one specimen 

agreeing with the description, from U.S.S.R., Ural Mts., Sverdlovsk region. 

1 1. Stereocaulon depreaultii DEL. 

ex NYL. Synopsis Methodica Lichenum, 1 (2): 249 (1860); ASAH. J. Jap. Bot. 35: 292 (1960); YOSHIM. 

Lichen Flora of Japan in Colour: 176 (1974). - Sfereocaulon paschale subsp. depreaulfii (DEL.) LAMB, 

Ann. Rep. Nat. Mus. Canada, 1952-53, Bull. no. 132: 273 (1954). 

Typus: Canada, Newfoundland, no exact locality given, leg. DESPREAUX (date ?) (CH-NYL, no. 39814, 

holotypus). What appear to be isotype specimens are present in ASAH (with date 1828), FH, PC (with 

date 1827), RB (as from "Petit-nord de Terre-neuve", the old name for the Great Northern Peninsula) 

and s. 

Exsicc. : ASAH. Lich. Japon. EXS. Fasc. VI (1959) no. 294 (ASAH, FH, LD, UPS). - KUROK. Lich. 

Rarior. el Crif. Exs. Fasc. 111 (1971) no. 139 (ASAH, FH, LD, UPS). 

Icon.: ASAH. 1960, fig. 4A. -YOSHIM. 1974, fig. 76i; PI. 33, fig. 325. 

Mat. chim. : atranorin and lobaric acid. 

Disfrib.: Canada (Newfoundland and possibly Nova Scotia; see below) and Japan. Possibly 

elsewhere in eastern N. America, but these finds rather doubtful. It is an example of the disjunct type 

of eastern N. American and Asiatic distribution found in certain mosses (IWATSUKI, 1958) and a few 

lichens, e. g. Sfereocaulon fennesseense MAGN. and Lecanora perconcinna HUE (syn. L. microbola LAMB, 

1954, p. 29111). See also W. L. CULBERSON, 1972. 

Identity established by examination of the holotype of L. perconcinna in PC-HUE, 1966.

21 6 Journ. Hattori Bot. Lab. No. 43 1977 

Remarks: there is another specimen of the same species in H-NYL (no. 39980) labelled 

"Stereocaulon Despreauxii Del.", on which NYLANDER has written "alpinum Halifax Jardin". 

Halifax is a town in Canada, Nova Scotia, and Jardin perhaps the name of the collector. The 

holotype specimen also bears the name "tortuosum Delise" in an older handwriting, perhaps 

that of DELISE himself, and a specimen of the same species (perhaps isotype material) in FH 

bears the same name with the addition of "Delise in herb. et monogr. ined." The epithet 

tortuosum remained a herbarium name (nomen nudum) until 1900, when it was published by 

HULTING (1900, p. 17), who validated it by a short diagnosis supplied by NYLANDER, and 

quoted Stereocaulon subintricans NYL. as a synonym. It seems that the Swedish specimen 

to which HULTING attached the name St. tortuosum (in GB) must be regarded as the holotype, 

rather than any of the DELISE material. It was studied both by MAGNUSSON (1926, p. 83) 

and by the present author, neither of whom could identify it with certainty. It also seems 

that the authority for St. tortuosum should be NYL. ex HUL~NG rather than DEL., because 

DELISE used the name in reference to St. depreaultii. See also under St. tortuosum and St. 

subintricans in Appendix 2 (p. 321). 

12. Stereocaulon depressum (FREY) LAMB 

ex POELT, Be~timmung~~~hlii~~e1 Europaischer Flechten: 645 (1969); DAHL et KROG, Macrolichens of 

Denmark, Finland, Norway and Sweden: 139 (1973). - Stereocaulon fastigiatum f. depressum MAGN. 

Goteborgs K. Vetensk.- o. Vitterh.-Samh. Handl. ser. 4, 30 (7): 36 (1926) (nom. inval., provisor.). - 

Stereocaulon botryosum f. depressum FREY, Rabenhorsts Krypt.-F1. 9, Abt. IV, 1. Halfte: 126 (1932) 

(val. pub].). 

Typus: Norway, Nordland, Umbukten, Krabbfjall, altit. ca. 600 m s. m., leg. A. H. MAGNUSSON, 

1924, no. 9040 (UPS-MAGN, holotypus). 

Facultative synonyms: Stereocaulon fastigiatum f. confluens MAGN. Gijteborgs K. Vetensk.- o. 

Vitterh.-Samh. Handl. ser. 4, 30 (7): 35 (1926). - Stereocaulon botryosum f. confluens (MAGN.) FREY, 


Exsicc.: MALME, Lich. Suec. Exs. Fasc. XL (1926) no. 979A, pr. p., as St. fastigiatum (LD. In s 

this number is St. botryosum ACH. em. FREY). 

Mat. chim.: atranorin, often with traces of some unidentified substances, including a 

fatty acid. (Porphyrilic and lobaric acids absent.) 

Distrib. : Fennoscandia, in northern and alpine locations: Sweden, Norway, Finland. Also found 

in E. Greenland and Jan Mayen. 

Remarks: closely related to St. groenlandicum (DAHL) LAMB, which has a more complex 

chemistry. MAGNUSSON'S original publication of the taxon is regarded as invalid (provisional 

name, cfr. STAFLEU et al., 1972, Art. 34), and so FREY is taken to be the original author. 

13. Stereocaulon esterhuysenae LAMB 

Lilloa, 26: 417 (1953). 

Typus: South Africa, Cape Province, Tulbagh Division, Great Winterhoek, altit. 1500 m s, m., leg. 

E. ESTERHUYSEN, 1951, no. 19862 pr. p. (CAN, holotypus; BOL, isotypus). 

Icon.: LAMB 1953, PI. 11, fig. 6 (the holotype). 

Mat. chim.: atranorin, stictic acid, norstictic acid (the latter in low concentration). 

Distrib. : South Africa. 

Remarks: the type material is a juvenile, dorsiventral, flattened condition; mature plants 

are erect, less or not dorsiventral, with rigid and ligneous, firmly attached pseudopodetia, 

somewhat resembling St. botryosum ACH. em. FREY in habitus. May be difficult to distinguish


from St. glabrum (MULL. ARG.) VAIN., but has a different morphology in the juvenile state. 

14. Stereocaulon evolutum GRAEWE 

ex TH. FR. Bot. Not.: 181 (1865); MAGN. Goteborgs K. Vetensk.- o. Vitterh.-Samh. Handl. ser. 4, 

30 (:7) 30 (1926); FREY, Rabenhorsts Krypt.-F1. 9, Abt. IV, 1. Halfte: 142 (1932). - Stereocaulon 

evolutum [var.] a. typicum TH. FR. Lichenographia Scandinavica, Pars prima: 45 (1871). 

Typus: "I vestra delen af Skaraborgs Ian", Vastergotland, Frammestad, leg. P. H. F. GRAEWE, 

1864 (UPS-TH FR, holotypus or isotypus). 

Facultative synonyms: Stereocaulon evolutum f. planum MAGN. Goteborgs K. Vetensk.- o. Vitterh.- 

Samh. Handl. ser. 4, 30 (7): 32 (1926). - Stereocaulon lusitanicum MAGN. ex TAV. Broteria, 16 (43): 

147 (1947). 

Exsicc.: ABB. Lich. Armor. Spect. Exs. Fasc. V (1932) no. 84, as St. paschale (LD, UPS). AND. 

Lich. Exs. Bohem. Bor. Fasc. I1 (1930) no. 87, pr. p., as St. globosus (PR. Mixed with St. tomentosum 

var. compactum FREY). - CLAUD. et HARM. Lich. Gall. praecip. Exs. Fasc. X (1908) no. 479, as St. 

paschale (FH). - CROMB. Lich. Brit. Exs. Cent. I1 (1877) no. 120 (ups. But in BM it is St. dactylophyNum 

FLORKE). - DEL. Lich. de France, Fasc. I (1828) no. 17, as St. dactylophyNum (FH). - FR. Lich. Suec. 

Exs. Fasc. I11 (1824) no. 89, pr. p., as St. paschale B. conglomeratum (FH. But in s it is St. paschale (L.) 

HOFFM., and in LD and ups a mixture of St. evolutum and St. paschale). - HAMBERG, Kryptogam- 

Herbarium (1895) no. 50, as St. paschale (UPS). - HAV. Lich. Norveg. Occid. Exs. Fasc. I1 (1913) no. 

33 (FH, LE). - Krypt. Exs. Vindob. Cent. XXI (1913) no. 2065 (FH, LE, PR, US, WELC). - MAGN. Lich. 

Sel. Scand. Exs. Fasc. X (1937) no. 237, as St. evolutum f. planum (DEGEL, LE, PR). - MALME, Lich. 

Suec. Exs. Fasc. XXVIII (date ?) no. 677 (WIS). - OLIV. Herb. Lich. Orne et Calvados (Lich. Exs.) 

Fasc. VII (1882) no. 314, as St. coralloides var. conglomeratum (DUKE); Fasc. IX (1884) no. 407, as St. 

pileaturn (DUKE). - RABENH. Lich. Europ. Exs. Fasc. XXXII (1870) no. 858 (FH-TUCK). - ROUM. 

Lich. Gall. Exs. Cent. V (1883) no. 423, as St. coralloides var. conglomeratum (UPS). - SAMP. Lich. 

Portugal, Fasc. I11 (1923) no. 286, as St. coralloides (LD). - TAV. Lich. Lusit. Sel. Exs. Fasc. I (1951) 

no. 12, as St. lusitanicum (LD, TENN). 

Icon.: ALVIN et KERSH. 1963, PI. 54, lower fig. - BRIGHTM. et NICHOLSON 1966, figs. 8, 8A. - 

FREY 1932, fig. 19 d-f; PI. 11, fig. 7. - GALL. 1913, figs. 120-123. - KERSH. 1960, fig. 4. - Oz. et CLAUZ. 

1970, fig. 391 A, B. - TAV. 1947, fig. 1, as St. lusitanicum. 

Mat. chim.: atranorin and lobaric acid. 

Distrib.: a European suboceanic species, occurring mainly along the coastal regions of western 

Europe from Scandinavia and the Faeroe Islands in the north to Spain, Portugal, Madeira and the 

Canary Islands in the south; penetrating eastwards in Europe as far as Finland, central Germany and 

Czechoslovakia. Sweden, Norway, Denmark, Faeroe Islands, Finland, Belgium, England, Scotland, 

Ireland, France, Spain, Portugal, W. Germany, Czechoslovakia, Madeira, Canary Islands. Unverified 

records: Italy (JATTA, 1900; 1909-11); U. S. S. R., Murmansk Distr. (DOMBROVSKAYA, 1970), Estonian 

SSR (TRASS, 1958). Transbaikalia (RASSADINA, 1936). Erroneous record: Canada, Quebec (LEPAGE, 

1958, =St. saxatile MAGN.). 

Remarks: when sterile, as is commonly the case, St. evolutum may be rather difficult to 

distinguish from St. saxatile MAGN., but the pseudopodetia are stronger and more woody 

and perfectly glabrous, and the phyllocladia are digitate-squamulose, many of them forming 

phyllocladioid branchlets, and often of a characteristic caesious-gray color. If apothecia 

are present, the short broad spores are diagnostic. The pycnoconidia also seem to be different 

in the two species. 

Fo. Iaxum (FREY) LAMB 

ex POELT, Be~timmung~schliisse1 Europaischer Flechten: 643 (1969). - Stereocaulon spissum var. laxum


FREY, Rabenhorsts Krypt.-F1. 9, Abt. IV, 1. Halfte: 141 (1932). - Stereocaulon evolutum var. Iaxum 

(FREY) LAMB ex GRWM. Catalogus Lichenum Germaniae: 24 (1963). 

Typus: Germany, Prov. Hannover, Giersfeld, "Granitblocke des Hunengrabes", leg. H. SANDSTEDE 

1888 (M-ARN, holotypus; isotypes in herb. SANDSTEDE and in B, the latter destroyed in World War 11). 

Exsicc. : HAV. Lich. Exs. Norveg. Fasc. VII (1905) no. 264, as St. evolutum (BG). 

Icon.: FREY 1932, PI. I, fig. 6, as St. spissum var. laxum (holotype or isotype specimen). 

Remarks: characterized by laxly caespitose pseudopodetia. Fairly common in western 

Europe. 

15. Stereocaulon glabrum (MULL. ARG.) VAIN. 

Lichens, in: Res. Voy. Belgica, Botanique: 16 (1903); D. C. LINDS. Brit. Antarct. Survey Sci. Repts. 

89: 50 (1974). - Stereocaulon alpinum var. glabrum MULL. ARC. Lichens, in: Mission Scientifique du 

Cap Horn, 1882-1883, 5, Botanique: 151 (1889). 

Typus: Chile, Magallanes, Hardy Peninsula, Orange Bay, leg. HYADES (G-~LL, holotypus). 

Facultative synonyms: Stereocaulon patagonicum LAMB, Farlowia, 4: 454 (1955). - Stereocaulon 

colensoi var. reagens RAS. Ann. Bot. Soc. Zool. Bot. Fenn. Vanamo, 2 (1): 24 (1932). - Pilophoron 

colensoi var. reagens (RAs.) ZAHLBR. Catalogus Lichenum Universalis, 10: 381 (1939). - Stereocaulon 

patagonicum f. reagens (RAs.) LAMB, An. Parques Nac. [Buenos Aires], 1958, 7: 99 (1959). -Stereocaulon 

patagonicum f. subirregulare LAMB, Farlowia, 4: 456 (1955). 

Icon.: ASAH. 1967, fig. 8. -LAMB 1955, fig. 21, as St. patagonicum; fig. 22, as St. patugonicum f. 

subirregulare. - D. C. LINDS. 1974, PI. 111, fig. f. -VAIN. 1903, PI. 11, fig. 8. 

Mat. chim. : the species occurs in two chemical strains. 

Strain I, with atranorin, stictic acid and norstictic acid (the latter in low concentration). 

The typical and common strain. 

Strain II, with atranorin, norstictic acid and connorstictic acid (stictic acid absent). One 

specimen only of this strain seen: Chile, Prov. Malleco, Dept. Angol, Parque Nacional de 

Nahuelbuta, leg. M. MAHU, 1971 (no. 2320) (FH, USGO-MAHU). 

Distrib. : Chile (Magallanes, Valdivia, Chiloe, Ayskn, Bio-Bio, Osorno, Malleco), Juan Fernandez, 

Argentina (Tierra del Fuego, Santa Cruz, Chubut, Rio Negro, Neuqukn,) South Georgia, W. Antarctica 

(South Shetlands) and Tristan da Cunha. 

Remarks: specimens from the southernmost parts of S. America (including the type 

specimen), and also from alpine locations further north, are mostly very reduced and depauperate; 

the optimally developed state of the species occurs in Argentine and Chilean 

Patagoaia, at comparatively low elevations (St. patagonicum LAMB). The taxa reagens RA~. 

and subirregulare LAMB represent states somewhat less depauperated than the type material 

of the species. The isolated occurrence in Tristan da Cunha is paralleled by the similar 

distribution of another S. American species, St. implexum TH. FR., but in that case the insular 

population is chemically different. 

Fo. flabellans (LAMB) LAMB (n. comb.) 

Basionym: Stereocaulonpaschale var. alpinum f.flobellans LAMB, Farlowia, 4: 457 (1955). 

Typus: Argentina, Patagonia, Rio Negro, summit of Mt. Rigi near Lago Frias, altit. ca. 1750 m s. 

m., leg. I. M. LAMB, 1950 (no. 5787) (CAN, holotypus; FH, isotypus). 

Icon. : LAMB 1955, fig. 23, as St. paschale var. alpinum f. flabellans. 

Remarks: known only from the type locality. 

16. Stereocaulon glareosum (SAV.) MAGN. 

Goteborgs K. Vetensk.- o. Vitterh.-Samh. Handl. ser. 4, 30 (7): 60 (1926); FREY, Rabenhorsts Krypt.-


F1. 9, Abt. IV, 1. Halfte: 170 (1932); POELT, Bestimmungsschli~ssel Europaischer Flechten: 641, 645 

(1969). - Stereocaulon tomentosum f. glareosum SAV. Bull. Jard. Imp. Bot. Pierre le Grand (Izv. Imp. 

Bot. Sada Petra Velikago), 14: 121 (1914). 

Typus: U. S. S. R., Kamtchatka, Sczapina River, leg. V. P. SAVICZ, 1909 (no. 6186 [115]) (LE, 

holotypus). 

Facultative synonyms: Stereocaulon paschale f. ramuliferum NYL. ex STIZENB. Flora, 48: 93 (1865) 

(nomen nudum), NYL. Lichenes Lapponiae Orientalis: 111 (1866)12 (cum descript.). - Stereocaulon 

glareosum f. congestum MAGN. Goteborgs K. Vetensk.- o. Vitterh.-Samh. Handl. ser. 4, 30 (7): 63 

(1926). 

Exsicc. : FELLM. Lich. Arct. no. 47 (1865), as St. paschale f. ramuliferum (H, H-NYL). - HAV. Lich. 

Exs. Norveg. no. 684 (1949) (BG). - Krypt. Exs. Vindob. Cent. I1 (1896) no. 152b, as St. alpinum (the 

specimen from Finnish Lapland) (BM, COLO, FH, PR, US). - Lichenotheca Fenn. Fasc. XXXIX (1956) 

no. 963, as St. saxatile (FH). - MAGN. Lich. Sel. Scand. Exs. Fasc. IX (1935) no. 211 (DEGEL, FH, LE, 

PR). - POELT et al. Lich. Alpium Fasc. VI (1961) no. 111 (czu, LD); Fasc. VIII (1961) no. 147 (GZU, 

LD). - RAS. Lich. Fenn. Exs. Fasc. IV (1936) no. 199 (CAN, UPS). 

Icon.: DAHL et KROG 1973, fig. 43. - FREY 1932, fig. 27; 1959, fig. 12; 1969, fig. 5 (opposite p. 

52). - HALE 1969, fig. 385. - KERSH. 1960, fig. 12. 


Distrib.: boreal-arctic, alpine-subalpine, circumpolar, extending across Europe, Asia and N. 

America, also in the northern part of S. America. Sweden, Norway, Finland, Switzerland, Austria, 

Italy, France, Scotland, U. S. S. R. (Estonian SSR, Kola Peninsula, Ural Mts., Altai, E. Siberia, Transbaikalia, 

Kamtchatka, Extreme Orient), Mongolia, Kashmir, Nepal, Spitsbergen, Iceland, Greenland, 

Canada (Quebec, Labrador, Alberta, British Columbia, N. W. Territ., Yukon Territ.), U. S.A. (Colorado, 

Montana, Wyoming, Oregon, Washington, Alaska), Mexico, Colombia, Ecuador, Peru. 

Remarks: the cephalodia of this species are very characteristic. 

Var. brachyphylloides LAMB 

ex LLANO, J. Wash. Acad. Sci. 41: 196 (1951). 

Typus: U. S. A., Alaska, Brooks Range, Anaktuvuk Pass, altit. 1000 m s. m., leg. G. A. LLANO 

and N. WEBER, 1949 (no. 527) (us, holotypus; CAN, isotypus). 

Exsicc. : Lich. Exs. Univ. Colorado Mus. Fasc. I1 (1963) no. 66, as St. glareosum (LD, LE). 

Remarks: seems to be as widely distributed as the typical species. 

17. Stereocaulon glaucescens TUCK. 

Boston J. Nat. Hist. 3: 302 (1841). - Stereocaulon vesuvianum var. glaucescens (TUCK.) LAMB ex 

HALE, Amer. Midl. Naturalist, 51: 245 (1954) (comb. inval.). 

Typus: U. S. A., New Hampshire, The Notch, leg E. TUCKERMAN, 1839 (FH-TUCK, holotypus). 

Mat. chim. : atranorin, lobaric acid, and an unidentified compound. 

Distrib.: N. E. North America, mostly in mountainous or alpine situations. The typical species 

occurs in New Hampshire, Vermont and QuCbec; the variety caespitosulum extends over a wider area 

including New York State, Massachusetts, Maine, Ontario, Nova Scotia and Newfoundland. Some 

specimens seen from Michigan and Minnesota strongly resemble var. caespitosulum, but their identity 

is not fully established. 

Remarks: although TUCKERMAN himself (1845, p. 101; 1847, p. 238) reduced this species 

to synonymy with St. denudatum (=St. vesuvianum var. nodulosum (WALLR.) LAMB), it is quite 

different from that species, belonging to sect. Stereocaulon and obviously closely related 

to St. saxatile MAGN. The resemblance to certain states of St. vesuvianum PERS. (and even 

l2 Reprinted in Not. Sallsk. Fauna FI. Fenn. Forh. 8: 101-192 (1882).


more so to St. vanoyei DUVIGN.) is superficial and coincidental. When typically developed, 

it is noteworthy for its rather large, apically produced, convex and pileate-squamulose phyllocladia, 

but the var. caespitosulum more resembles St. evolutum GRAEWE and St. saxatile 

MAGN.; from the latter it is distinguished by its strongly ligneous, firmly affixed, non-tomentose 

pseudopodetia. The spores of var. caespitosulum are 3-septate, 20-33 x34 p; the typical 

species has not been found with apothecia. 

Var. caespitosulum (NYL.) LAMB (n. comb.)13 

Basionym: Stereocaulon denudatum var. caespitosulum NYL. Synopsis Methodica Lichenum, 1 (2): 

247, footnote (1860). - Stereocaulon depreaultii var. caespitosulum (NYL.) LAMB ex THOMS. Bryologist, 

57: 284 (1954) (comb. inval., basionym not cited), ex IMSH. Bryologist, 60: 223 (1957) (val.). -Stereocaulonpaschale 

subsp. depreaultii var. caespitosulum (NYL.) LAMB, Ann. Rep. Nat. Mus. Canada, 1952- 

53, Bull. no. 132: 274 (1954). 

Typus: U. S. A., "ad terram in montibus Novae Angliae", leg. E. TUCKERMAN (date ?) (TUCK. 

Lich. Amer. Septentr. Exs. no. 114 in H-NYL, holotypus. The isotypes seen in various herbaria are 

identical). 

Exsicc.: Lich. Canad. Exs. no. 64 (1970) (CAN). - MERR. Lich. EXS. Ser. 11, Fasc. I (1925) no. 7, 

as St. paschale var. conglomerata (c, COLO, FH, UPS-MAGN, US, WIS). - TUCK. Lich. Amer. Septentr. Exs. 

Fasc. V (1854) no. 114, as St. denudatum (FH, H-NYL, TENN, US, isotype material). 

18. Stereocaulon grande (MAGN.) MAGN. 

ex NILSS. Ark. f. Bot. 24A (3): 71 (1931); POELT, Bestimmungsschliissel Europaischer Flechten: 647 

(1969); DAHL et KROG, Macrolichens of Denmark, Finland, Norway and Sweden: 141 (1973). - 

Stereocaulon paschale var. grande MAGN. Goteborgs K. Vetensk.- o. Vitterh.-Samh. Handl. ser. 4, 30 

(7): 49 (1926). - Stereocaulon alpinum var. grande (MAGN.) FREY, Bull. Murith. Soc. Valais Sci. Nat. 

54: 85 (1937). 

Typus: several localities in Sweden mentioned, no holotype designated. As lectotype we have 

selected the specimen from Jamtland, par. Kall, Sundet, leg. A. H. MAGNUSSON, 1914 (UPS-MAGN, 

lectotypus, lectotypif. nov.). It was marked as type specimen by MAGNUSSON himself. 

Exsicc.: Lichenotheca Fenn. Fasc. XXXI (1953) no. 758, as St. tomentosum (FH); Fasc. XLlV 

(1958) no. 1080, as St. paschale (FH). - MAGN. Lich. Sel. Scand. Exs. Fasc. X (1937) no. 238 (PR, s). - 

Rds. Lich. Fenn. Exs. Fasc. VIII (1939) no. 396, as St. squamescens (GB, LD). - VEZDA, Lich. Sel. Exs. 

Fasc. XIX (1966) no. 465, as St. paschale (FH). 

Icon.: FREY 1932, PI. 111, fig. 4 pr. p. (right-hand specimen only).14 


Distrib.: boreal-circumpolar, apparently not occurring in central Europe. Sweden, Norway, 

Finland, U. S. S. R. (Kola Peninsula, Sayan Mts., Kamtchatka), W. Greenland, Canada (Ontario, 

QuBbec, Nova Scotia, Newfoundland, Labrador, Manitoba, Saskatchewan, British Columbia, N. W. 

Territ.), U. S. A. (Maine, New Hampshire, Oregon, Alaska). 

Remarks: a rather unsatisfactory species from the taxonomic point of view, being intermediate 

between St. paschale (L.) HOFFM. and St. alpinum var. erectum FREY, and partaking 

of the characters of both. Only when well and typically developed is it clearly distinguishable. 

Its characteristic features, in approximate order of constancy and reliability, are: (1) phyllo- 

l3 This combination made on label of Lich. Canad. Exs. no. 64 (1970), but invalid (basionym not 

cited). 

l4 The description given by FREY, 1932, p. 155, is not reliable for this species, as it refers in part to 

St. alpinum var. erectum FREY. His illustrations loc. cit., apart from the one listed above, which is of 

part of the type (lectotype) material of St. grande, depict specimens of St. alpinum var. erectum.


cladia mostly of the paschale-type (but somewhat coarser); (2) cephalodia pulvinate to subglobose, 

smooth or verruculose-divided, usually pale br'own, containing either Arostoc or 

Stigonema; (3) apothecia large, terminal, often subdividing; (4) usually of tall growth, 4- 

8 cm high. MAGNUSSON (1946, p. 92) states that St. grande has longer conidia than St. 

paschale. AHTI (1964, p. 24) points out that St. grande seems to be ecologically distinguished 

from St. paschale and St. alpinum. While undoubtedly more than a mere ecotypic modification, 

St. grande is perhaps not entitled to the rank of a proper species. 

19. Stereocaulon groenlandicum (DAHL) LAMB 

Occas. Papers Farlow Herb. Crypt. Bot. no. 5: 10 (1973). - Stereocaulon rivulorum var. groenlandicum 

DAHL, Meddel. Gronland, 150 (2): 117 (1950). 

Typus: S. W. Greenland, Julianehaab Distr., Igalikofjord, Igdleriigssalik, altit. 700-800 m s. m., 

leg. E. DAHL, 1937 (0, holotypus; c, us, isotypes). 

Icon.: DAHL 1950, PI. 11, fig. 2, as St. rivulorum var. groenlandicum. -LAMB 1973, figs. 9, 10a, b, 

c, 11. 

Mat. chim.: the full chemical spectrum (as in the type material) consists of atranorin, 

perlatolic acid, anziaic acid, miriquidic acid, and some unidentified substances. Miriquidic 

acid is obviously an accessory substance, as many of the specimens seen lack it. A strongly 

deficient phase with only atranorin certainly identifiable is also of moderately common occurrence, 

which makes distinction from St. depressum (FREY) LAMB difficult. 

Distrib. : boreal-arctic, probably circumpolar. Rare in N. Scandinavia: Sweden (Ase~e Lappmark) 

and Norway (Nordland, Nord-Trondelag, Tromso, Sogn og Fjordane). More common in the arctic 

and subarctic regions: Spitsbergen, Greenland, Canada (N. W. Territ.), U. S. A. (Alaska). 

Remarks: most closely related to the mainly Fennoscandian species St. depressum (FREY) 

LAMB, which always contains atranorin only. However, as mentioned above, a deficient 

phase of St. groenlandicum with atranorin only is sometimes found, and the question arises 

as to whether the two species may be synonymous, although St. depressum is usually distinguishable 

in its morphology (more flattened and dorsiventral growth-habitus). In the 

meantime, it seems best to treat them as separate species. Smaller, more stunted specimens 

of St. groenlandicum not clearly showing the characteristic ligneous development of the 

pseudopodetia are morphologically hardly distinguishable from St. rivulorum MAGN. 

20. Stereocaulon incrustatum FLORKE 

Deutsche Lichenen, Lief. IV: 12 (1819); TH. FR. Monographia Stereocaulorum et Pilophororum: 360 

(1858); HUE, NOUV. Arch. Mus. Hist. Nat. ser. 3, 10: 255 (1898), as St. tomentosum var. incrustatum; 

MAGN. Gijteborgs K. Vetensk.- o. Vitterhdamh. Handl. ser. 4, 30 (7): 59 (1926); FREY, Rabenhorsts 

Krypt.-FI. 9, Abt. IV, 1. Halfte: 172 (1932); RAs. Suomen Jakalakasvio: 68 (1951); YOSHIM. Lichen 

Flora of Japan in Colour: 177 (1974). - Stereocaulon tomentosum var. incrustatum (FLORKE) SCHAER. 

Lichenum Helveticorum Spicilegium, Sect. VI: 276 (1833). - Stereocaulon tomentosum [subsp.] * 

incrustatum (FLORKE) NYL. Synopsis Methodica Lichenum, 1 (2): 245 (1860). 

Typus: Germany, Schmargendorf in Grunewald near Berlin, "an der Erde in den Tannenwaldungen", 

leg. H. G. FLORKE, 1816 (UPS-TH FR, S, isotypes; the holotype in herb. FLORKE not seen by us). 

The material distributed with the original description in FLORKE'S exsiccat Deutsch. Lich. no. 77 bears 

no indication of locality, but is probably also isotype material. FLORKE, loc. cit., also mentioned 

specimens from Rostock, so lectotypification on the Schmargendorf material is necessary (lectotypif. 

nov.). 

Facultative synonyms: Patellaria tomentosa /3. erecta WALLR. Flora Cryptogamica Germaniae,


Pars prior: 439 (1831). -Stereocaulon abduanum ANZI, Comment. Soc. Crittogamolog. Ital. 2 (1) 5 

(1864). - Stereocaulon tomentosum var. abduanum (ANZI) OLIV. Mem. Soc. Sci. Nat. Cherbourg, 36: 

162 (1906-07). - Stereocaulon incrustatum var. abduanum (ANZI) FREY, Rabenhorsts Krypt.-F1. 9, 

Abt. IV, 1. Halfte: 172 (1932). - Stereocaulon incrustatum var. elatum OKSN. Zhurn. Inst. Bot. Vseukrakjins'k. 

Akad. Nauk (Journ. Bot. Acad. Sci. RSS Ukraine), 1 (2): 314 (1940). 

Exsicc.: ANZI, Lich. Rar. Langob. Exs. Fasc. I (1861) no. 14 (FH-TUCK, s). - ARN. Lich. EXS. 

(Lich. Jurae) no. 1565 (1892) (BM, FH, PC). - BARTH, Herbarium Transsylvanicum Fasc. I (1873) no. 4, 

as St. tomentosum (M). - BARTLING et HAMPE, Veget. Cellul. German. Septentr., Lich. Decas I1 (date ?) 

no. 17, pr. p., as St. tomentosum (M. Mixed with St. paschale (L.) HOFFM.). - Erb. Crittog. Ztal., ed. 

DE NOTARIS, CESATI, BAGLIETTO etc. Fasc. IV (1859) no. 194 (FH). - Erb. Crittog. Ztal., publ. Soc. 

Critlogamolog. Ztal., Ser. 11, Fasc. VII (1870) no. 319 (FH). - FLORKE, Deutsch. Lich. Fasc. IV (1819) 

no. 77 (probably isotype material) (UPS). -Flora Romaniae Exs. Cent. XV (date ?) no. 1409, as St. 

corraloides [sic] (GB, s, us). - FUNCK, Cryptog. Gewrichse des Fichtelgebirg's, Ed. 11, Heft 30 (1824) no. 

624 (BM, FH). - HEPP, Flechten Europas, Band VI, Heft 11 (1857) no. 301 (FH-TUCK, PC, s). - MAGN. 

Lich. Sel. Scand. Exs. Fasc. VII (1933) no. 156 (DEGEL, PR); Fasc. XI1 (1939) no. 287 (ups). - RABENH. 

Lich. Europ. Exs. Fasc. V (1856) no. 136 (FH-TUCK, GB, PR, s); Fasc. XVI (1859) no. 455 (FH-TUCK, PR, 

s). - REICHENB. et SCHUB. Lich. EXS. Fasc. VI (1826) no. 141 (PC). - ROUM. Lich. GUN. Exs. Cent. I1 

(1880) no. 162 (ups. FREY, 1932, p. 162, states that the example in M-ARN is St. alpinum var. gracilentum 

(TH. FR.) MAGN.). - TOBOL. Lichenotheca Polon. Fasc. IX (1957) no. 184 (FH, LD, S. Not typical). - 

WARTMANN et SCHENK, Schweizer. Kryptog. Cent. VII, Fasc. XIV (1869) no. 652 (FH). - VBZDA, Lich. 

Sel. Exs. Fasc. LII (1975) no. 1282 (BRNU); Fasc. LIV (1975) no. 1340 (BRNU). - ASAH. Lich. Japon. 

Exs. Fasc. VI (1959) no. 295 (ups, us). - Plantae Varsav. Exs. Fasc. I1 (1964) no. 39 (LD, s). 

Icon. : MIG. 1927-31 (1928), PI. 72c, figs. 6, 7; PI. 73, fig. 2. - FREY 1932, fig. 28a; PI. 111, fig. 9; 

1959, fig. 11; 1969, fig. 6 (opposite p. 53). - AHLN. 1932, fig. 1. - KERSH. 1960, fig. 11. -0~s~. 1968, fig. 146. - Oz. et CLAUZ. 1970, fig. 395 D, E. - YOSHIM. 1974, fig. 77c; PI. 33, fig. 330. - PASICH 

1973, figs. 1, 2 (plants with abnormal production of soralia). 

Mat. chim.: atranorin and lobaric acid, or quite frequently in a deficient phase containing 

atranorin only. By ASAHINA (1934, p. 60) and DWIGNEAUD (1942, p. 82) erroneously stated 

to contain stictic acid, this apparently derived from the statement by ZOPF (1907, p. 404) that 

"Pseudopsoromsaure" (=stictic acid) is present. 

Distrib.: of mainly continental distribution in Europe, Asia and N. America. Sweden, Denmark, 

Norway, Finland, France, Germany, Switzerland, Austria, Italy, Czechoslovakia, Hungary, Poland, 

Rumania, U. S. S. R. (Estonian SSR1q Ukraine, central and eastern Siberia, Transbaikalia, Kamtchatka, 

Extreme Orient), Mongolian People's Republic, U. S. A. (Colorado, Alaska), Korea, Japan. 

Unverified records from Belgium (DE WILDEMAN and DURAND, 1898-1907) and U. S. S. R., Kola 

Peninsula and Murmansk District (TRASS, MAGI and PARN, 1963; DOMBROVSKAYA, 1970). Probably 

erroneous record from the British Isles (JAMES, 1965, p. 147); 2 English specimens seen by us in a folder 

labelled Stereocaulon incrustatum in BM are poorly developed states of either St. condensatum HOFFM. 

St. glareosum (SAV.) MAGN. 

Remarks: closely related to St. glareosum (SAV.) MAGN. St. tomentosum f. walamoense 

NYL. may refer to St. incrustatum (see p. 325). PASICH (1973) reported the occurrence of 

soralia in several specimens of St. incrustatum from Poland, and showed that their abnormal 

occurrence is due to the influence of external conditions, probably as a result of atmospheric 

pollution by dust. 

Fo. gracile FREY 

Rabenhorsts Krypt.-F1. 9, Abt. IV, 1. Halfte: 176 (1932) ("'gracilis", corrected to gracile in Errata, p. 

'Wap of its distribution in the Estonian SSR given by TRASS, 1970, fig. 47.


41 1). 

Typus: Germany, Rhon Mts. (borders of Bayern and Hesse), exact locality not indicated, leg. 

MARTIUS (date ?) (M-ARN, holotypus). 

Icon.: FREY 1932, P1. 111, fig. 10. 

Remarks: distinguished by its dwarf proportions, but otherwise typical. Rare; in addition 

to the type, we have seen one specimen from Norway referable here. 

21. Stereocaulon intermedium (SAV.) MAGN. 

Goteborgs K. Vetensk.- o. Vitterh.-Samh. Handl. ser. 4, 30 (7): 23 (1926). - Stereocaulon coralloides 

f. intermedium SAV. Bot. Materialy, Notul. System. ex Inst. Cryptog. Hort. Bot. Petropol. 2 (11): 163 

(1923). 

Typus: U.S.S.R., Kamtchatka, 4 localities mentioned, leg. V. P. SAVICZ, 1908 and 1909. MAG- 

NUSSON, loc. cit., stated "Type specimen in my herbarium"; this specimen (UPS-MAGN) may be considered 

as the lectotype. It is from Poperechnyi Mountain by Koryatskaya River, leg. V. P. SAVICZ, 1908 

(no. 5654 [98]); an isolectotype duplicate is present in FH. 

Facultative synonym (inval.): Stereocaulon pseudosasakii ASAH. ex SAT^, Bull. Yamagata Agric. 

Coll. no. 1 : 46 (1949); ASAH. Misc. Bryol. Lichenol. 2 (5); 61 (1961) (nomen nudum). 

Exsicc.: ASAH. Lich. Japon. Exs. Fasc. I1 (1955) no. 95, as St. pseudosasakii (FH, UPS, US). - MAC. 

Canad. Lich. Ser. I (1893-94) no. 130, as St. coralloides (MSC, LD, s, us, WELC); Ser. I1 (1909-12) no. 21 1, 

as Stereocanlon [sic] coralloides (oc). 

Icon. : DEGEL. 1938, P1. IV, fig. b. - YOSHLM. 1974, fig. 76h; P1. 33, fig. 324. 


Distrib. : mainly northern amphipacific, but with disjunct occurrence also in eastern N. America 

(Newfoundland). U. S. S. R., (Kamtchatka, Sakhalin), Bering and St. Paul Islands in the Bering Sea, 

Japan (Hondo, Hokkaido), Canada (British Columbia, Yukon Territ., Newfoundland), U. S. A. 

(Alaska, Washington, Oregon, California). The 4 Newfoundland specimens seen and ascribed to this 

species show all the essential characters of the species. Unconfirmed record from U. S. S. R., Sayan 

Mts. region (RASSADINA, 1961). 

Remarks: closely related to the similarly amphipacific species St. tennesseense MAGN. 

The cephalodia are characteristic (pale greenish-gray, verrucose-tuberculate, usually large 

and conspicuous, with Nostoc or Stigonema). 

Fo. compactum LAMB (n. f.) 

Diagn. : Habitu humile, compactum, crustoso-pulvinatum. 

Typus: U. S. A., Alaska, Aleutian Islands, Rat Island, Amchitka, near Constantine Harbor, leg. 

G. A. LLANO, 1949 (no. 1277) (US, h010typ~~; CAN, ~SO~YPUS). 

Remarks: seen from U.S.A. (Alaska, Washington), U.S.S.R. (Kamtchatka) and Canada 

(British Columbia and Newfoundland). 

22. Stereocaulon leprocauloides LAMB 

ex W. WEB. Bryologist, 74: 193 (1971). 

Typus: New Guinea, Morobe Distr., Mt. Kaindi, near Wau, altit. 15W1800 m s. m., "along 

roadside cut-banks", leg. W. A. WEBER and D. MCVEAN, 1968 (FH, holotypus; co~o, isotypus). 

Exsicc. : Lich. Exs. Univ. Colorado Mus. Fasc. VIII (1971) no. 299 (FH, isotype material). 

Icon.: W. WEB. 1971, fig. 2. 

Mat. chim.: atranorin and lobaric acid (cfr. Fox, FOLLMANN et HUNECK, 1971). 

~istrib. : New Guinea. 

Remarks: position within the genus somewhat doubtful; may possibly belong to sect.

224 Journ. Hattori Bot. Lab. No. 43 

Stereocladiurn. 

23. Stereocaulon myriocarpum TH. FR. 

De Stereocaulis et Pilophoris Commentatio: 15 (1857); Monographia Stereocaulorum et Pilophororum : 

34 (1858); LAMB, Ergebn. Forsch.-Unternehmen Nepal Himalaya, 1 (4): 350 (1966). - Stereocaulon 

myriocarpum [f.] a. typicum TH. FR. De Stereocaulis et Pilophoris Commentatio: 15 (1857). - Stereocaulon 

tomentosum [subsp.]* St. myriocarpum (TH. FR.) NYL. Synopsis Methodica Lichenum, 1 (2): 244 

(1 860). 

Typus: 3 localities mentioned in Mexico, all collected by LIEBMANN, no holotype designated. 

DODGE (1929, p. 128) selected as lectotype the specimen from El Pelado, Sierra de OajacalB, altit. 9- 

10000 feet (27W3100 m s. m.), leg. LIEBMANN, 1841 (no. 76) (UPS-TH FR, lectotypus; c, isolectotypus). 

Facultative synonyms: Stereocaulon myriocarpum [var.] p. orizabae TH. FR. De Stereocaulis et 

Pilophoris Commentatio: 15 (1857). - Stereocaulon orizabae (TH. FR.) VAIN. Dansk Bot. Ark. 4 (11): 

7 (1926). - Stereocaulon tomentosum subsp. myriocarpum var. orizabae (TH. FR.) LAMB ex ASAH. 

Lichens, in: KIHARA, Fauna and Flora of Nepal Himalaya, 1952-53, 1: 50 (1955) (comb. inval.). - 

Stereocaulon tornentosum [subsp.]*** St. myriocarpoides NYL. Synopsis Methodica Lichenum, 1 (2): 

245 (1860). - Stereocaulon myriocarpoides (NYL.) HUE, NOUV. Arch. Mus. Hist. Nat. ser. 3, 2: 247 

(1890). 

Exicc.: CUMM., WILLIAMS et SEYMOUR, Decades N. Amer. Lich. Decade XXX (1899) no. 292, as 

St. tomentosum (DUKE, MICH, MSC, OC, WELC). - ELENK. Lich. Flor. Ross. Fasc. IV (1910) no. 162a, pr. 

p., as St. denudatum var. genuinum (FH. But in UPS-MAGN it is St. vesuvianum var. nodulosum f. umbricolum 

(FREY) LAMB). - Krypt. Exs. Vindob. Cent. XVII (1909) no. 1655c, as St. tornentosum (co~o, FH, M, s, 

UPS-MAGN, US). - KUROK. Lich. Rarior. et Crit. Exs. Fasc. 111 (1971) no. 143 (FH, LD). - Lich. EXS. 

Univ. Colorado Mus. Fasc. VIII (1971) no. 304 (COLO, FH). - LINDIG, Lich. Novo-Granatenses (1863 ?)I7 

no. 2586 (FH-TUCK). - VEZDA, Lich. Sel. Exs. Fasc. XXXVIII (1970) no. 931 (FH); Fasc. LVII (1976) 

no. 1417 (BRNU, FH). 

Icon. : TH. FR. 1858, PI. IX, fig. 1. 

Mat. chim.: atranorin, stictic acid, sometimes constictic acid, usually also some norstictic 

acid (cfr. HUNECK, 1974). Doubtfully ventosic acid (see below). A dubious compound, 

"myriocarpic acid", isolated and named by AGHORAMURTHY, SARMA and SESHADRI, 1961, 

from "Stereocaulon rnyriocarpum var. orizabae". According to C. CULBERSON (1969, pi 

156), "structure not known". The specimens investigated by the authors were from two 

different localities in the Himalayas. One of these contained atranorin and the substance 

called "myriocarpic acid" (the PD reaction is not mentioned); the other specimen was stated 

to contain atranorin, stictic acid and ventosic acid. D-arabitol was also found in both 

samples. Dr. S. HUNECK kindly analyzed for us bulk material of two collections of St. 

rnyriocarpum from the Himalayan region, and failed to find any substance corresponding to 

"myriocarpic acid", only atranorin and stictic acid; "vermutlich handelt es sich bei der 

Myriocarpsaure der indischen Autoren nur um unreine Stictinsaure, sodass Myriocarpsaure 

gar nicht existiert" (in litt., 1972). 

Distrib.: two disjunct centers of distribution: one in South, Central and North America and the 

other in Asia. Mexico, Costa Rica, Panama, Venezuela, Colombia, Ecuador, Peru, W. Indies (Dominican 

Republic), U. S. A. (New Mexico, Colorado, California, Montana, Washington, Alaska), Canada 

(British Columbia); N. India, Nepal, Sikkim, Bhutan, Tibet, Pakistan, mainland China, Formosa, 

Mongolia, U. S. S. R. (Sayan Mts., Altai Mts., Transbaikalia, Extreme Orient), New Guinea. Unveri- 

- 

l8 =Oaxaca,=Sierra Madre del Sur; in Oaxaca State. 

l7 Probably not a true exsiccat. LYNGE 

by SAYRE (1969). 

(1915-20) lists it doubtfully as such, but it is not included


fied record: Java (ZAHLBRUCKNER, 1956, as St. myriocarpoides). Erroneous record: Argentine Patagonia 

(RASANEN, 1939, as St. myriocarpoides),=St. glabrum (MULL. ARG.) VAIN. 

Remarks: perhaps too closely related to St. tomentosum to qualify as a good distinct 

species. Distinction from St. tomentosum var. alpestre FLOT. is often difficult, especially in 

North America. The distinguishing characters against St. tomentosum are the thinner and 

smoother tomentum and the more developed cephalodia (subglobose, botryose-divided, 

blueish-gray). Although largely sympatric with St. tomentosum, it shows a distinctive bicentric 

distribution pattern. It is completely lacking in Europe. In New Guinea a morphologically 

similar but chemically different species also occurs (St. papuanum LAMB). 

Var. altaicum LAMB (n. var.) 

Diagn.: Robustum, pseudopodetiis eximie validis, firmis, lignosis, substrato perarcte 

affixis, ad 2 mm crassis, usque 6(-8) cm longis, tomento tenui et laevigato vestitis vel glabrescentibus. 

Cephalodia prominentia, pulvinato-subglobosa, subbotryosa; aliter quoque 

velut in specie typica. 

Typus: U. S. S. R., Siberia, Altai Mts., Gub. Semipalatinsk, Katon-Karagai, reg. alpina, leg. P. 

KRYLOV and L. SERGIEVSKAYA, 1928 (no. 8) (UPS-MAGN, holotypus). 

Remarks: appears to be a well-defined regional infraspecific entity, characterized by its 

very strong, woody, firmly affixed pseudopodetia. Six different collections have been seen, 

all from the Altai and Sayan Mountains region. It was distinguished by MAGNUSSON as a 

new species bearing the name here given, but never published by him. 

24. Stereocaulon papuanum LAMB (n . sp.) 

Diagn.: Pseudopodetia erecta, laxe caespitosa, irregulariter vel subdichotome ramosa, 

sat robusta, longit. 5-8 cm, crassit. ad 2(-3) mm, substrato arcte vel demum laxe affixa, 

indurata et quasi lignosa, ecorticata, glabra vel glabrescentia, fete tota longitudine phyllocladiis 

densiuscule vestita; ea verrucoso-subglobosa, parva, interdum leviter papillaeformielongata 

(sed nunquam rite coralloidea), pseudopodetiorum basin versus magis squamulosa 

vel fere digitata. Cephalodia sat numerosa, pulvinata, 0.4-0.8(-1.0) mrn lata, aeruginosoalbicantia, 

tandem obsolete verruculosa, algas Nostocaceas continentia. Apothecia numerosa, 

lateralia, sessilia (nonnulla etiam terminalia), parva, ad 1 mm lata, rufonigrescentia, plana 

et tenuiter fuscomarginata, tandem modice convexa et immarginata. Conus centralis compactus, 

incolor, hyalinus; hypothecium itidem incoloratum. Hymenium 60-75 p altum, 

paraphysibus discretis, fuscocapitatis. Asci clavati, longit. 50-65 p, sporis octonis, rectis 

vel subrectis, aciculari-fusiformibus, 3(-5)-septatis, 32-40 p longis, 3-4 p latis. Atranorinum 

et acidum colensoinicum juxta substantias quatuor ignotas continet (K+flavescens, PD-vel 

levissime flavescens). -Pro forma magis robusta St. myriocarpi facile censendum esset, 

sed materiis chimicis dissimilibus plane abhorrens. 

Typus: New Guinea, Papua, Central Division, Owen Stanley Range, Mt. Albert Edward, altit. 

3680 m s. m., "large clump on a shrub growing in open grassland", leg. L. J. BRASS, 1933 (no. 4394a) 

(LIL, holotypus; CAN, 

NY, isotypes). 

Mat. chim. : atranorin, colensoinic acid, and traces of 4 unidentified substances. 

Distrib. : New Guinea. 

Remarks: morphologically equivalent to a large robust state of St. myriocarpum TH. FR. 

It is interesting to note that many specimens of the latter in New Guinea are similarly robust. 

Seems to occur indiscriminately on rocks and on lignum. 11 collections seen, all from high

elevations. 


25. Stereocaulon pileatum ACH. 

Lichenographia Universalis: 582 (1810); MAGN. Goteborgs K. Vetensk.- o. Vitterh.-Samh. Handl. ser. 

4,30 (7): 68 (1926); FREY, Rabenhorsts Krypt.-F1.9, Abt. IV, 1. Halfte: 98 (1932); HALE, HOW to Know 

the Lichens: 200 (1969); POELT, Be~timmung~~chlii~~e1 Europaischer Flechten: 634 (1969); DAHL et 

KROG, Macrolichens of Denmark, Finland, Norway and Sweden: 133 (1973). - Stereocaulon cereolinum 

[f.] a. pileatum (AcH.) TH. FR. De Stereocaulorum et Pilophororum Commentatio: 19 (1857). - 

Stereocaulonpaschale [var.] 8. pileatum (AcH.) WAHLENB. Flora Suecica, pars 11: 855 (1826). - Cereoluspileatus 

(AcH.) BOIST. Nouvelle Flore des Lichens, 2e. Partie: 34 (1903). 

Typus: Switzerland, exact locality not stated, leg. SCHLEICHER, (date ?) (H-ACH, holotypus). 

Facultative synonyms: Stereocaulon coridensatum var. sorediatum HARM. ex CROZ. Bull. Acad. Int. 

GBogr. Bot. 22: 157 (1913). - Stereocaulon pileatum var. sorediiferum NYL. ex KIEFF. Bull. Soc. Hist. 

Nat. Metz, 19: 13 (1895). - Stereocaulon pileatum f. sorediiferum (NYL.) HARM. Lichens de France, 

Fasc. 111: 369 (1907). - Cereolus pileatus var. sorediifer (NYL.) BOIST. Nouvelle Flore des ~ichens; 

2e. Partie: 34 (1903). - Stereocaulon pileatum f. sorediiferum subf. terrestre HARM. Lichens de France, 

Fasc. 111: 369 (1907) (nomen nudum). - Stereocaulon pileatum f. macrum MAGN. Goteborgs K. Vetensk.- 

o. Vitterh.-Samh. Handl. ser. 4,30 (7): 70 (1926). - Stereocaulon saxonicum BACHM. Hedwigia, 

67: 109 (1927). - Stereocladium saxonicum (BACHM.) SCHADE, STOLLE et RIEHM. Lich. Sax. Exs. Decas 

XXXIII, no. 330 (1927). 

Exsicc.: ANZI, Lich. EXS. Ztal. Super. Fasc. I (1865) no. 30a, b, as St. cereolinum (FH-TUCK). - ARN. 

Lich. Exs. (Lich. Jurae) no. 916a-c (1882) (FH, us); no. 1515 (1891) (FH. In BM doubtful, perhaps another 

species). - ARN. Lich. Monac. Exs. no. 143 (1891) (FH). - ASAH. Lich. Japon. Exs. Fasc. VI (1959) no. 

298 (UPS, us). - CLAUD. et HARM. Lich. Gall. praecip. Exs. Fasc. IX (1908) no. 425 (FH); Fasc. X (1908) 

no. 474, as St. condensatum var. sorediatum (FH). - Erb. Crittog. Ztal., ed. DE NOTARIS, CESATI, BAGLI- 

ETTO etc. Fasc. XXV (1865) no. 1228, as St. cereolinum (FH). - FR. Lich. Suec. Exs. Fasc. I11 (1824) no. 

88, pr. p. (L, mixed with St. condensatum HOFFM. In FH-TUCK it is all St. condensatum). -HARM. 

Lich. in Lotharingia (date ?) no. 165 (FH). - HAV. Lich. EXS. Norveg. Fasc. XI11 (1914) no. 459 (BG). 

- HOWE, Lich. Novae Angliae, Decade I11 (1911) no. 23 (FH). - KORB. Lich. Sel. German. Fasc. X 

(1864) no. 271, as St. cereolinum (M). - Krypt. Exs. Vindob. Cent. XXIII (1915) no. 2275, pr. p., as 

St. alpinum var. tyroliense (St. saxonicum on 2nd. corrected label) (co~o, FH, US. But in PR it is a mixture 

of St. pileatum and St. nanodes TUCK., and in S it is entirely St. nanodes). - MAGN. Lich. Sel. Scand. 

Exs. Fasc. XI1 (1939) no. 288 (CAN). - MALME, Lich. Suec. Exs. Fasc. XXXVII (date ?) no. 918, pr. 

p. (WIS. The specimen from Netherlands St. pileatum, that from Sweden St. nanodes TUCK.). - MASS. 

Lich. Ztal Exs. Fasc. VI (1856) no. 181, as St. condensatum (BM, FH-TUCK, RB). - MOUG. et NESTL. 

Stirpes Cryptog. Vogeso-Rhenanae Fasc. X (1833) no. 947 (FH-TUCK). - OLN. Herb. Lich. Orne et 

Calvados (Lich. Exs.) Fasc. VII (1882) no. 312 (DUKE). - Reliquiae Farlowianae no. 483 (1922) (FH, 

PR, WIS). - SAMP. Lich. Portugal Fasc. I11 (1923) no. 285 (LD, UPS-MAGN). - SCHADE, STOLLE et RIE- 

HMER, Lich. Sax. Exs. Decas XXXIII (1927) no. 330, pr. p., as Stereocladium saxonicum (GB. But in 

DEGEL it is St. nunodes f. carinthiacum (FREY) LAMB). - STENHAM. Lich. Suec. Exs. Fasc. I11 (1860) no. 

85, as St. cereolinum (FH-TUCK, LD). - SUZA, Lich. Bohemoslovak. Exs. Fasc. I1 (1927) no. 42 (FH, 

UPS-MAGN). - TAV. Lich. Lusit. Sel. Exs. Fasc. VIII (1964) no. 190, as St. pileatum f. sorediiferum (LD). 

- TUCK. Lich. Amer. Septentr. Exs. Fasc. V (1854) no. 113, as St. condensatum (FH, TENN. In us doubtful, 

probably another species). - WESTEND. et WALLAYS, Herb. Cryptog. Belge Fasc. XXVII (1859) no. 1316, 

as St. cereolinum (FH). - VEZDA, Lich. Sel. Exs. Fasc. I (1 960) no. 16 (LD) ; Fasc. XVII (1 965) no. 41 7 

(LD. With slight intermixture of St. nanodes TUCK.); Fasc. LV (1975) no. 1368 (BRNU). 

Icon. : FREY 1932, fig. 12; fig. 13, as St. saxonicum; 1959, fig. 3 (partly representing f. pumilum (NYL) 

LAMB); 1969, fig. 9 (opposite p. 53). - NEARING 1947, fig. VII-10. - KERSH. 1960, fig. 14. - OKSN. 

1968, fig. 144. - HALE 1969, fig. 38 1. - DAHL et KROG 1973, fig. 39. - YOSHIM. 1 974, fig. 75b ; PI. 33, 

fig. 323.


Mat. chim.: atranorin and lobaric acid, as already stated by ZOPF (1907).ls 

Distrib.: widely distributed in the northern hemisphere. Sweden, Norway, Finland, Belgium, 

Luxembourg, England, Scotland, Ireland, Faeroe Islands, France, Portugal, Germany, Austria, Switzerland, 

Italy, Czechoslovakia, U. S. S. R. (Ukraine), Canada (Qukbec, New Brunswick, Nova Scotia, 

Newfoundland), U. S. A. (Massachusetts, Vermont, New Hampshire, Maine, New York, N. Carolina, 

Michigan, Minnesota, Hawaii), West Indies (Dominican Republic), Japan. Unverified records: 

Poland (TOBOLEWSKI, 1955, 1965); Jugoslavia (KuSAN, 1953); U. S. S. R., Siberia, Irkutsk region (OKS- 

NER, 1940). ALMBORN (1948) reckons it to the southern element in Scandinavia, and it is certainly 

lacking in the northern provinces. 

Remarks: earlier authors frequently used the name Stereocaulon cereolinum ACH. for 

this species. But according to TH. FRIES (1871, p. 56), Stereocaulon cereolinum was based 

on a mixture, not including St. pileatum, and consisting mainly of Pilophoron cereolus (AcH.) 

TH. FR. The typification of Stereocaulon saxonicum BACHM. is yet to be settled (see p. 312), 

but from his descriptions and illustrations (1926, 1927, mostly under the designation of 

"Stereocaulon tiroliense" or "Stereocladium tiroliense") it clearly refers to St. pileatum, as 

was already suspected by GAMS (1967, p. 114). St. pileatum only very rarely occurs on substrata 

other than rock; a few specimens have been seen growing on sandy soil, but as they 

were distinctly capitate-sorediate, there was no risk of mistaking them for St. condensatum 

HOFFM. Ecologically, this species is of interest in being one of the few lichens capable of 

withstanding atmospheric pollution, and in the British Isles it is actually spreading into 

urban and industrial areas (KERSHAW, 1963; HAWKSWORTH, 1975). 

Fo. pumilum (NYL.) LAMB 

ex POELT, Bestimmungsschlussel Europaischer Flechten: 634 (1969). - Stereocaulon coralloides var. 

conglomeratum f. pumilum NYL. ex HARM. Lichens de France, Fasc. 111: 361 (1907). - Stereocaulon 

subcoralloides f. pumilum (NYL.) ZAHLBR. Catalogus Lichenum Universalis, 4: 668 (1927). - Stereocaulon 

coralloides var. pumilum (NYL.) OLIV. Mem. Acad. Cienc. y Art. Barcelona, ser. 3,16: 475 (1921) 

(acc. to ZAHLBR. IOC. cit.; not seen by us). 

Typus: recorded from 2 localities in France. Lectotypified by DODGE (1929, p. 114) on the specimen 

collected by RIPART (from Haute-Vienne, Bessines) and said to be in herb. RIPART (no. 108). 

Isolectotype seen by us in H-NYL; leg. RIPART, 1865, labelled by NYLANDER as 'LStere~~aul~n coralloides 

Schrad. f. pumilum", and the only specimen of that name in his herbarium. 

Facultative synonym: Stereocaulon pileatum f. ramifcans MAGN. Goteborgs K. Vetensk.- o. 


Exsicc.: LARBAL. Lichen-Herbarium Fasc. I (1879) no. 6, as St. pileatum (EM). 

Icon. : FREY 1959, fig. 3 (in part), as St. pileatum. 

Remarks: widely distributed but rare. Existing descriptions off. pumilum are not helpful, 

but MAGNUSSON'S description, as f. ramificans, is good. 

Var. nipponicum LAMB (n. var.) 

Diagn.: A specie typica differt hypothecio bene infuscato et sporis angustioribus et 

nonnihil longioribus: 28-36(40) p longis, (2.5-)3 p latis. Ceterum ut in typo. 

Typus: Japan, Kiushiu, Prov. Higo, Aida, on granitic rock, leg. K. MAEBARA, 1927 (herb. ASAH. 

no. 648) (CAN, h010typ~~; ASAH, ~SO~YPUS). 

la There have been certain discrepant reports: DUVIGNEAUD, 1942, stated St. pileatum to contain 

atranorin only, and RAMAUT and SCHUMACKER, 1962, claimed Belgian material to contain atranorin, 

lobaric acid and norstictic acid; probably a mixture of species was here involved.


Remarks: to date known only from the type material. The typical species, which also 

occurs in Japan (cfr. ASAH. Lich. Japon. Exs. no. 298), has a colorless to sometimes very 

pale yellow-brownish, but never deeply pigmented hypothecium, and its spores are 20-30 x 

4-5 LA. 

26. Stereocaulon pseudodepreaultii ASAH. 

J. Jap. Bot. 35 (10): 293 (1960); YOSHIM. Lichen Flora of Japan in Colour: 176 (1974). 

Typus: Japan, Honshu, Fukui Prefecture, Prov. Wakasa, Mt. Aobayama, leg. M. TOGASHI, 1956 

(ASAH, h010typ~S ; FH, ~SO~YPUS). 

Icon.: ASAH. 1960, fig. 4B. 


Distrib. : Japan. 

Remarks: resembles St. depreaultii DEL. in the shape of the phyllocladial granules, but 

is otherwise very different in being smaller, less branched, with smaller apothecia and paler 

cephalodia. The anatomical structure of the apothecia, however (pigmented central cone, 

etc.) shows some similarity to that found in St. depreaultii. 

27. Stereocaulon rivulorum MAGN. 


F1. 9, Abt. IV, 1. Halfte : 166 (1 932) ; POELT, Be~timrnung~~chlii~~el Europaischer Flechten : 6471648 

(1969). 

Typus: Sweden, Lycksele Lappmark, par. Tarna, Syterbacken, altit. 450m s. m., leg. A. H. 

MAGNUSSON, 1924 (no. 8237) (UPS-MAGN, lectotypus; lectotypified by TONSBERG, 1977). MAGNUSSON 

cited 10 specimens in the protologue from Sweden, Norway and Spitzbergen, without designating a 

holotype. At least 6 of these other paratype specimens belong to the lobaric acid strain of the species, 

acc. to TONSBERG, op. cit. (see below). 

Exsicc.: ARN. Lich. Exs. (Lich. Jurae) no. 651b (1878 ?), pr. p., as St. alpinurn (var.) (ups. But in 

PC and s it is St. botryosurn ACH. em. FREY); no. 1363a (1888), as St. alpinurn (FH, MICH. But in PC it is 

St. alpinurn LAUR.). - Krypt. EXS. Vindob. Cent. XXXI (1928) no. 3058 (Strain 111°) (COLO, FH, PR, S, 

UPS-MAGN, US). - Lichenotheca Fennica Fasc. XLVII (1958) no. 1153, as St. saxatile (LD). - MAGN. 

Lich. Sel. Scand. Exs. Fasc. X (1937) no. 239 (PR, sZ0). - ZAHLBR. Lich. Rarior. Exs. no. 306 (1933) (s). 

Icon. : LYNGE 1928, PI. VI, figs. 5-6. - FREY 1932, fig. 26; 1959, fig. 13. - HALE 1969, fig. 388. 

Mat. chim.: T@NSBERG (1977) has shown that 3 or possibly 4 chemical strains exist in 

St. rivulorum : 

Strain I (the typical strain, according to T@NSBERG'S lectotypification), with atranorin, 

perlatolic acid and anziaic acid. 

Strain 11, with atranorin and lobaric acid. 

Strain 111, with atranorin, stictic acid and norstictic acid (only tentatively assigned to 

St. rivulorum, and perhaps a distinct species). 

Strain IV, with atranorin only demonstrable. This is perhaps merely a deficient phase 

of one of the foregoing strains. 

Strain I has been found in Scandinavia, Iceland, Greenland and Alaska. Strain I1 and 

l0 The example in UPS-MAGN chromatogrammed; according to TONSBERG (1977) that in o also 

belongs to the lobaric acid strain, but in TRH it is a mixture of the typical (perlatolic) and lobaric acid 

strains. 

20 The example in PR belongs to Strain 11, but according to TONSBERG, IOC. cit. those in o and TRH 

are the typical (per~atd~ic) strain.


Strain IV are widely distributed in boreal mountains and in the Arctic. The tentative Strain 

I11 is known only from Greenland and Jan Mayen. 

Specimens morphologically resembling St. rivulorum but containing miriquidic acid in 

addition to atranorin, perlatolic acid and anziaic acid appear to belong to a different species, 

St. groenlandicum (DAHL) LAMB (p. 221). 

Distrib.: the total distribution of St. rivulorum, without taking into account the separate chemical 

strains, is northern circumpolar, arctic-alpine, extending southwards to the Alps of Central Europe, the 

Altai Mts. of Russia, and Mongolia. At present the distribution pattern of the various strains cannot 

be given in detail, as relatively few specimens have been investigated chromatographically. Sweden, 

Norway, Finland, Switzerland, Austria, U. S. S. R. (Kola Peninsula, Murmansk Distr., N. E. Siberia, 

Novaya Zemlya, Yakutia with the New Siberian Islands, Franz Josef Archipelago, Altai Mts., Extreme 

Orient, Kamtchatka), Mongolian People's Republic, Spitzbergen, Iceland,22 Jan Mayen, Greenland, 

Canada (QuCbec, Manitoba, Alberta, British Columbia, Yukon Territ., N. W. Territ.), U. S. A. (Colorado, 

Washington, Alaska). Erroneous record: Italy (CENGIA SAMBO, 1933),=St. alpinum LAUR. 

Remarks: most closely related to St. groenlandicum (DAHL) LAMB and St. glareosum 

(SAV.) MAGN. St. paschale [f.] c. subcrustosum FR. may be an early synonym of St. rivulorum, 

to judge from the description (see p. 324). St. rivulorum f. nana CENGIA SAMBO is a doubtful 

taxon (see p. 324). This species finds its optimal habitat in seepage channels, and is frequently 

found in places with late snow cover. Ecological notes on its occurrence in Iceland 

are given by KRISTINSSON (1974) and in Switzerland by AMMANN (1971). 

Fo. pulvinarium (SAV.) LAMB (n. comb.) 

Basionym: Stereocaulon alpinum f. pulvinarium SAV. Bot Materialy, Notul. System. ex Inst. Cryptog. 

Hort. Bot. Petropol. 2 (11): 169 (1923). 

Typus: U. S. S. R., Kamtchatka, between Sczapina River and Kronotzkoje Lake, leg. V. P. 

SAVICZ, 1909 (no. 6325 [103])23 (LE, lectotypus, lectotypif. nov.; UPS-MACN, isolectotypus). 2 other 

Kamtchatka collections (paratypes) are cited in the original description. 

Remarks: a low, crustose, mat-like form of St. rivulorum with very short pseudopodetia. 

Seen also from Novaya Zemlya and Greenland. As none of the material has been examined 

chromatographically, it is not possible to state to which chemical strain the specimens belong. 

St. alpinum f. pulvinarium was recorded from Italy by CENGIA SAMBO (1933); the material 

not seen by us, and the determination probably incorrect. 

28. Stereocaulon sasakii ZAHLBR. 

Feddes Repert. 33: 48 (1933); SATB, Cladoniales (I), in: NAKAI, T. et M. HONDA, Nova Flora Japonica: 

88 (1941); YOSHIM. Lichen Flora of Japan in Colour: 177 (1974). 

Typus: Formosa (Taiwan), Mt. Niitaka (=Mt. Morrison), on soil, leg. S. SASAKI, 1927 (no. 414 

acc. to label) (w, holotypus). 

Ex~icc.:~~ ELENK. Lich. Flor. Ross. Fasc. I1 (1907 ?) no. 73b, as St. alpinum (BM, FH, LE). 

Icon.: YOSHIM. 1974, fig. 77a, b; P1. 33, fig. 329; fig. 76k, 1, as St. myriocarpum; PI. 24, fig. e, P1. 33, 

fig. 327, as St. myriocarpum. 

Mat. chim. : atranorin and lobaric acid; occasionally in a deficient phase with atranorin 

only. 

22 ACC. to TQNSBERG (1977) also occurs in the Faeroe Islands. 

23 In SAVICZ'S original publication the collecting number, apparently by a typographical error, is 

given as 6235 (103). 

z4 The specimen distributed in KUROK. Lich. Rarior. et Crit. Exs. no. 145 as "Stereocaulon sasakii" 

is St. paschale (L.) HOFFM.


Distrib.: the typical species has an E. Asiatic distribution, occurring in Formosa, Japan, China, 

Java and the Himalayan region (N. India, Nepal, Tibet). The var. tomentosoides has its main center of 

distribution in western N. America, but also penetrates rarely into the E. Asiatic region. The var. 

simplex is of rare occurrence in both these main areas. 

Remarks: St. sasakii is morphologically analogous to St. tomentosum FR., but differs 

chemically in containing lobaric acid instead of stictic acid. This chemical difference is 

correlated with a different distributional pattern. The typical species has considerable 

resemblance to St. myriocarpum TH. FR.; the var. tomentosoides is morphologically more or 

less equivalent to St. tomentosum FR. ZAHLBRUCKNER mistakenly interpreted the apothecia 

as lecanorine. "St. myriocarpum (lobaric acid strain)" in YOSHIM. (1974, p. 177) refers to 

this species. 

Var. tomentosoides LAMB (n. var.) 

Diagn.: Habitu Stereocaulo tomentoso FR. persimile, facie sat dorsiventrali, phyllocladiis 

congestis et tandem inciso- vel digitato-squarnulosis; cephalodia evolutiora subglobosa, 

aeruginoso-glaucescentia aut interdum fuscescentia, forma ut in St. myriocarpo TH. FR. 

Apothecia parva, lateralia aut sublateralia; structura interna, sporis etc. ut in forma typica 

et St. tomentoso. 

TYPUS: U.S.A., Washington Mason Co., Webb Hill Road near Cranberry Lake, on the ground, 

leg. C. WESTMAN, 1954 (no. 7793) (FH, holotypus). 

Exsicc.: Krypt. Exs. Vindob. Cent. XLI (1957) no. 4020, as St. tomentosum (FH, L, s); Cent. XLIV 

(1962) no. 4338, as St. myriocarpum var. orizabae (FH, w). - Lich. Canad. Exs. no. 65 (1970), as St. 

myriocarpum (CAN, FH). - MAC. Canad. Lich. Ser. I (1893-94) no. 45, pr. p., as St. paschale (WIS. 

The material is however heterogeneous, 2 widely separated localities being given on the label; in LD, 

us and WELC it is St. saxatile MAGN.). 

Distrib.: Canada (British Columbia, Yukon Territ.), U. S. A. (Washington, Oregon, California, 

Colorado, New Mexico, Minnesota, Michigan, Alaska), U. S. S. R. (Arakamtchetchene Island in 

Bering Strait), Japan, N. India, Nepal. 

Remarks: may eventually prove to be specifically distinct from St. sasakii; the different 

main distributional areal would lend support to this view. 

Var. simplex (RIDD.) LAMB (n. comb.) 

Basionym: Stereocaulon tomentosum var. simplex RIDD. Bot. Gaz. 50: 298 (1910). 

Typus: U. S. A., Washington, Mt. Rainier region, "on sandy river bottom", leg. T. C. FRYE, 

1904 (FH-RIDD, holotypus; MICH, WELC, isotypes). 

Icon. : RIDD. 1910, fig. 4, as St. tomentosum var. simplex. 

Distrib. : U. S. A. (Washington, Alaska), Formosa. RIDDLE mentions also specimens from Oregon 

and British Columbia, which we have not seen. 

Remarks: distinguished by the simple to sparingly branched pseudopodetia and the 

scanty and scattered phyllocladia in a thick spongy tomentum. Atranorin and lobaric acid 

demonstrated in the type specimen by chromatography. 

29. Stereocaulon saxatile MAGN. 

Goteborgs K. Vetensk.- o. Vitterh.-Sarnh. Handl. ser. 4, 30 (7): 41 (1926); FREY, Rabenhorsts Krypt.- 

F1. 9, Abt. IV, 1. Halfte: 136 (1932); DUNCAN, Introduction to British Lichens: 80 (1970). - Stereocaulon 

evolutoides var. saxatile (MAGN.) LAMB, Canad. J. Bot. 29: 581 (1951) (nom. illegit.). 

Typus: several localities in Sweden (Medelpad) cited, all collected by E. ERIKSSON. In UPS-MAGN 

one of these is marked as "Typus" by MAGNUSSON, and should therefore be designated as the lectotype.


It is from Medelpad, Alna socken, Gubben, "p& vittrad sten", leg. E. ERIKSSON, 1926 (UPS-MAGN, 

lectotypus, lectotypif. nov.). 

Facultative synonyms: Stereocaulon paschale var. evolutoides MAGN. Goteborgs K. Vetensk.- o. 

Vitterh.-Samh. Handl. ser. 4, 30 (7): 50 (1926). - Stereocaulon evolutoides (MAGN.) FREY, Rabenhorsts 

Krypt.-F1. 9, Abt. IV, 1. Halfte: 145 (1932). - Stereocaulon paschale subsp. evolutoides (MAGN.) 

LAMB, Ann. Rept. Nat. Mus. Canada, 1952-53, Bull. no. 132: 274 (1954). Stereocaulon paschale 

subsp. evolutoides f. Iaxatum LAMB, Ann. Rept. Nat. Mus. Canada, 1952-53, Bull. no. 132: 275 (1954). 

- Stereocaulon evolutoides f. glabrescens LAMB ex THOMS. Bryologist, 57: 284 (1954) (nomen nudum). 

- Stereocaulon paschale f. taeniarum MAGN. Goteborgs K. Vetensk.- o. Vitterh.-Samh. Handl. ser. 

4, 30 (7): 48 (1926). - Stereocaulon tomentosum var. graphiticola RAs. Lichenotheca Fenn. no. 247 

(1947). 

Exsicc.: AND. Lich. EXS. Bohem. Bor. Fasc. I (1929) no. 27, as St. globosus (PR). - CUMM., WIL- 

LIAMS et SEYMOUR, Decades N. Amer. Lich. Decade I11 (1892 ?) no. 25, as St. paschale (DUKE, OC, US, 

WELC, WIS. But in c it is a mixture of St. paschale (L.) HOFFM. and St. tomentosum FR.). - CUMM. et al. 

Lich. Bor.-Amer. Decade XVI (1898) no. 151, as St. paschale (NEBC, PR, S, WELC). - FOLLM. Lich. EXS. 

Sel. Mus. Hist. Nut. Casselensi Fasc. VIII (1975) no. 158 (KASSEL). - HOWE, Lich. Novae Angliae 

Decade I11 (191 1) no. 22, as St. paschale (FH. With intermixture of St. tomentosum FR.). - Lichenotheca 

Fenn. Fasc. V (1947) no. 103, as St. squamescens (LD. But in CAN and ups it is St. paschale (L.) HOFFM.); 

Fasc. X (1947) no. 247, as St. tomentosum f. graphiticola (FH, LD); Fasc. XLI (1957) no. 1009, as St. 

squamescens (LD). - MAC. Canad. Lich. ser. 1 (1893-94) no. 45, as St. paschale (LD, us, WELC. But in 

WIS it is St. sasakii var, tomentosoides LAMB). - MAC. Canad. Lich. ser. I1 (1909-12) no. 133, as St. 

paschale (the specimen from Nova Scotia) (WELC). - MAGN. Lich. Sel. Scand. Exs. Fasc. VII (1933) 

no. 158, as St. evolutoides (LE, PR). - MALME, Lich. Suec. Exs. Fasc. XXI (1915) no. 508, pr. p., as St. 

coralloides (PR, WIS, mixed with St. dactylophyllum FLORKE; S, mixed with St. evolutum GRAEWE). - 

MERR. Lich. EXS. Fasc. I1 (1909) no. 40, as St. paschale (DUKE, FH, L, US). - PIS~T, Lich. Slovak. Exs. 

Fasc. VI (1968) no. 142, as St. evolutoides (LD). - SCHADE, STOLLE et RIEHMER, Lich. Sax. Exs. Decas 

XXVI (1927) no. 260, pr. p., as St. tomentosum (BM. Mixed with St. dactylophyllum var. occidentale 

(MAGN.) GRUMM.). - TOBOL. Lichenotheca Polon. Fasc. VI (1955) no. 113, as St. paschale (LD. But in 

FH it is apparently St. paschale (L.) HOFFM.). - VBZDA, Lich. Bohemoslovak. Exs. Fasc. I (1957) no. 

44, as St. tomentosum (FH). -VBZDA, Lich. Sel. Exs. Fasc. XI11 (1965) no. 312, as St. evolutoides 

(BRNU, LD, M); Fasc. LII (1975) no. 1290 (BRNU). 

Icon.: FREY 1932, fig. 22; PI. 11, fig. 8, as St. evolutoides. - KERSH. 1960, fig. 8, as St. evolutoides. 

- HALE 1969, fig. 383. - DAHL et KROG 1973, fig. 48. - FOLLM. 1975, fig. 1. 


Distrib. : northern suboceanic, "probably essentially middle boreal to hemiboreal, amphiatlantic" 

(AHTI, 1964, p. 24). Does not occur in the Arctic regions, nor in the southern hemisphere or Asia. 

Sweden, Norway, Denmark, Finland, Belgium, Netherlands, England, Scotland, Germany, Czechoslovakia, 

Poland, U. S. S. R. (Leningrad region, Karelia ladogensis, Estonian SSR), Canada (Ontario, 

Quebec, Nova Scotia, Newfoundland, Labrador, Manitoba), U. S. A. (New York, New Jersey, 

Connecticut, Massachusetts, Vermont, New Hampshire, Maine, Michigan, Wisconsin, Minnesota). 

Remarks: closely related to St. paschale (L.) HOFFM. A very variable species in its 

morphology, especially in N. America. St. paschale var. evolutoides, syn. St. evolutoides, 

represents the common and biologically typical state of the species; St. saxatile was based 

on a somewhat unusual state characterized by firm and wide attachment to rock, irregularly 

concrescent phyllocladial squamules, and in the persistence of the primary thallus; it is however 

the prior species-epithet, and must be used instead of evolutoides provided the 2 taxa are 

regarded as synonymous, which to us seems unquestionable. F. laxatum LAMB was published 

to designate plants of lax habitus, with loosely spreading prostrate pseudopodetia, but is 

only a modification completely transitional to the typical species. St. paschale f. taeniarum


MAGN. belongs to St. saxatile, being an environmentally produced modification due to development 

among erect moss-stems and lichens (Polytrichum, Cladinae, Cornicularia etc.) which 

constrain the pseudopodetia to grow separate and f vertically. RASANEN (1939a) claimed 

that St. paschale f. squamescens NYL. refers to this species, and used the epithet Stereocaulon 

squamescens (NYL.) RKs. for it in preference to St. evolutoides, an illegitimate procedure if 

NYLANDER'S forma should really belong here, which seems somewhat doubtful (see p. 323). 

Fo. sorediatum (MAGN.) LAMB (n. comb.) 

Basionym: Sfereocaulon paschale f. sorediafum MAGN. Goteborgs K. Vetensk.- o. Vitterh.-Samh. 

Handl. ser. 4, 30 (7): 51 (1926). 

Typus: Sweden, Varmland, par. Sillerud, near Jarnsjon, leg. A. H. MAGNUSSON, 1912 (UPS-MAGN, 

holotypus). 

Remarks: the type specimen belongs to St. saxatile, and is typical of the species as it 

commonly occurs except in the presence of a number of globose, farinose, well-delimited 

soralia mostly crowded together in one place on the surface. MAGNUSSON regarded it as a 

monstrous condition. In addition to the type material, we have seen 2 other specimens, both 

also from Varmland but from different localities, and they are exactly similar. This very 

local occurrence suggests the possibility of dissemination from a single mutant. 

Fo. paschaleoides (HAv.) LAMB 

ex POELT, Be~timrnung~schIii~~e1 Europaischer Flechten: 644 (1969). - Stereocaulon evolutoides var. 

paschaleoides HAV. Arbok Univ. Bergen, Naturvitensk. Rekke, no. 12: 17,20 (1954). 

Typus: Norway, Granvin, Gamladaemmo, near Skilsaete, "on moss- and lichen-covered gneiss 

boulders", leg. J. HAVAAS (date ?) (UPS-MAGN, CAN, isotypes). 

Exsicc.: Lichenotheca Fenn. Fasc. XXXI (1953) no. 757, as St. tomentosum (FH); Fasc. LII (1960) 

no. 1278, as St. paschale (FH). 

Remarks: a puzzling form agreeing in morphological characters with St. saxatile, but 

pale-colored as in St. paschale (L.) HOFFM., not slate-gray as is normal for St. saxatile. The 

tomentum is however gray. It seems to be almost intermediate between the two species, or 

might even be regarded as belonging to St. grande (MAGN.) MAGN. Specimens referable 

here have been seen from Sweden, Norway, Finland, Estonian SSR, Canada (Ontario, Labrador, 

Manitoba) and U.S.A. (New York, Wisconsin). 

30. Stereocaulon sibiricum LAMB (n. sp.) 

Diagn. : Pseudopodetia erecta, caespitosa, substrato arcte affixa, 1.5-2.5 cm longa, 0.4- 

1.0 mm crassa, teretia, rigida et quasi lignosa, decorticata, omnino glabra, irregulariter ramosa. 

Phyllocladia lateralia, apices pseudopodetiorum versus aggregata, basin versus rarescentia; 

minuta, primum granuliformia, ad 0.1 mm lata, dein coralloideo-ramosa, ad 0.4 mm longa 

(ut in St. subcoralloidi NYL.) aut etiam interdum lobato- vel digitato-squamulosa. Cephalodia 

numerosa, conspicua, lateraliter sessilia vel brevissime pedicellata, grisea, fuscocinerascentia, 

fusca aut aeruginoso-nigrescentia, pulvinata subglobosave, 0.5-1.0 (-2.0) mm lata, superficie 

scabrida vel minute tuberculato-verruculosa, Stigonema continentia. Soredia desunt. Thal- 

lus primarius mox evanescens, e phyllocladiis parvis, digitato-ramosis, leviter complanatis, 

sax0 horizontaliter adpressis. Apothecia terminalia, parva, ad 1 mm lata, fusca vel fusco- 

nigra, primum plana et margine integro pseudothallino pallidiori circumdata, tandem pileato- 

convexa et immarginata. Conus centralis incolor, compactus, gelatinosus. Hypothecium


fere incolor (pallide isabellinum). Hymenium 55-65 p altum, paraphysibus discretis, fuscocapitatis. 

Asci clavati, 4 0 p longi. Sporae 6-8nae, parallelae, rectae, 3-septatae, 3CL 

40 p longae, 3.CL3.5 p latae. Pycnoconidia bacilliformia, recta aut subrecta, 4-5 xca. 0.8 p. 

Atranorinum, acidum sticticum et acidum norsticticum continet (Kfflavescens, PD+ 

aurantiaco-rubescens). - Quoad habitus St. subcoralloidi NYL. et St. sterili (SAV.) LAMB 

valde simile est. 

Typus: U. S. S. R., central Siberia, Krasnoyarsk Territory, Evenkiiskii Distr., valley of Baikitin 

River, on rock, leg. N. SEMENCZUK, 1957 (no. 26) (FH, holotypus; UPS-MAGN, isotypus). 

Mat. chim. : atranorin, stictic acid and norstictic acid. 

Distrib. : U. S. S. R. (central Siberia, Kamtchatka). Seen from 4 different localities in the Krasnoyarsk 

Territory, and 1 from Kamtchatka. 

Remarks: resembles St. subcoralloides NYL. and St. sterile (SAV.) LAMB, but has a tendency 

for some of the phyllocladia to become flattened-squamulose, and is chemically different 

(stictic acid instead of lobaric acid). Was distinguished by MAGNUSON in his herbarium 

under an unpublished name. 

3 1 . Stereocaulon spathulifrum VAIN. 

Ark. Bot. 8 (4): 36 (1909); POELT, Be~timmung~~chlii~~e1 Europaischer Flechten: 635 (1969); DAHL et 

KROG, Macrolichens of Denmark, Finland, Norway and Sweden: 133 (1973). - Stereocaulon fastigiaturn 

f. spathuliferurn (VAIN.) MAGN. Goteborgs K. Vetensk.- o. Vitterh.-Samh. Handl. ser. 4, 30 (7): 37 

(1926). - Stereocaulon botryosurn f. spathuliferurn (VAIN.) FREY, Rabenhorsts Krypt.-F1. 9, Abt. IV, 1. 

Halfte: 125 (1932). - Stereocaulon botryosurn var. spathuliferum (VAIN.) LAMB, Canad. J. Bot. 29: 581 

(1951). 

Typus: Norway, Granvin, Hardanger, Nesheimshorgen, altit. 980m s. m., on rock, leg. J. J. 

HAVAAS, 1900 (TUR-VAIN, no. 3926, holotypus). 

Exsicc.: HAV. Lich. Exs. Norveg. Fasc. XIX (date ?) no. 610 (BG). - HAV. Lich. Norveg. Occid. 

Exs. Fasc. VII (1938) no. 154 (LD, PR). - MAGN. Lich. Sel. Scand. Exs. Fasc. XVI (1952) no. 392, as 

St. botryosurn var. dissoluturn (CAN, LD. Somewhat transitional to f. dissoluturn (MAGN.) LAMB). 

Mat. chim.: atranorin, stictic acid, norstictic acid, and several unidentified substances, 

two of which may be constictic and consalazinic acids. 

Distrib.: boreal-arctic, amphiatlantic-bicentric or possibly circumpolar. Not seen from central 

Europe. Sweden, Norway, Finland, Scotland, U. S. S. R. (Murmansk Distr.), Iceland, Greenland, 

Canada (British Columbia), U. S. A. (Oregon, Washington, Alaska). Erroneous record: Austria 

(MAGNUSSON, 1926),=St. botryosurn ACH. em. FREY, St. rivulorurn MAGN. and St. alpinurn LAUR. 

Remarks: superficially has some resemblance to St. coniophyllum LAMB and St. nanodes 

TUCK., but not closely related to either. Differs from St. botryosum ACH. em. FREY both 

morphologically and chemically. 

Fo. dissolutum (MAGN.) LAMB (n. comb.)a5 

Basionyrn: Stereocaulon fastigiaturn var. dissoluturn MAGN. Goteborgs K. Vetensk.- o. Vitterh.- 

Samh. Handl. ser. 4,30 (7): 36 (1926). - Stereocaulon botryosurn f. dissoluturn (MAGN.) FREY, Raben- 

horsts Krypt.-F1. 9, Abt. IV, 1. Halfte: 125 (1932). - Stereocaulon botryosurn var. dissoluturn (MAGN.) 

MAGN. Forteckning over Skandinaviens Vaxter, 4. Lavar: 49 (1936). 

Typus: 4 collections cited (3 from Sweden, 1 from Norway), no holotype designated. As lectotype 

we have selected the specimen from Sweden, Lycksele Lappmark, par. Tarna, Bjorkfors, Syterbacken, 

26 This combination made in POELT, Be~timmung~~chlii~~e1 

Europaischer Flechten: 635 (1969), but 

there invalid (full basionym reference not given).


leg. A. H. MAGNUSSON, 1924 (no. 8379) (UPS-MAGN, lectotypus, lectotypif. nov.). The other specimens 

cited in the protologue are different, belonging to St. botryosum f. congestum (MAGN.) FREY and St. 

rivulorum MAGN. 

Exsicc.: MALME, Lich. Suec. Exs. Fasc. XL (1926) no. 979B, as St. fastigiatum var. dissolutum (LD, 

s, WIS). 

Icon.: FREY 1932, PI. I, fig. 7, as St. botryosum f. dissolutum. 

Remarks: widely distributed and much more common than the typical species, from 

which it differs in the lack of apical spathulate expansions, the soredia being effusely spread 

over the pseudopodetial apices. It has a considerable resemblance to the Antarctic species 

St. antarcticum VAIN. 

Fo. globuliferum (MAGN.) LAMB 

ex POELT, Be~timrnung~~chlii~~el Europaischer Flechten: 635 (1969). - Stereocaulon fastigiatum f. 

globuliferum MAGN. Goteborgs K. Vetensk.- o. Vitterh.-Samh. Handl. ser. 4,30 (7): 37 (1926). - Stereocaulon 

botryosum f. globuliferum (MAGN.) FREY, Rabenhorsts Krypt.-F1.9, Abt. IV, 1. Halfte: 125 (1932). 

Typus: Sweden, Lycksele Lappmark, par. Tarna, Bjorkfors, Syterbacken, altit. 450 m s. m., on 

rock, leg. A. H. MAGNUSSON, 1924 (no. 8236) (UPS-MAGN, holotypus). 

Remarks: apical soredia concentrated in & well-defined subglobose masses. Rare; 

known only from the type and one specimen seen from Alaska. 

Fo. pygmaeum (MAGN.) LAMB (n. comb.)as 

Basionym: Stereocaulon botryosum f. pygmaeum MAGN. Ark. Bot. 33A (1): 91 (1946). 

Typus: Sweden, Lycksele Lappmark, par. Tarna, Brakkfjallet, altit. 600 m s. m., on irrigated rock, 

leg. A. H. MAGNUSSON, 1924 (no. 8084) (UPS-MAGN, holotypus). 

Remarks: like a stunted, partly prostrate state of f. dissolutum with granular soredia 

effusely concrescent into a dense crust. Known only from the type specimen and two specimens 

seen from U.S.A. (Washington and Oregon). 

32. Stereocaulon sterile (SAV.) LAMB 

ex KROG, Norsk Polarinst. Skr. no. 144: 89 (1968); Occas. Papers Farlow Herb. Crypt. Bot. no. 2: 1 

(1972); ibid. no. 5: 1 (1973). - Stereocaulon evolutum f. sterile SAV. Bot. Materialy, Notul. System. ex 

Inst. Cryptog. Hort. Bot. Petropol. 2 (11): 165 (1923). 

Typus: U. S. S. R., Kamtchatka, Truby Volcano near Sel'devaya Bucht, leg. V. P. SAVICZ, 1908 

(no. 2263) (LE, lectotypus; lectotypified by LAMB, 1973). 

Icon.: LAMB 1973, figs. 2a, b, c, 3,4, 5, 6,7a, b. 


Distrib. : northern amphipacific: U. S. S. R. (Kamtchatka), Canada (British Columbia), U. S. A. 

(Alaska, Washington, Oregon, California). 

Remarks: morphologically resembling St. subcoralloides (NYL.) NYL., St. sibiricum 

LAMB, St. tennesseense MAGN., and in certain states also St. evolutum GRAEWE, to which latter 

however it is not closely related. 

33. Stereocaulon subcoralloides (NYL.) NYL. 

Flora, 57: 6, footnote (1874); MAGN. Goteborgs K. Vetensk.- o. Vitterh.-Samh. Handl. ser. 4, 30 (7): 

-- 

26 This combination made in HUNECK, Phytochemistry, 11 : 1494 (1972), but there invalid (basionym 

reference not given).


28 (1926); FREY, Rabenhorsts Krypt.-F1.9, Abt. IV, I. Halfte: 200 (1932); DAHL et KROG, Macrolichens 

of Denmark, Finland, Norway and Sweden: 136 (I 973). - Stereocaulonpaschale f. subcoralloides NYL. 

Lichenes Scandinaviae : 64 (1 861). 

Typus: no type specimen designated ("id inscripsi in Mus. Fenn."). No specimen labelled by 

NYLANDER as St. paschale f. subcoralloides could be found by us in H (Finnish lichen herbarium and 

general lichen herbarium) or H-NYL. Therefore it is necessary to select a neotype, and for this we have 

chosen a specimen in H-NYL (no. 39994) from Finland, Korpilahti, leg. E. LANC (VAINIO) in 1874, 

labelled by NYLANDER Stereocaulon subcoralloides (neotypif. nov.). 

Exsicc.: Krypf. Exs. Vindob. Cent. XXI (1913) no. 2066 (FH, PR, S, WELC). - Lichenotheca Fenn. 

Fasc. XXV (1949) no. 618 (UPS-MACN)?~ - NORRL. et NYL. Herb. Lich. Fenn. Fasc. I1 (1875) no. 85, 

as St. coralloides (FH-TUCK). - RAS. Lich. Fenn. Exs. Fasc. I1 (1935) no. 56 (ups). 

Icon. : DAHL et KROC 1973, fig. 45. 

Mat. chim.: atranorin and lobaric acid (cfr. DUVIGNEAUD, 1942 and HUNECK, 1974). 

Disfrib. : boreal circumpolar. Sweden, Norway, Finland, U. S. S. R. (Karelia ladogensis, Kola 

Peninsula, Krasnoyarsk Territ., W. Siberia, Transbaikalia, Kamtchatka), Canada (Ontario, QuBbec, 

Nova Scotia, Newfoundland), U. S. A. (New York, Maine, Alaska). Erroneous records: England 

(MACNUSSON, 1926), Scotland (WATSON, 1942). The specimen from England mentioned by MACNUS- 

SON (in EM, annotated by him as St. subcoralloides) is a small state of St. dactylophyllum FL~RKE. The 

Scottish record of WATSON is based on a specimen of St. subdenudafurn HAV. (FH, commun. U. DUNCAN). 

Remarks: closely related to St. sibiricum LAMB, less closely to St. tennesseense MAGN. 

Rarely fertile. St. coralloides ,B. conglomeratum TH. FR. is given as a synonym of this species 

by MAGNUSSON (1926) and ZAHLBRUCKNER (1926-27). However, TH. FRIES (1857, p. 17) 

ascribes the epithet to E. FRIES, who had described it as St. paschale P. conglomeratum, which 

is a nomen confusum (see p. 329). VAINIO (1940, p. 10) made the combination Stereocaulon 

conglomeratum ("TH. FR.") VAIN. Both he and TH. FRIES used the epithet in the sense of the 

present species, to which however it is inapplicable. 

Fo. sorediascens LAMB (n. f.) 

Diagn.: Phyllocladia passim soredioso-dissoluta, sorediis pulvinatis, albidis, farinosis, 

0.5-1.5 mm latis, plus minusve confluentibus. 

Typus: U. S. S. R., Karelia ladogensis, Kurkijoki, Kuupala, Jaavuori, leg. V. RASANEN, 1931 

(CAN, holotypus). 

Remarks: known only from the type specimen (sterile). 

34. Stereocaulon supervestiens MAGN. 

ex MAGN. et ZAHLBR. Ark. Bot. 31A (6): 37 (1944). 

Typus: U. S. A., Hawaii, Oahu, Waianae Range, Kealea trail, leg. CRANWELL, SELLING and SKOT- 

TSBERC, 1938 (no. 5640) (s, holotypus; CB, isotypus). 

Zcon.: MAGN. et ZAHLBR. 1945, P1. VII, fig. 2. 


Distrib. : Hawaiian Islands. 

Remarks: a somewhat unclear species of doubtful taxonomic position, known only from 

4 collections. As it has some similarities to St. octomerellum MULL. ARG., it may actually 

belong to sect. Denudata, subsect. Botryoideum. The extensive and persistent primary thallus, 

with numerous dark cephalodia, somewhat resembles that of St. condensaturn HOFFM., but 

the phyllocladia of which it is composed are smaller, regularly grain-like and equal-sized, not 

27 Lichenotheca Fenn. no. 1230, as St. subcoralloides, is St. paschale (L.) HOFFM. (FH).


becoming squamuliform or subdivided. Growing on naked soil. 

35. Stereocaulon tennesseense MAGN. 

ex DEGEL. Ark. Bot. 30A (3): 48 (1941): ASAH. J. Jap. Bot. 36: 48 (1961); YOSHIM. Lichen Flora of Japan 

in Colour: 176 (1974). 

Typus: U. S. A., Tennessee, Great Smoky Mts., near Alum Cave, altit. 1515 m s. m., on moist 

rock, leg. G. DEGELIUS, 1939 (DEGEL, holotypus; US, isotypus). 

Exsicc.: KUROK. Lich. Rar. et Crit. Exs. Fasc. I1 (1969) no. 97 (FH, LD). 

Icon. : ASAH. 1961, figs. 3,4; 1961a, fig. 3. - YOSHIM. 1974, fig. 76j; PI. 33, fig. 326. 


Distrib.: disjunct-bicentric: eastern N. America and E. Asia. U. S. A. (New York, Virginia, N. 

Carolina, Tennessee), Canada (Newfoundland), Japan (Hokkaido). St. tennesseense is listed by W. 

CULBERSON (1972) as a representative of this disjunct distribution type, together with 11 other lichen 

species. Its northern occurrence in Newfoundland may place it with a group of several Newfoundland 

plants with distinctly southern affinities, the "Coastal Plain Element" of DAMMAN (1965). 

Remarks: resembles St. dactylophyllum FL~RKE, but is more closely related to St. subcoralloides 

(NYL.) NYL. 

Var. nigrofastigiatum LAMB (n. var.) 

Diagn. : Pseudopodetia fastigiato-erecta, eximie rigida lignosaque, pro maxima parte 

vel saltem basin versus fusconigricantia. Phyllocladia praesertim in apicibus pseudopodetiorum 

congesta, digitato-coralloidea. 

Typus: U. S. A., New York State, Mt. Colden near Lake Placid, altit. ca. 850 m s. m., on rocks of 

talus slope, leg. J. L. LOWE, 1952 (CAN, holotypus). 

Remarks: appears to be rather common in the Adirondack Mountains region of New 

York State, and one specimen has also been seen from N. Carolina. 

36. Stereocaulon tomentosum FR. 

Schedulae Criticae de Lichenibus Exsiccatis Sueciae, part 111: 20 (1 825); RIDD. Bot. Gaz. 50: 296 (1910); 

MAGN. Goteborgs K. Vetensk.- o. Vitterh.-Samh. Handl. ser. 4, 30 (7): 56 (1926); AND. Die Strauchund 

Laubflechten Mitteleuropas: 124 (1928); FREY, Rabenhorsts Kryp~F1.9, Abt. IV, 1. Halfte: 179 

(1932). - Stereocaulon paschale [var.] P. tomentosum (FR.) DUBY, Aug. Pyrami de Candolle Botanicon 

Gallicum, ed. 2, pars 2: 618 (1830). - Stereocaulon paschale [subsp.] b. tomentosurn BRANTH et ROSTR. 

Bot. Tidsskr. 3: 162 (1869). - Stereocaulon tonzentosum [var.] P. inciso-crenatum SCHAER. Enumeratio 

Critica Lichenum Europaeorum: 181 (1850). - Stereocaulon tomentosum f. incisocrenatum (SCHAER.) 

FREY, Rabenhorsts Krypt.-Fl. 9, Abt. IV, 1. Halfte: 184 (1932). - Stereocaulon tomentosurn [var.] a. 

campestre KijRB. Systema Lichenum Germaniae: 11 (1855). - Patellaria tomentosa [var.] y. decumbens 

WALLR. Flora Cryptogamica Germaniae, Pars prior: 440 (1831). 

Typus: Sweden, exact locality not stated. Isotype material distributed in FRIES, Lich. Suec. Exs. 

no. 90. The copy of this in ups may be considered the holotype. 

Facultative synonyms: Stereocaulon tomentosum [subsp.I2. magellanicum TH. FR. De Stereoc. et 

Pilophor. Comment.: 31 (1857). - Stereocaulon paschale var. magellanicum (TH. FR.) Nn. Mbm. Soc. 

Sci. Nat. Cherbourg, 5: 96 (1857). - Stereocaulon magellanicum (TH. FR.) ZAHLBR. Catalogus Lichenum 

Universalis, 4: 651 (1927)28. - Stereocaulon cupriniforme NYL. Flora, 48: 21 1 (1865). - Stereocaulon 

tomentosurn f. cupriniforme (NYL.) VAIN. Meddeland. Soc. Fauna F1. Fenn. 6: 100 (1881). -Stereocaulon 

tomentosum var. cupriniforme (NYL.) OLN. Mkm. Soc. Sci. Nat. Cherbourg, 36: 162 (190647). 

28 Apparently a lapsus rather than an intentional new combination.


- Stereocaulon tomentosum f. truncatum FREY, Rabenhorsts Krypt.-FI. 9, Abt. IV, 1. Halfte: 185 

(1932). 

Exsicc. ANZI, Lich. Rarior. Venet. Fasc. I (1863) no. 19 (FH-TUCK, s). - ARN. Lich. Monac. Exs. 

no. 334 (1894) (FH, PC). - BREUT. Flora German. Exs. Ser. 11, Cent. I1 (1843) no. 109 (FH, s). - BRITZ. 

Lich. Exs. FI. Augsburgs Fasc. IV (1903) no. 149 (BM, FH. In PC with slight intermixture of St. incrustaturn 

FLORKE); Suppl. V (1907) no. 787, as St. alpinum (BM, FH, M, PR). - CLAUD. et HARM. Lich. Gall. 

praecip. Exs. Fasc. IX (1908) no. 423 (FH). - CLEMENTS, Crypt. Format. Coloradens. Cent. IV (1907) no. 

306, as Stereocaulum [sic] paschale (co~o). - ELENK. Lich. Flor. Ross. Fasc. I (1901) no. 17, as St. 

tomentosum var. campestre (FH). - FLAG. Lich. Franche-Compte' Fasc. VI (1884) no. 253, pr. p. (FH, 

PC. Both mixed with St. dactylophyllum FL~RKE. FREY, 1932, p. 180, states the copy in G to contain also 

St. alpinum LAUR.). -Flora Exs. Austro-Hungar. Fasc. VIII, Cent. XVI (1885) no. 1539 (PC, PR, S, 

us). -Flora Hungar. Exs. Cent. I, Lichenes 6 (1912) no. 16 (PR, S, US). - FR. Lich. Suec. Exs. Fasc. 

I11 (1824) no. 90 (FH-TUCK, S; isotype material). - FUNCK, Cryptog. Gewachse des Fichtelgebirg's, 

Heft 1 (1801) no. 20, pr. p., as St. paschale (PR. Mixed with St. dactylophyllum FLORKE); ed. 11, Heft 41 

(1838) no. 841, as St. tomentosum B. alpestre (BM, FH). -HARM. Lich. in Lotharingia no. 157 (date ?), 

as St. tomentosum var. alpinum (FH). - HAV. Lich. EXS. Norveg. Fasc. VII (1905) no. 265 (BG). - 

HEPP, Flechten Europas Band VI, Heft 11 (1857) no. 302, as St. tomentosum B. inciso-crenatum (FH- 

TUCK, PC, s). - HOWE, Lich. Novae Angliae Decade 111 (lqll) no. 22, pr. p., as St. paschale (FH. Mixed 

with St. saxatile MAGN.). - KAVINA et HILITZ. Crypt. Cech. Exs. Fasc. VI (1937) no. 273 (PR). - 

Krypt. Exs. Vindob. Cent. XVII (1909) no. 1655 (FH, PC, WELC. In PR and s, with intermixture of St. 

incrustarum FLORKE); Cent. XLII (1958) no. 4145 (BM, FH, s). - Lichenotheca Fenn. Fasc. XXXVIII 

(1956) no. 935 (FH); Fasc. XXXIX (1956) no. 962 (FH); Fasc. XLVI (1958) no. 1140 (FH). - LOJKA, 

Lichenotheca Regn. Hungar. Exs. Fasc. I (1882) no. 12 (FH-TUCK, PC, s). - MAC. Canad. Lich. ser. I1 

(1909-12) no. 133, as St. paschale (the specimen from Ottawa) (DUKE); no. 134 (the specimen from Nova 

Scotia, Cape Breton) (DUKE). - MALME, Lich. Suec. Exs. Fasc. XXVII (date ?) no. 651 (s). - NORRL. 

et NYL. Herb. Lich. Fenn. Fasc. I1 (1875) no. 86 (FH-TUCK). - PETRAK, Flora Bohem. et Morav. Exs. 

ser. 11, Abt. 2, Flechten (1924) no. 80 (FH, PR. In s it is referable to the var. compactum FREY). - Plants 

of Wyoming, ed. Univ. of Wyoming, Laramie, no. 2690 (date ?) (co~o, us). - RABENH. Lich. Europ. 

Exs. Fasc. V (1856) no. 133 (FH-TUCK, PR, s); Fasc. XVI (1859) no. 454 (PR, s). - RAS. Lich. Fenn. Exs. 

Fasc. XI1 (1940) no. 570 (ups); Fasc. XV (1943) no. 736 (ups). - ROUM. Lich. Gall. Exs. Cent. VI 

(date ?) no. 559 (ups). - SCHULZ, Flora Gall. et German. Exs. Cent. XI1 (1848) no. 1194, as St. tomentosum 

var. /3. majus (M). - STENHAM. Lich. Suec. Exs. Fasc. I11 (1860) no. 79 (FH-TUCK, LD, US). - SUZA, 

Lich. Bohemoslovak. Exs. Fasc. V (1930) no. 135 (PR). -TUCK. Lich. Amer. Septentr. Exs. Fasc. I 

(1847) no. 23 (FH, L). 

Icon.: AND. 1928, P1. XVIII, fig. 8. ERN., NADv. et SERV. 1956, P1. VI, fig. 4. - ELENK. 1906, 

PI. 111, figs. 5-7. -FINK 1910, PI. 19. - FREY 1932, fig. 29a, b, c; PI. 111, fig. 7; fig. 8, as f. truncatum. 

-HALE 1969, fig. 386. - KERSH. 1960, fig. 10. - MIG. 1927-31, Lief. 268-277 (1928), P1. 75, fig. 1. - 

NEAR. 1947, fig. vii-8. - OKSN. 1968, fig. 147ax. - Oz. et CLAUZ. 1970, fig. 393. - RAs. 1951, PI. 11, 

fig. 11. - SAT^ 1941, fig. 27. 

Mat. chim.: atranorin, stictic acid, norstictic acid; possibly also constictic and consalazinic 

acids. Working apparently with wrongly determined or mixed collections has given 

rise to some discrepant statements: RAMAUT (1962) and RAMAUT and SCHUMACKER (1962) 

reported the presence of lobaric acid. BRUUN (1973) reported lobaric acid and bourgeanic 

acid (an aliphatic compound, known to occur in a few exotic Stereocaulon species) from a 

Norwegian specimen of St. tomentosum, but a supposedly duplicate sample previously identified 

by us failed to show either of these compounds. 

Distrib.: boreal-montane-temperate, circumpolar but avoiding the high arctic regions, extending 

20 Flora Romaniae Exs. Cent. X (1931) no. 905, as St. tomentosum, is Cladonia squamosa (SCOP.) 

HOFFM. (s, US).


into the southern hemisphere along the Andean chain to the southernmost part of S. America. Generalized 

distribution map in LAMB, 1951, fig. 9. In S. E. Asia its place is taken by the very closely related 

St. myriocarpum TH. FR. Sweden, Norway, Denmark, Finland, Belgium, Scotland, France, Spain, 

Germany, Swit~erland,~~ Austria, Czechoslovakia, Hungary, Poland, U. S. S. R. (Kola Peninsula, 

Murmansk Distr., Estonian SSR, Belorussian SSR, Lening~ad region, Karelia ladogensis, Smolensk 

ObIast, Ural Mts., Altai Mts., Sayan Mts. Krasnoyarsk Temt., Transbaikalia, Kamtchatka, Extreme 

Orient, Sachalin), Mongolian People's Republic, Iceland, Greenland, Canada (Ontario, Quebec, New 

Brunswick, Prince Edward Island, Nova Scotia, Newfoundland, Manitoba, Saskatchewan, Alberta, 

British Columbia, N. W. Territ., Yukon Territ.), Miquelon Island, U. S. A. (New York, Connecticut, 

Massachusetts, Maine, New Hampshire, Vermont, Michigan, Wisconsin, Iowa, Minnesota, Montana, 

Idaho, Colorado, Wyoming, California, Washington, Alaska), China (Yiinnan), Korea, Japan (Hokkaido), 

Venezuela, Colombia, Ecuador, Peru, Bolivia, Chile (Magallanes, Brunswick Peninsula), Argentina 

(Tierra del Fuego, Santa Cruz, Rio Negro; the var. capitatum in Prov. TucumBn), Falkland Islands. 

Unverified records: Rumania (Mo~uzr, PETRIA and MANTU, 1967), Bulgaria (NIKOLOFF, 1931), U. S. 

S.R., Georgia (PACHUNOFF, 1934) and Moscow region (MULLER ARG., 1878), Manchuria P SAT^, 1952), 

Tibet (HUE, 1898-1901), U.S.A., Ohio (HUE, 1898-1901). Erroneous records: Himalayas (KREMPE- 

LHUBER, 1868, as St. tomentosum var. alpestre),=St. myriocarpurn TH. FR.; Chile, Juan Fernandez 

(ZAHLBRUCKNER, 1924, as St. paschale var. mageNanicum),=mixture of 2 indeterminable species, not 

St. tomentosum; Antarctic Peninsula (DARBISHIRE, 191 2), = St alpinum LAUR. ; South Georgia (ZAHL- 

BRUCKNER, 1917, as St. tomentosum var. magellanicum),=St. alpinum LAUR. (cfr. also Du RIETZ, 

1926a). 

Remarks: St. tomentosum var. inciso-crenatum SCHAER., var. campestre KORB., and 

Patellaria tomentosa var. decumbens WALLR., being founded on the same type material as the 

species, are obligate synonyms of the latter; see Appendix 1, p. 307. 

Fo. botryocarpum (MAGN.) LAMB 

ex POELT, Bestirnmungsschliisse1 Europaischer Flechten: 639 (1969). - Stereocaulon botryocarpum 

MAGN. Bot. Not.: 307 (1939). 

Typus: Sweden, Gotland, Gotska Sandon, on sand dunes, leg. TH. ARWIDSSON, 1934 (S, holotypus). 

Icon.: GALLQE 1972, PI. 1, figs. 3-5; PI. 59, figs. 481487; PI. 60, figs. 488494; PI. 61, figs. 495- 

501; PI. 62, figs. 502-506; PI. 63, figs. 507-510; PI. 64, figs. 511-516; PI. 65, figs. 517-520; PI. 66, figs. 

521-526; PI. 67, figs. 527-535; PI. 68, figs. 536-540; PI. 69, figs. 541-545; PI. 70, figs. 546-551; PI. 71, 

figs. 552-554; all as St. tomentosum. 

Remarks: distinguished by the very short, semi-prostrate, dorsiventral pseudopodetia 

with dark gray, smoothly felted tomentum and crustose-concrescent phyllocladia. PD+ 

orange-red. The characters of botryosely divided apothecia and large cephalodia emphasized 

by MAGNUSSON are accidental and inconstant. Characteristic of movable sand dunes, and 

analogous to St. alpinum var. gracilentum (TH. FR.) MAGN. In addition to 3 specimens from 

the type locality, we have seen one from Canada, Prince Edward Island, on sand dunes, 

referable to this form. 

Var. alpestre FLOT. 

Flora, Jahrg. 19, 1, Beiblatt 1 : 17 (1836); FREY, Rabenhorsts Krypt.-FI. 9, Abt. IV, 1. Halfte: 185 

(1932); Ber. Schweiz. Bot. Ges. 69: 195 (1959). - Stereocaulon alpinum [f.] P. alpestre (FLoT.) TH. FR. 

Monogr. Stereoc. et Pilophor.: 358 (sep. 54) (1858). - Stereocaulon tomentosum f. alpestre (FLoT.) 

Nn. Synopsis Methodica Lichenum, 1 (2): 244 (1860). - Stereocaulon tomentosum [subsp.l**. al- 

pestre (FLoT.) HUE, Nouv. Arch. Mus. Hist. Nat. ser. 3, 2: 247 (sep. 39) (1890). 

30 Only the var. alpestre seen; cfr. also FREY (1959).


Typus: Polish-Czechoslovakian border, Riesengebirge, leg. FLOTOW (date ?) (B, holotypus, destroyed 

in World War I1 and not seen by us; it was however seen by FREY and his description of 1932 is 

based on it). 

Exsicc. : Krypt. Exs. Vindob. Cent. XIX (191 1) no. 1855b, as St. tomentosum (COLO, s, us). - 

THOMS. Lich. Arct. nos. 43, 46 (1960) (w~s). 

Icon.: FREY 1932, fig. 29e; 1959, fig. IOe. 

Remarks: morphologically puzzlingly similar to St. alpinum LAUR., but reliably distinguished 

by the PD+red reaction and, if fertile, by the small lateral apothecia. Also the 

tomentum is usually heavier than in St. alpinum. But distinction from St. myriocarpum 

TH. FR. may be difficult. FLOTOW'S original description is very good. Common and as 

widely distributed as the typical species, and more frequently found than the latter in very 

exposed situations. 

Var. cornpacturn FREY 

Rabenhorsts Krypt.-FI. 9, Abt. IV, 1. Halfte: 187 (1932). 

Typus: E. Germany, Sachsen, Plauen, Vogtland, Brambach, leg. A. SPINDLER, 1908 (HBG, holotypus; 

FREY, isotypus). 

Facultative synonym: Stereocaulon tomentosum f. tectorum TOM. Mem. Inst. Agronom. Woronesch 

(MBm. Inst. Agronom. de 1'Empereur Pierre I t~ Voroneje), 3: 127 (1918). 

Exsicc.: AND. Lich. Exs. Bohem. Bor. Fasc. I1 (1930) no. 87, pr. p., as St. globosus (PR. Mixed 

with St. evolutum GRAEWE). - HAHN, Flechten-Herbarium (1884) no. ,9, as St. tomentosum (DEGEL). - 

K ~ K Lich. , Bohem. Fasc. 111 (1913) no. 126, as St. tomentosum (PR). - SCHADE, STOLLE et RIEHM. 

Lich. Sax. Exs. Decas VIII (1925) no. 76, as St. tomentosum (BM, PR); no. 77, as St. tomentosum (BM). - 

VBZDA, Lich. Sel. Exs. Fasc. LII (1975) no. 1283 (BRNU). 

Icon.: FREY 1932, fig. 29d; 1959, fig. 10d. -TOM. 1918, PI. 11, figs. 1-3, as St. tomentosum f. tectorum. 

Remarks: distinguished by its dark gray color, flattened and dorsiventral habitus, and 

often dark (grayish) tomentum of the underside; strongly resembling St. saxatile MAGN. 

Fairly common and widely distributed in Europe, Russia and N. America. 

Var. capitaturn LAMB (n. ~ar.)~l 

Diagn. : Apothecia pro majore parte terminalia et magna, usque 2(-3) mm lata; minora 

et lateralia passim quoque occurrunt. Ceteris notis ut in forma typica. 

Typus: Argentina, Prov. Tucumin, Valle de Tafi, W. slope of Cumbre Potrerillo, altit. 3700 m s. 

m., in rocky alpine Stipa pasture, leg. I. M. LAMB, 1947 (no. 5309) (CAN, holotypus). 

Remarks: differs from the typical species in having many of the apothecia terminal and 

large, pileate and convex-immarginate, sometimes irregularly lobate. Small lateral apothecia 

of the normal type for the species may occur also on the same thallus. Habitus typical of 

St. tomentosum, with heavy tomentum; phyllocladia mostly resembling those of the var. 

alpestre. Forming clumps up to 30 cm across, with pseudopodetia upright in the center, 

more or less decumbent at the periphery. Several collections made in the same region from 

elevations of 2000 to 3700 m s. m. Collected in 1872-74 by LORENTZ and HIERONYMUS, who 

worked in the same region of Argentina (Cienega, Prov. Tucumin); specimens are present 

in UPS-TH FR and M, determined as "Stereocaulon myriocarpum P. Orizabae" and listed by 

KREMPELHUBER under that name in Flora, 61 (1878). Apparently endemic to the Precordillera 

of N. W. Argentina. 

31 Published as nomen nudum in GRASSI, 1950, p. 365.


37. Stereocaulon uliginosum LAMB (n. sp.) 

Diagn.: Pseudopodetia erecta, caespitosa vel dispersa, fragilia, irregulariter ramosa, basi 

emorientia, 2.5-4.0 cm alta, 0.5-1.0 mrn crassa, albicantia, glabra, in apicibus confertim 

sorediosa. Phyllocladia sparsa, praesertim in parte inferiori pseudopodetiorum evoluta, 

deplanato-squamulosa et digitato-ramulosa, albicantia, 0.5-0.8 mm lata, iis Stereocauli 

rivulorum MAGN. haud dissimilia. Soralia copiosa, terminalia, albida, conferta, globulosa, 

0.5-1.3 rnm lata, vulgo confluentia, subtiliter farinosa, fere laevigata. Cephalodia sparsa, 

minuta et inconspicua, pseudopodetiis lateraliter insidentia, pulvinato-subglobosa, ca. 0.5 mm 

lata, leviter fuscescentia, obsolete verruculosa vel fere laevigata, Stigonema aut Nostoc foventia. 

Apothecia haud visa. Atranorinum et acidum lobaricum continet (K flavesc., PD pallide 

flavesc.). - Habitu, crescendi mod0 et materiis chimicis cum St. alpino LAUR. necnon St. 

paschali (L.) HOFFM. congruens, sed soraliis ut in St. capitellato MAGN. instructurn. 

Typus: Iceland, N. Thing., Sandiris, Thistilfjordur Di~tr.~~, altit. 70 rn s. m., in heathland, intermixed 

with St. alpinum LAUR., leg. H. KRISTINSSON, 1968 (no. 16423 pr. p.) (FH, holotypus; AK, isotypus). 


Distrib. : Iceland, W. Greenland, possibly Sweden. 

Remarks: a sorediate species resembling a tall state of St. capitellatum MAGN. (syn. St. 

farinaceum MAGN.), but with fragile, basally emorient pseudopodetia of the paschale-type 

and chemically different. Not seen fertile. It has the same habitat ecology as St. paschale 

(L.) HOFFM., to which it is probably closely related. Found in 2 localities in Iceland by 

KRISTINSSON, in heath or grassland, forming pure stands or intermixed with St. alpinum 

LAUR. The W. Greenland specimen (Sdndre Strdmfjord, leg. SKYTTE CHRISTIANSEN, 1946, 

in c and FH) was growing on ground among mosses in Cassiope tetragona heath. The Swedish 

specimen (Torne Lappmark, leg. J. POELT, 1951, in GZU and FH) can only be tentatively assigned 

here; it agrees morphologically and chemically, but was growing on rock and is scanty and 

not very well developed. 

Sect. 2. Stereocladium (NYL.) DODGE 

38. Stereocaulon nanodes TUCK. 

Amer. J. Sci. Arts, ser. 2, 28: 201 (1859); Genera Lichenum: 144 (1872); Synopsis of the North American 

Lichens, Part I: 234 (1882); NYL. Synopsis Methodica Lichenum, 1 (2): 251 (1860); RIDD. Bot. 

Gaz. 50: 303 (1910); DODGE, Ann. Cryptog. Exot. 2 (2): 128 (1929). 

Typus: U. S. A., New Hampshire, White Mountains, 3 localities mentioned (Crystal Falls, Saco 

Falls, upper gorge of the Arnrnonoosuck), "rocks near water", leg. E. TUCKERMAN. AS lectotype we 

have selected the specimen from "in the gorge of the Arnrnonoosuck at the last ford", 1848 (FH-TUCK, 

lectotypus, lectotypif. nov.). 

Facultative synonyms: Stereocladium tyroliense NYL. Flora, 58: 302 (1875). - Stereocaulon alpinum 

var. tyroliense (NYL.) ARN. Verh. K. K. Zoo1.-Bot. Ges. Wien, 27: 549, 566 (1877). - Stereocaulon 

tomentosum var. tyroliense ("ARN."; NYL.) OLIV. Mem. Soc. Sci. Nat. Cherbourg, 36: 162 (1906-07). - 

Stereocaulon tyroliense (NYL.) LETT. Hedwigia, 60 (3): 106 (1918). - Stereocaulon nanodes f. tyroliense 

(NYL.) LAMB ex MAQUINAY, LAMB, LAMBINON et RAMAUT, Physiol. P1. (Copenhagen), 14 (2): 285 

(1961) (comb. inval.); ex POELT, Be~timmung~schlii~~e1 Europaischer Flechten: 634 (1969) (val.). - 

Stereocaulon tyroliense var. lapponicum MAGN. Goteborgs K. Vetensk.- o. Vitterh.- Samh. Handl. 

ser. 4, 30 (7): 74 (1926). - Stereocaulon hypopetrueurn VAIN. ex Us. Ann. Acad. Sci. Fenn. ser. A, 34 

(4): 60, footnote (1931). 

32 Transliterated from the Icelandic characters.


Exsicc.: ARN. Lich. EXS. (Lich. Jurae) no. 652a, b (date ?), as St. alpinum var. tyroliense (ups); 

no. 1541 (1892), as Stereocladium tyroliense (EM). - Krypt. Exs. Vindob. Cent. XXIII (1915) no. 2275, 

pr. p., as St. alpinum var. tyroliense (St. saxonicum on 2nd. emended label) (s. But in co~o and FH it is 

St. pileatum AcH., and in PR a mixture of St. nanodes and St. pileatum). - MAGN. Lich. Sel. Scand. Exs. 

Fasc. I (1927) no. 6, as St. tyroliense var. lapponicum (ups); Fasc. X (1937) no. 240, as St. saxonicum 

(CAN, LD, PR, S, UPS-MAGN). - MALME, Lich. Suec. Exs. Fasc. XXXVII (date ?) no. 918, pr. p., as St. 

pileatum (the specimen from Sweden) (wrs). - PISUT, Lich. Slovak. EXS. Fasc. IX (1972) no. 211 (BRNU). 

- RAs. Lich. Fenn. Exs. Fasc. V (1936) no. 201, as St. tyroliense var. lapponica (UPS). - SUZA, Lich. 

Bohemoslovak. Exs. Fasc. 1V (1929) no. 105, as St. saxonicum (UPS-MAGN); Fasc. VII (1932) no. 198, 

as St. tyroliense (PR). - V~ZDA, Lich. Bohemoslovak. Exs. Fasc. I1 (1957) no. 45, as St. tyroliense (FH, 

LD). - VEZDA, Lich. Sel. Exs. Fasc. XVII (1965) no. 416 (LD); Fasc. XXXlX (1971) no. 965 (FH, LD); 

Fasc. LII (1975) no. 1289 (FH). 

Icon.: FREY 1932, fig. 15; PI. I, fig. 2, as St. tyroliense; PI. I, fig. 3, as St. tyroliense var. lapponum 

[sic]; 1959, fig. 4. - Oz. et CLAUZ. 1970, fig. 387A, B. - DAHL et KR~C 1973, fig. 38. 

Mat. chim. : atranorin and lobaric acid. Occasionally in a deficient phase with atranorin 

only (cfr. HUNECK, 1966). RAMAUT and SCHUMACKER (1962) reported atranorin, lobaric 

acid and stictic acid, probably as a result of working with heterogeneous material. 

Distrib. : northern and central Europe; very locally in N. E. North America (Mt. Washington area 

in New Hampshire). Sweden, Norway, Finland, Belgium, England, Germany, Switzerland, Austria, 

Italy, Czechoslovakia, U. S. S. R. (Karelia ladogensis), U. S. A. (New Hampshire). Unverified records: 

Poland (TOEOLEWSKI, 1965; NOWAK, 1968); Argentina (RASANEN, 1941). Erroneous records: Newfoundland 

(ECKFELDT, 1895), =mixture of St. pileatum ACH. and Cladonia decorticata (FLORKE) SPRENG, 

fide RIDDLE (1910); Peru (LEIGHTON, 1866),=Leprocaulon congestum (NYL.) LAMB et WARD. 

Remarks: originally collected in 1848, it still occurs in the locus classicus (Amrnonoosuck 

River, New Hampshire), where we collected good and typical material in 1966. St, tyroliense 

var. Iapponicum MAGN. represents the optimally developed state, and St. tyroliense (NYL.) 

LETT. the commonly occurring incompletely developed state; St. hypopetraeum VAIN. represents 

an intermediate condition. Some ecological notes on its occurrence in Switzerland 

are given by FREY (1959). St. nanodes is commonly a pioneer plant on substrata rich in heavy 

metals, such as slag heaps, and is highly resistant to atomospheric pollution. It may occur 

on calamine slag substrata with a zinc content of up to 700 p.p.m., and concentrates that 

element in the thallus up to 3300 p.p.m. (MAQUINAY, LAMB, LAMBINON and RAMAUT, 1961; 

LAMBINON, 1964). POELT (1972) records its occurrence in Berlin, obviously as a result of 

quite recent colonization; some specimens were growing on the rusty surface of disused iron 

railroad tracks. It is usually saxicolous, but occasionally occurs also on lignum; it was 

found in Tirol growing on wooden vineyard poles regularly sprayed with copper compounds 

(POELT and HUNECK, 1968). 

Fo. carinthiacum (FREY) LAMB 

ex FREY, Ber. Schweiz. Bot. Ges. 69: 189 (1959); POELT, Bestimmungsschlii~~el Europaischer Flechten: 

634 (1969). - Stereocaulon carinthiacum FREY, Rabenhorsts Krypt.-FI. 9, Abt. IV, 1. Halfte: 113 

(1932). 

Typus: Austria, Karnten, "Sieben Hiigel" near Klagenfurt, leg. J. STEINER (no. 195; date ?) (B, 

holotypus, destroyed in World War 11, and not seen by us). FREY collected more material in the type 

locality in 1931, and it is preserved in CAN, FREY and UPS-MAGN. 

Facultative synonym: Stereocaulon ostrobottniae MAGN. ex AHLN. Acta Soc. Fauna FI. Fern. 62 

(8): 7 (1940). 

Exsicc.: SCHADE, STOLLE et RIEHM. Lich. Sax. Exs. Decas XXXIII (1927) no. 330, pr. p., as Stereo-


cladium saxonicum (DEGEL. But in GB it is St. pileatum AcH.). 

Icon.: FREY 1932, fig. 16, as St. carinthiacum; 1959, fig. 5. 

Remarks: in this form the distinct bifacial flabellate primary squamules are imperfectly 

developed or only rarely present in the youngest stages; the apices of the pseudopodetia tend 

to branch and soon become effusely sorediate on all sides. Not uncommon; specimens seen 

from Sweden, Norway, Finland, Austria, Switzerland, Germany and Czechoslovakia. 

Fo. schadeanum LAMB (n. f.) 

Diagn. : Habitu pulvinato, compacto, squamulis primariis supra apices pseudopodetiorum 

elevatis et superficiem crustosam efficientibus. Pseudopodetia brevissima, congeste ramificantia, 

plus minusve denigrata. 

Typus: E. Germany, Sachsen, Freiberg, near Brand, "auf Gneisssteinchen und Erde auf einer 

Bergwerkshalde", altit. 510 m s. rn., leg. A. SCHADE, 1927 (no. 1767) (UPS-MAGN, holotypus; CAN, 

isotypus). 

Facultative synonym (inval.): "Stereocaulon incognitum ad int." SCHADE, Sitzungsber. Naturwiss. 

Ges. Isis (Dresden), Jahrg. 1932: 159 (1933). 

Remarks: differs from the "tyro1iense"-condition in its compact-pulvinate growth-habit, 

with the primary unifacial-sorediate squamules all raised up on the ends of very short, congestedly 

branched, usually blackened pseudopodetia, and forming a more or less evenly 

crustose surface to the pulvinulae. SCHADE'S epithet "incognitum", referring to material 

from the type locality, is a nomen nudum proposed only provisionally. Rare; seen from 4 

localities in Germany and Czechoslovakia. 

Sect. 3. Denudata JATTA 

Subsect. A. Denudata 

39. Stereocaulon vesuvianum PERS. 

Ann. Wetterauischen Ges. Gesammte Naturk. 2: 19 (1810); TORN. Lichenographia Sicula: 108 (1849); 

JATTA, Sylloge Lichenum Italicorum: 98 (1900); YOSHIM. Lichen Flora of Japan in Colour: 178 (1974). 

- Stereocaulon botryosum [subsp.] ,B. vesuvianum (PERs.) ACH. Synopsis Methodica Lichenum: 285 

(1814). - Stereocaulon alpinum var. vesllvianum (" Vesuviana") (PERs.) LINK, Grundriss der Krauterkunde, 

Vierter (Praktischer) Theil: 168 (1833)33. - Stereocaulon denudatum [var.] B. vesuvianum (PERs.) 

LAUR. ex HEPP, Flechten Europas, Band I, Heft 1, no. 2 (1853). - Stereocaulon denudatum [subsp.It 

vesuvianum (PERs.) TH. FR. De Stereoc. et Pilophor. Comment.: 28 (1857). 

Typus: Italy, Mt. Vesuvio, leg. ?, date ? (L-PERS, holotypus; H-ACH, isotyp~s).~~ 

Exsicc. : ASAH. Lich. Japon. Exs. Fasc. I (1954) no. 45, as St. vesuvianum var. umbonatum (UPS, us). 

- BORN~LLER, Plant. Exs. Canar. (1901) no. 3245, as St. denudatum (PR); no. 3506, as St. denudatum 

var. pulvinatum (PR). - Erb. Crittog. Ital., publ. Soc. Crittogamolog. Ital. Ser. 11, Fasc. I (1868) no. 20 

(FH, LD, M). - FOLLM. Lich. EXS. Sel. Mus. Bot. Berol. Fasc. IV (1970) no. 76 (KASSEL); Lich. Exs. Sel. 

Mus. Hist. Nut. Casselensi Fasc. VII (1975) no. 138 (KASSEL). - HEPP, Flechten Europas Band 1, Heft 1 

(1853) no. 2, as St. denudatum B. vesuvianum (FH, PC. In FH-TUCK with intermixture of St. vulcani 

(BORY) AcH.). - JATTA, Lich. Ital. Merid. Exs. (1874-75) no. 45 (M). - Krypt. Exs. Vindob. Cent. 

33 Not seen. Reference from ZAHLBRUCKNER, 1926-27. 

34 Of the 3 syntype specimens in herb. PERSOON, 2 are labelled "Ex Sicilia"; the third has no indication 

of locality, but is annotated "Act. Soc. Vetteraviae"; it must be from Mt. Vesuvio and therefore 

the holotype of the species. The isotype specimen in herb. ACH. is labelled "Vesuvii crater Persoon".


XXXVI (date ?) no. 3576, as St. denudatum (co~o, FH, US); Cent. XLIV (1963) no. 4337, as St. denudarum 

var. vesuvianum (FH, LD). - MANDON, Lich. Mudsre (date ?) no. 44, as St. denudatum (PC. With 

slight admixture of St. azoreum (SCHAER.) NYL.). - MASS. Lich. Ztal. EXS. Fasc. 1 (1855) no. 10 (FH- 

TUCK). - RABENH. Lich. Europ. Exs. Fasc. VI (1857) no. 160 (FH-TUCK, LD, M, PR). - SAMP. Lich. 

Portugal Fasc. 111 (1923) no. 287, as St. denudatum (LD). - SCHAER. Lich. Helvet. Exs. Fasc. XXVII 

(date ?) no. 652, as St. denudatum ,B. vesuvianum (FH. In some other copies it is St. flavireagens GYELN, 

=St. vulcani (BORY) ACH. acc. to DEGELIUS, 1941, p. 28). - TREV. Lichemtheca Veneta Fasc. IV (1869) 

no. 145 (ups). - VBZDA, Lich. Sel. Exs. Fasc. XXX (1969) no. 731 (FH, LD). 

Icon.: NYL. 1860, PI. VII, figs. 30a-d, as St. denudatum var. vesuvianum. - LICOPOLI 1871, PI. I, 

figs. 1-18. - POLI 1964, PI. XVI, fig. 11 (field photograph). - YOSHIM. 1974, fig. 77e; PI. 33, fig. 332. 

Mat. chim.: atranorin, stictic acid, usually also norstictic acid. RAMAUT and SCHU- 

MACKER (1962) claimed that a Belgian specimen of var. denudatum (=var. nodulosum (WALLR.) 

LAMB) contained atranorin, stictic acid and lobaric acid. A substance called 4,6-dihydroxy-2 

methoxy-3-methylacetophenone was isolated and characterized from St. vesuvianum by 

BOLOGNESI, CHIOCCARA and SCHERILLO (1974). 

Distrib.: widely distributed in both hemispheres, especially in volcanic regions. Material attributed 

to the typical species has been seen from Italy, Portugal, Switzerland, England (see Remarks 

below), U.S.S.R. (Kamtchatka), Iceland, Spitsbergen, Greenland, U.S.A. (Alaska, Hawaii), Formosa, 

Japan, New Zealand, W. and central Africa, Cape Verde Islands, Madeira, Canary Islands, Azores, 

Rkunion, Colombia, Chile and Argentina. The distribution of the various infraspecific taxa shows 

considerable differences, and is separately enumerated for each. 

Remarks: a very polymorphic species which has been broken down, sometimes not very 

satisfactorily, into a number of infraspecific taxa. The commonest of these in the northern 

hemisphere is var. nodulosurn (WALLR.) LAMB (syn. St. denudatum FLORKE). The distribution 

pattern of the whole complex is almost cosmopolitan. Some of the taxa may be sorediate, 

but this character seems to have liitle taxonomic importance in this species. 

St. vesuvianum (var. vesuvianum) is an early colonizer of lava flows; studies on this aspect 

of its ecology have been made by LICOPOLI (1871), POLI (1964), KLEMENT (1965) and KRISTINS- 

SON (1974a). KLEMENT, op. cit., distinguishes a special association, the Stereocauletum 

vesuviani, of which it is the dominant component. Information and field photographs of 

the association in the Canary Islands are given by FOLLMANN (1976), who states that initial 

stages may be observed even on quite recent lava flows which have not completely cooled 

down. In most places where it occurs, the typical species, unlike some of its varieties, grows 

on volcanic scoria; of interest in this respect is the only known collection from the British 

Isles: England, Kent, Lydd-on-Sea, "on clinker by railway halt", coll. P. W. JAMES and E. 

FREY, 1958 (BM); the artificial substratum closely resembles volcanic scoria. 

The spores of St. vesuvianum and its varieties are 3-5 (-6)-septate, (25-) 30-50 (-55) x 

2.5-3.5 p. TOMASELLI and TRUGLIO (1966) claimed that Italian material may have very 

stout spores, up to 12 p in breadth. We are unable to confirm this statement. 

Fo. santorinense (J. STEIN.) LAMB 

ex POELT, Bestimm~ngsschliisse1 Europaischer Flechten : 637 (1969). - Stereocaulon denudatum subsp. 

santorinense J. STEIN. Verh. 2001.-Bot. Ges. Wien, 69: 90 (1920).36 

Typus: Greece, Santorin, on 1866 lava flow, leg. SCHIFFNER and V. WETTSTEIN, 1911 (no. 7257) 

(w, holotypus). 

36 ZAHLBRUCKNER, 

1926-27, p. 666 (1927) quotes it as Stereocaulon santorinense; apparently not an 

intentional combination.


Exsicc.: ZAHLBR. Lich. Rarior. Exs. no. 198 (1921), as St. santorinense (FH). 

Remarks: analogous to the f. sessile of var. nodulosum. Rare; seen from Greece and the 

Azores. 

Var. noriulosum (WALLR.) LAMB 

ex POELT, Be~timmung~~chlu~sel Europaischer Flechten: 633 (1969); YOSHIM. Lichen Flora of Japan in 

Colour : 178 (1974). - Patellaria paschalis [var.] a. nodulosa WALLR. Flora Cryptogamica Germaniae, 

Pars prior: 441 (1831). - Stereocaulon denudatum f. nodulosa "Wallr." (WALLR.) VAIN. Meddeland. 

Soc. Fauna FI. Fenn. 14: 20, 1886 (1888). 

Typus: listed by WALLROTH from several localities in Germany, the only one of which mentioned 

by name is "Bructero Hercyn.", referring to the Brocken in the Harz Mountains of central Germany. 

The type material seen by us in STR is unlocalized, and VAINIO (1886a) also does not mention any locality 

for the type material seen by him in B. Probably a neotype from the same region should be designated. 

Facultative synonyms: Stereocaulon denudatum FLORKE, Deutsche Lichenen, Lief. 4: 13 (1819). - 

Baeomyces denudatus (FLORKE) HEPP, Lichenen-Flora von Wurzburg: 89 (1824). - Stereocaulon paschale 

[var.] 6. denudatum (FLORKE) SCHAER. Lichenum Helveticorum Spicilegium, Pars prima, Sect. 

VI: 274 (1833). - Stereocaulon paschale [f.] 6. denudatum (FLORKE) MUDD, A Manual of British Lichens: 

66 (1861). - Stereocaulon vesuvianum var. denudatum (FLORKE) LAMB ex POELT, Verh. Zoo1.- 

Bot. Ges. Wien, 95: 111 (1955). - Patellariapaschalis [var.] /3. bracteata WALLR. Flora C~yptogamica 

Germaniae, Pars prior: 442 (1831). - Stereocaulon denudatum f. bracteata "Wallr."; (WALLR.) VAIN. 

Meddeland. Soc. Fauna F1. Fenn. 14: 20 (1886). - Stereocaulon denudatum [status] b. validum LAUR. 

ex FR. Lichenographia Europaea Reformata: 205 (1831). - Stereocaulon denudatum [f.] b. validum 

(LAuR.) LAUR. ex RABENH. Deutschlands Kryptogamen-Flora, 2, Erste Abth., Lichenen : 11 1 (1845). - 

Stereocaulon denudatum var. validum (LAuR.) "TH. FRIES"; OLIV. MBm. Soc. Sci. Nat. Cherbourg, 36: 

163 (1907). - Stereocaulon denudatum [status] a. tenue LAUR. ex FR. Lichenographia Europaea Reformata: 

205 (1831). - StereocauIon denudatum [f.] a. tenue (LAuR.) LAUR. ex RABENH. Deutschlands 

Kryptogamen-Flora, 2, Erste Abth., Lichenen: 111 (1845). - Stereocaulon denudatum [f.] /3. commune 

[subf.] aa. tenue (LAuR.) TH. FR. De Stereoc. et Pilophor. Comment.: 27 (1857). - Stereocaulon 

vesuvianum f. tenue (LAuR.) OKSN. Flora Lishainikiv Ukraini, 2: 359 (1968) (comb. inval.). - Stereocaulon 

denudatum [f.] b. capitatum FLOT. ex KORB. Systema Lichenum Germaniae: 13 (1855). - Stereocaulon 

vesuvianum var. capitatum (FLoT.) LAMB ex HAK. Arch. Soc. 2001. Bot. Fenn. "Vanamo", 9: 

116 (1955) (comb. inval.); P. JAMES, Lichenologist, 3: 98 (1965) (val.). - Stereocaulon vesuvianum var. 

denudatum f. capitatum (FLoT.) POELT, Verh. Zoo1.-Bot. Ges. Wien, 95: 11 1 (1955) (comb. inval.). - 

Stereocaulon denudatum [f.] P. commune TH. FR. De Stereoc. et Pilophor. Comment. : 27 (1857). - Stereocaulon 

denudatum [status] c. digitatum LAUR. ex FR. Lichenographia Europaea Reformata: 205 (1831). 

- Stereocaulon denudatum [f.] c. digitatum (LAuR.) RABENH. Deutschlands Kryptogamen-Flora, 2, 

Erste Abth., Lichenen: 11 1 (1845). - Stereocaulon denudatum [f.] /3. commune [subf.] /3/3. digitatum 

(LAuR.) TH. FR. De Stereoc. et Pilophor. Comment.: 27 (1857). - Stereocaulon denudatum [var.] a. 

genuinum TH. FR. Lichenographia Scandinavica, Pars prima: 50 (1871). - Stereocaulon nabewariense 

ZAHLBR. ex YAS. Bot. Mag. (Tokyo), 29: 320 (1915); Ann. Mycol. 14: 56 (1916) ("nabewaziense"). - 

Stereocaulon vesuvianum f. nabewariense (ZAHLBR.) ASAH. Lich. Japon. Exs. Fasc. 11, no. 96 (1955) 

(comb. inval.). 

Exsicc.: ABB. Lich. Armor. Spect. Exs. Fasc. IV (1929-30) no. 62, as St. denudatum (LD, UPS). - 

AND. Lich. Exs. Bohem. Bor. Fasc. IV (1931) no. 186, as St. denudatum (PR. Somewhat transitional to 

f. umbonatum (WALLR.) LAMB). - ANZI, Lich. Rar. Langob. Exs. Fasc. I (1861) no. 15, as St. denudatum 

var. capitatum (FH-TUCK). - ARN. Lich. EXS. (Lich. Jurae) no. 1576 (1893), as St. denudatum (BM, PC). 

ASAH. Lich. Japon. Exs. Fasc. I1 (1955) no. 96, as St. vesuvianum f. nabewariense (ups, us). - BARTLIN~ 

et HAMPE, Veget. Cellul. German. Septentr., Lich. Decas IV (date ?) no. 34, as St. denudatum (M). - 

BREUT. Flora German. Exs. Ser. 11, Cent. I1 (1843) no. 105, as St. denudatum (FH). - DESM. PIantes


Cryptogames de France. ed. 2, ser. 11, Fasc. X (1857) no. 494, as St. denudatum (FH). - FLORKE, 

Deutsch. Lich. Fasc. IV (1819) no. 79, as St. denudatum (PR, UPS. Isotype material of St. denudatum 

FLORKE). - FOLLM. Lich. Exs. Sel. Mus. Hist. Nut. Casselensi Fasc. VIII (1975) no. 159 (KASSEL). - 

FR. Lich. Suec. Exs. Fasc. XI1 (1852) no. 346, as St. denudatum (FH-TUCK). - FUNCK, Cryptog. 

Gewachse des Fichtelgebirg's, Ed. 11, Heft 32 (1826) no. 662, as St. denudatum (FH). -HARM. Lich. 

GUN. Rar. Exs. Fasc. I1 (1909) no. 65, as St. denudatum (FH, PC). - HAV. Lich. EXS. Norveg. Fasc. 

VII (1905) no. 263, as St. denudatum (BG). - HAV. Lich. Norveg. Occid. Exs. Fasc. I1 (1913) no. 32, as 

St. denudatum (FH). - HEPP, Flechten Europas Band X, Heft 19 (1860) no. 546, as St. denudatum a 

validum (FH-TUCK, PC). - W. JOHNS. North of England Lich.-Herb. Fasc. VI (1896) no. 211, as St. 

denudatum (BM). - KUROK. Lich. Rarior. et Crit. Exs. Fasc. I (1966) no. 48, as St. vesuvianum 

(FH). - KUTAK, Lich. Bohem. Fasc. IV (1913) no. 171, as St. denudatum (PR, u). - LARBAL. Lichen- 

Herbarium Fasc. I (1879) no. 7, as St. tomeutosnm var. alpinurn (EM, PC); Fasc. VII (1880) no. 244, as 

St. denudatum (EM). - LEIGHT. Lich. Btri. Exs. Fasc. XI11 (1876) no. 387, as "Stereocaulon with 

cephalodia" (ups). - Lichenotheca Fenn. Fasc. V (1947) no. 104, as St. denudatum f. digitata (FH); 

Fasc. XXXI (1953) no. 759, as St. denudatum f. digitata (FH). - Lich. Canad. Exs. no. 68 (1970) 

(CAN). - Lich. EXS. Sect. Bot. Mus. Hist.-Nut. Hungar. (1969) no. 84, as St, denudatum (LD). - 

MALME, Lich. Suec. Exs. Fasc. XXI (1915) no. 510, as St. denudatum (WIS). - MIG. Cryptog. 

German., Austr. et Helvet. Exs., Flechten Fasc. LXII (1933) no. 343, as St. denudatum (FH, PR, 

u). - Mouc. et NESTL. Stirpes Cryptog. Vogeso-Rhenanae Fasc. V (1 81 5) no. 466, as St. botryosum 

(FH-TUCK, LE). -NORRL. et NYL. Herb. Lich. Fenn., Continuatio (1921) no. 805, as St. denudatum 

(UPS). - OLIV. Herb. Lich. Orne et Calvados (Lich. Exs.) Fasc. VII (1882) no. 311, as St. denudatum 

(DUKE). - PISUT, Lich. Slovak. Exs. Fasc. 111 (1965) no. 63, as St. vesuvianum (LD). - RABENH. Lich. 

Europ. Exs. Fasc. V (1856) no. 135, as St. denudatum (PR). - RAS. Lich. Fenn. Exs. Fasc. IV (1936) no. 

200, as St. denudatum (UPS); Fasc. VIII (1939) no. 398, as St. denudatum var. umbonata f. saxicola 

(UPS. Partly consisting off. umbonatum (WALLR.) LAMB). - SCHADE, STOLLE et RIEHM. Lich. Sax. Exs. 

Decas XXVI (1927) no. 259, as St. coraZZoide~~~ (EM, DEGEL, GB, LD, WELC. Another sample in DEGEL 

has intermixture of St. dactylophyllum var. occidentale (MAGN.) GRUMM.). - SOMM. Plant. Cryptog. 

Norveg. Fasc. IV (1830) no. 163, as St. denudatum (FH). - STENHAM. Lich. Suec. Exs. Fasc. I11 (1860) 

no. 83, as St. denudatum (BM, FII-TUCK, LD, US). - TOBOL. Lichenotheca Polon. Fasc. V (1954) no. 85, 

as St. denudatum (LD). - VEZDA, Lich. Bohemoslovak. Exs. Fasc. I (1957) no. 43, as St. denudatum 

(FH. In LD it has some intermixture of St. evolutum GRAEWE). - V~ZDA, Lich. Sel. Exs. Fasc. XLIII 

(1972) no. 1070, as St. vesuvianum (LD); Fasc. LII (1975) no. 1284 (BRNU). - ZW. Lich. Exs. Fasc. 

XVIII (date ?) no. 909, 909 bis, as St. denudatum (PC). 

Icon.: RIDD. 1910, fig. 6, as St. denudatum. - YAS. 1925, PI. XIII, fig. 3, as St. nabewariense. - 

MIG. 1927-31 (1928), PI. 75, fig. 2, as St. denudatum. -AND. 1928, PI. XVIII, fig. 9, as St. denudatum. 

- MAGN. 1929, PI. VI, fig. 3, as St. denudatum. -FREY 1932, fig. 20; PI. 11, fig. 6, as St. denudatum; 

PI. I, fig. 5, as St. denudatum f. capitatum. SAT^ 1941, fig. 22, as St. nabewariense. -NEAR. 1947, 

fig. VII-9, as St. denudatum. - RAs. 1951, PI. XVI, fig. 3, as St. denudatum. - KERSH. 1960, fig. 15, as 

St. vesuvianum. - ALVIN et KERSH. 1963, PI. 54, upper fig., as St. vesuvianum. - BRIGHTMAN et NI- 

CHOLSON 1966, figs. 7, 7A, as St. vesuvianum. - Oz. et CLAUZ. 1970, figs. 388, 389 A-D, as St. vesuvianum. 

- DAHL et KROG 1973, fig. 37, as St. vesuvianum. 

Distrib.: very widely distributed, almost cosmopolitan, but not occurring in S. E. Asia. Sweden, 

Norway, Denmark, Finland, Belgium, British Isles, Faeroe Islands, France, Germany, Switzerland, 

Austria, Italy, Czechoslovakia, Hungary, Poland, U.S.S.R. (Murmansk Distr., Kola Peninsula, Komi 

ASSR, Karelia ladogensis, Lapponia petsamensis, Novaya Zemlya, Kamtchatka), Bering Straits 

region, Iceland, Spitsbergen, Greenland, Jan Mayen, Canada (Qdbec, Labrador, Newfoundland, 

British Columbia), Miquelon Island, U. S. A. (Oregon, Alaska), Mexico, Colombia, Ecuador, Peru, 

Bolivia, Japan, New Zealand, W. Africa, Tristan da Cunha. Unverified records: Rumania (MORUZI, 

38 All the copies seen by us have the printed epithet "coralloides" struck out and "spissum Nyl." 

substituted in handwriting.


PETRIA and MANTU, 1967), U.S.S.R., Ukraine (OKSNER 1968), Taimyr Peninsula (ELENKIN, 1909), 

Madeira and Cape Verde Islands (TUCKERMAN, 1874, perhaps St. vulcani OJORY) AcH.), U.S.A., Pennsylvania 

(MOZINGO, 1948), Formosa (ZAHLBRUCKNER, 1933). 

Remarks: very variable in its morphology. Unlike the typical species, it grows usually 

on plutonic, metamorphic or sedimentary rocks (not lava) and is therefore found in nonvolcanic 

regions. It has a tendency towards the production of farinose soredia at the tips 

of the pseudopodetia, sometimes very distinct and conspicuous (as in material called f. capitatum) 

but often only indistinctly or not at all developed, for which reason it does not seem 

justified to separate sorediate states taxonomically. Soredia are present in the isotype specimens 

of St. denudatum seen by us (FL~RKE, Deutsch. Lich. no. 79, in PR, UPS). NO soredia 

were seen in the type material of var. nodulosum examined. 

Fo. umbonatum (WALLR.) LAMB 

ex MAQUINAY, LAMB, LAMBINON et RAMAUT, Physiologia Plantarum, 14: 286 (1961) (comb. inval.); 

ex POELT, Be~timmung~~chlii~~e1 Europaischer Flechten: 637 (1969) (val.). - Patellaria paschalis 

[var.] y. umbonata WALLR. Flora Cryptogamica Germaniae, Pars prior: 442 (1831). - Stereocaulon 

denudatum f. umbonata "Wallr."; (WALLR.) VAIN. Meddeland. Soc. Fauna F1. Fenn. 14: 21, 1886 

(1888). - Stereocaulon denudatum var. umbonatum "Wainio"; (WALLR.) ZAHLBR. Catalogus Lichenum 

Universalis, 4: 644 (1927). - Stereocaulon vesuvianum var. umbonatum (WALLR.) LAMB Canad. J. Bot. 

29: 582 (1951). 

Typus: the remarks on the typification of var. nodulosum given above apply also here. 

Facultative synonym (doubtful): Stereocaulon denudatum [f.] d. compactum FLOT. Flora, 19, Beiblatter: 

55 (1836). 

Exsicc.: W. JOHNS. North of England Lich.-Herb. Fasc. VI (1896) no. 212, as St. denudatum var. 

pulvinatum (BM). - Lichenotheca Fenn. Fasc. XIX (1948) no. 471, as St. denudatum (TUR); Fasc. XLVII 

(1958) no. 1154, as St. paschale var. alpinum (TUR). 

Icon. : FREY 1932, PI. 11, fig. 3, as St. denudatum var. umbonatum. 

Distrib.: same distribution pattern as var. nodulosum, but penetrating to higher latitudes in both 

hemispheres. A specimen from Coats Land, Antarctica, in lat. 79'12' S. (BIRM) represents the southernmost 

known record for any species of Stereocaulon. 

Remarks: may be merely a compacted, crustose modification of var. nodulosum caused 

by severe environmental conditions. This taxon has been commonly referred to as var. or f. 

pulvinatum (SCHAER.), but that epithet is inapplicable, for reasons given in Appendix 2, p. 000. 

Fo. umbricolum (FREY) LAMB (n. comb.) 

Basionym: Stereocaulon denudatum var. umbricolum FREY, Rabenhorsts Krypt.-F1. 9, Abt. IV, 1. 

Halfte: 134 (1932). - Stereocaulon vesuvianum var. umbricolum (FREY) FREY, Ber. Schweiz. Bot. Ges. 

69: 191, 192 (1959) (comb. inval.); LAMB ex POELT, Be~timmung~schlii~~e1 Europaischer Flechten: 638 

(1969) (val.). 

Typus: Switzerland, Berner Oberland, Grimsel, road to Nagelisgratli, altit. 2200 m s. m., on granite 

rock, leg. E. FREY, 1930 (no. 1429) (FREY, holotypus; CAN, isotypus). 

Icon.: FREY 1932, P1. 11, fig. 5, as St. denudatum var. umbricolum; 1959, fig. 7a, as St. vesuvianum 

var. umbricolum. 

Distrib.: rare; seen from Switzerland, Sweden, Norway, Faeroe Islands and U.S.A. (Alaska). 

Remarks: possibly merely an ecotypic modification of shaded or moist situations. 

Fo. sessile (MAGN.) LAMB 

ex POELT, Bestimmungsschliisse1 Europaischer Flechten: 638 (1969). - Stereocaulon pileatum f. sessile


MAGN. Goteborgs K. Vetensk.- o. Vitterh.-Samh. Handl. ser. 4,30 (7): 71 (1926). 

Typus: Sweden, Vastergotland; 2 localities cited, no holotype designated. As lectotype we 

selected the specimen from Halleberg, leg. C. STENHOLM, 1893 (UPS-MAGN, lectotypus, lectotypif. nov.). 

MAGNUSSON himself designated it as the type in litt. The other (syntype) specimen from Floby is different 

(St. dactylophyllurn f. sessile LAMB, n. f.; see p. 214). 

Distrib. : rare; only 3 specimens seen, all from Sweden. 

Remarks: apothecia borne directly on the squamules of the primary thallus; analogous 

to the f. santorinense (J. STEIN.) LAMB of the typical species. 

Fo. depressum (MAGN.) LAMB (n. comb.) 

Basionym: Stereocaulon denudaturn var. depressurn MAGN. Goteborgs K. Vetensk.- o. Vitterh.- 

Samh. Handl. ser. 4, 30 (7): 82 (1926). - Stereocaulon vesuvianum var. depressum (MAON.) LAMB ex 

SOWT. Rev. Bryol. Lichenol. 27: 78 (1958) (comb. inval.); An. Parques Nacionales (Buenos Aires), 

7, 1958: 101 (1959) (val.). 

Typus: several localities in Sweden and Norway cited, no holotype designated. FREY (1932, p. 

134) selected as lectotype a specimen from Sweden, Lycksele Lappmark, par. Tarna, Brandsfjallet, leg. 

A. H. MAGNUSSON, 1924 (no. 8962) (UPS-MAGN, lectotypus). Other (paratype) material mentioned in 

the protologue is different, including St. spathuliferurn VAIN. and St. symphycheilum LAMB. 

Exsicc.: HAV. Lich. Exs. Norveg. Fasc. XI11 (1914) no. 458, as St. denudatum var. pulvinatum 

(BG). 

Icon.: FREY 1932, PI. 11, fig. 4, as St. denudatum var. depressurn. 

Distrib.: same distribution pattern as var. nodulosum, with the addition of U.S.S.R., Ural Mts. 

and Franz Josef Archipelago; Argentina, Rio Negro and Neuquen. 

Remarks: MAGNUSSON in his original description mentions the occasional occurrence of 

soralia. This refers to other species present in the mixed paratype material (St. spathuliferum 

and St. symphycheilum); the lectotype specimen is not sorediate. However, soredia 

do occur, although rarely, in some specimens of this form. A very large orbicular individual 

24 cm in diameter collected by V. AHMADJIAN in Sweden, Torne Lappmark in 1959 (FH) 

must probably have been several hundred years old. 

Fo. verrucosum LAMB (n. f.) 

Diagn.: Thallus compacto-caespitosus sed haud crustosus, ad 11 cm latus vel etiam 

major, dorsiventralis, pseudopodetiis e centro radiantibus, glabris. Phyllocladia conferta, 

verrucosa, 0.2-0.4 (4.5) rnm lata, absque maculis obscurioribus, simplicia vel nonnulla confluentia. 

Typus: U.S.A., Alaska, Aleutian Islands, Attu, altit. 300 m s. m., on rocks or shallow soil over 

outcrops, leg. G. A. LLANO, 1949 (no. 1463 pr. maj. p.) (us, holotypus). 

Distrib.: uncommon; seen from Sweden (see below), Iceland, Canada (British Columbia) and 

U.S.A. (Alaska). 

Remarks: distinguished from f. umbricolum (FREY) LAMB in the more congested and 

dorsiventral habitus and the larger verrucose phyllocladia, which resemble those of St. verruciferum 

NYL. The single Swedish specimen referred here differs from the type material 

only in the presence of granulose soredia at the apices of some of the pseudopodetia. 

Var. syrnphycheileoides LAMB 

ex VEZDA, Lich. Sel. Exs. Fasc. LII no. 1281 (1975). 

Typus: Czechoslovakia, Slovakia, Carpathians, Slovenske Rudohorie Mts., MniSek in valley of


Smolnik River near Smolnickh pila, altit. 500 m s. m., on old volcanic rocks, leg. A. KISZELY and A. 

V~ZDA, 1974 (FH, holotype; BRNU, isotype; isotype material distributed in VBZDA, Lich. Sel. Exs. no. 

1281). 

Facultative synonym: Stereocaulon vesuvianum var. surugaense ASAH. Lich. Japon. Exs. Fasc. I1 

no. 97 (1955) (nomen nudum). 

Exsicc.: ASAH. Lich. Japon. Exs. Fasc. I1 (1955) no. 97, as St. vesuvianum var. surugaense (UP& us). 

- VEZDA, Lich. Sel. Exs. Fasc. LII (1975) no. 1281 (isotype material) (BRNU). 

Distrib.: Czechoslovakia, U. S. A. (Alaska), Japan. 

Remarks: strongly resembles St. symphycheilum LAMB in its morphology, but has the 

characteristic chemical constituents of St. vesuvianum. 

Var. kilimandscharoense B. STEIN 

Jahresber. Schles. Ges. Vaterl. Cult. 66: 134 (1888). 

Typus: Africa, Tanzania, Mt. Kilimanjaro; 2 specimens cited, from altit. 3000m s. m., "am 

Senecio-Bach" and from altit. 4000 m s. m., "in den Grasflachen", no holotype designated. We select 

the former as lectotype (lectotypif. nov.). The specimen in STEIN'S herbarium (BRSL ?) not seen by us, 

but an isolectotype in UPS-TH FR studied. 

Facultative synonyms: Stereocaulon confluens MULL. ARG. Flora, 67; 614 (1884). - Stereocaulon 

vesuvianum var. confluens (MULL. ARG.) B. STEIN ex MEYER, Ostafrikanische Gletscherfahrten: 315 

(1890).37 - Stereocaulon confluens var. fuscescens MULL. ARG. Flora, 73: 336 (1890). - ~tereocaulon 

confluens f. persquamosa DWIGN. Lejeunia, MCm. no. 14: 84 (1956) (nomen nudurn). 

Exsicc.: ALMB. Lich. Afric. Fasc. IV (1974) no. 85, as St. confluens (FH). - V~ZDA, Lich. Sel. Exs. 

Fasc. LV (1975) no. 1373, as St. confiens (BRNU). 

Icon.: DUVIGN. 1956, figs. 5, 6, as St. confluens f. persquamosa; figs. 20,21, as St. denudatum. 

Distrib. : material attributed to this variety seen from Mexico, Venezuela, Argentina (Tierra del 

Fuego), Juan Fernandez, Japan, Formosa, Java, New Zealand, Tasmania, Africa (Cameroon, Zaire, 

Rwanda-Burundi, Uganda, Kenya), Madagascar, RBunion. 

Remarks: the anomalous distribution pattern raises some doubt as to whether this is a 

genetically homogeneous taxon. Occasional accidental states of var. nodulosum in Europe 

and elsewhere may mimic it strongly. The enlarged, plate-like or spathulate, compound 

phyllocladial expansions which are its diagnostic feature may occur either at the apices of the 

pseudopodetia or lower down. Soredia may be present (rarely in African specimens, cornmonly 

in Mexican specimens) and are farinose-pulverulent; in the following variety (egorescens) 

they are coarsely granulose. 

Anales Parques Nac. (Buenos Aires), 7, 1958: 100 (1959). - Stereocaulon obesum var. efflorescens 

Ns. Anales Soc. Ci. Argent. 128: 136 (1939). 

Typus: Argentina, Patagonia, Rio Negro, Nahuel Huapi, altit. 1700 m s. m., on sandy ground, leg. 

DONAT (date ?) (H-RAS, holotypus). 

Facultative synonyms: Stereocaulon violascens MULL. ARG. Rev. Mycol. (Toulouse), 1 : 164 (1 879). 

- Stereocaulon vesuvianum var. violascens (MULL. ARG.) LAMB, Anales Parques Nac. (Buenos Aires), 

7, 1958: 101 (1959). 

Distrib. : Argentina (Rio Negro, Chubut), Chile (incl. Juan Fernandez), Bolivia, Colombia, Ecuador, 

Venezuela, Tasmania, W. Indies (Dominican Republic), U. S. A. (Oregon). 

Remarks: apices of pseudopodetia coarsely granulose-sorediate, the granules derived 

by gradual diminution in size of the upper granulose phyllocladia. Pseudopodetia glabrous. 

37 Not seen; reference from ZAHLBRUCKNER, 1926-27.


Lower phyllocladia sometimes partly confluent-expanded as in var. kilimandscharoense. St. 

violascens was based on a decumbent, semi-crustose, dorsiventral state of purplish- or vio- 

lascent-cinereous color induced by very exposed habitat at high altitude. 

Var. obscurum (MULL. ARG.) LAMB (n. comb.) 

Basionym: Stereocaulon obscurum M~~LL. ARG. Flora, 74: 109 (1891). 

Typus: W. Africa, Cameroon, Mt. Cameroon, altit. 5000-12000 feet, on lava, leg. E. MANN, 1862 

(no. 15) (G-M~~LL, holotypus; isotypes in H-NYL, K). 

Distrib.: Africa. Rare; seen only from the type locality and from Kenya, Mt. Kenya. 

Remarks: resembling var. nodulosum, but phyllocladia dark-colored (caesious-gray to 

brownish) and pseudopodetia more or less brown-tomentose. Perhaps an ecotypic modification. 

Var. thyrsoideum LAMB (n. var.) 

Diagn. : Habitu et mod0 crescendi ut in var. eflorescenti(Rks.) LAMB et similiter in apicibus 

pseudopodetiorum granuloso-sorediatum, sed phyllocladiis inferioribus parvis, haud ultra 

0.5 rnrn latis (ut in St. obeso TH. FR.), macula centrali obscuriore praeditis. 

Typus: Ecuador, Prov. Imbabura, Volcin Imbabura, altit. 2750-3650 m s. m., on trachyte and trachytic 

tuff, leg. M. WAGNER, 1859 (no. 5531208) (M-KREMP, holotypus; FH, isotypus). 

Distrib. : Ecuador, known only from the type material. 

Remarks: somewhat flattened and dorsiventral, 5 cm diam., with pseudopodetia glabrous 

to faintly adpressed-tomentose, radiating outwards from a central holdfast area. Phyllocladia 

very crowded, ochraceous-whitish, verrucose to peltate, towards apices of pseudopodetia 

progressively smaller and finally transformed into granulose soredia. Sterile. Was 

distinguished by KREMPELHUBER in Flora, 44: 129 (1861) as Stereocaulon sphaerophoroides 

var. thyrsoideurn ? (nomen nudum). 

40. Stereocaulon anomalum LAMB 

ex ABB. MCm. Inst. Sci. Madagascar, S6r. B, Biol. VCg. 7: 9 (1956) (nomen nudum); ex VEZDA, Lich. 

Sel. Exs. Fasc. L no. 1247 (1974) (cum descript.). 

Typus: Madagascar, W. of Manantenina River (tributary of the Lokoho), E. summit of Marojejy 

massif, altit. 1850-2137 m s. m., on gneiss and quartzite rocks, leg. H. HUMBERT, 1948 (no. 2) (PC, holotypus; 

isotypes in FH and REN). 

Exsicc.: ABB. Lich. Madagasc. et Borbon. Sel. Exs. Fasc. I1 (1966) no. 37, as St. furfuraceum (FH. 

Not very typical). - V~ZDA, Lich. Sel. Exs. Fasc. L (1974) no. 1247 (BRNU). 

Mat. chim.: atranorin, stictic acid and norstictic acid. 

Distrib. : Madagascar and Africa (Tanzania and Zaire). 

Remarks: closely related to St. furfuraceurn DUVIGN. and St. fibrillosum LAMB. 

41. Stereocaulon arcticum LYNGE 

Skr. Norske Vidensk.-Akad. Oslo, Mat.-Naturv. K1. no. 6: 69 (1938); ibid. no. 7: 35 (1940); Skr. om 

Svalbard og Ishavet, no. 76: 30 (1939); POELT, Bestimmungsschliissel Europaischer Flechten: 637 

(1969); DAHL et KROG, Macrolichens of Denmark, Finland, Norway and Sweden: 131 (1973). - 

Stereocaulon vesuvianum var. arcticum (LYNGE) LAMB ex HALE, Amer. Midl. Naturalist, 51: 245 

(1954). 

Typus: Jan Mayen, Hageruphytta, leg. J. LID, 1930 (0, holotypus).


Facultative synonym: Stereocaulon alpinum var. Janii LYNGE ex DAHL, LYNGE et SCHOL. Skr. om 

Svalbard og Ishavet, no. 70: 39 (1937) (nomen nudum). 

Exsicc.: ELENK. Lich. Flor. Ross. Fasc. IV (1910) no. 162b, as St. denudatum var. pulvinatum (FH). 

- Lich. Groenlandici Exs. (ed. Mus. Bot. Hauniense) Fasc. I (1972) no. 49, as St. vesuvianum (LD). - 

RAS. Lich. Fenn. Exs. Fasc. VIII (1939) no. 397, as St. denudatum var. umbonata f. pulvinata (ups). 

Icon. : LYNGE 1938, PI. XII, fig. 3 ; PI. XIII, figs. 1,2; PI. XIV, fig. 1. 

Mat. chim.: atranorin, stictic acid, and small amounts of norstictic acid. 

Distrib. : boreal-arctic, circumpolar. Sweden (Jamtland), Norway (Nordland, Finnmark), Finland 

(Lapponia enontekiensis), U.S.S.R. (Lapponia petsamoensis, W. Siberia, Novaya Zemlya, Kamtchatka, 

Extreme Orient), Iceland, Faeroe Islands, Spitsbergen, Greenland, Jan Mayen, Canada (N. W. Territ.), 

U. S. A. (Alaska). 

Remarks: very similar to St. vesuvianum var. nodulosum (WALLR.) LAMB, but distinguished 

by its ecology, basally emorient pseudopodetia, and other characteristics. The distinguishing 

features between these two taxa tabulated by DAHL, 1950, p. 115. The presence of Nostoc 

in the cephalodia, a character emphasized by LYNGE, is usual but by no means constant; 

sometimes both Nostoc- and Stigonema-cephalodia occur on the same individual. Closely 

related to St. arenarium (SAV.) LAMB; "St. arcticum, chemischer Stamm mit Atranorin u. 

Dendroidin" in POELT, 1969, p. 647, refers to the latter species. ~~1snNss0N (1974) reports 

St. arcticum to be a common species in Iceland, occurring on soil in heathland, tundra mounds 

and river flats. 

Var. turjaZnse (RAs.) LAMB (n. comb.) 

Basionym: Stereocaulon denudatum subsp. turja2nsis RAs. Lich. Fenn. Exs. Fasc. VIII no. 399 

(1939). 

Typus: U.S.S.R., Lapponia petsamoensis, Pechenga (Petsamo), Vaitolahti, "in humo arenosa 

prope Mare glaciale", leg. V. RASANEN, 1938 (H-RAS, holotypus; isotypes distributed in Us. Lich. 

Fenn. Exs. no. 399). 

Exsicc.: RKs. Lich. Fenn. Exs. Fasc. VIII (1939) no. 399, as St. denudatum subsp. turjaEnsis (isotype 

material) (H, UPS). 

Remarks: pseudopodetia small, not over 12 mm in length. Looks like a miniaturized 

version of St. verruciferum NYL. Generally distributed over the same area as the typical 

species. 

42. Stereocaulon arenarium (SAV.) LAMB 

Occas. Papers Farlow Herbarium of Cryptogamic Botany, no. 2: 1 (1972). - Stereocaulon denudatum 

var. pulvinatum f. arenarium SAV. Bot. Materialy, Notul. System. ex Inst. Cryptog. Hort. Bot. Petropol. 

2 (11): 171 (1923). 

Typus: U.S.S.R., Kamtchatka, crater of Uzon Volcano S. of Kronotski Lake, leg. V. P. SAVICZ, 

1909 (no. 6419) (LE, lectotypusS8; UPS, isolectotypus). 

Icon.: LAMB 1972, figs. 2, 3, 4, 5, 6. 

Mat. chim. : atranorin and porphyrilic acid. 

Distrib.: arctic-subarctic, probably circumpolar. Greenland, Jan Mayen, U.S.S.R. (Kamtchatka) 

Canada (N. W. Territ., QuBbec), U. S. A. (Alaska). Unverified record: U.S.S.R., Lake Baikal (RAS- 

SADINA, 1936, as St. denudatum f. arenarium). 

Remarks: morphologically similar to St. arcticum LYNGE, but fundamentally different 

in chemical constitution; the PD reaction is faint yellow. 

s8 Lectotypified by LAMB, loc. cit.


43. Stereocaulon cornutum MULL. ARG. 

Flora, 69: 252 (1886); VAIN. Ann. Acad. Sci. Fenn. ser. A, 6 (7): 24 (1915). 

Typus: Jamaica, mountains near Gordon-Town, leg. J. HART (date ?), commun. W. JOSHUA 

(G-M~~LL, holotypus, not seen; K, isotypus; probably the same isotype material distributed in LOJKA, 

Lichenotheca Univ. no. 154). 

Exsicc.: LOJKA, Lichenotheca Univ. Fasc. IV (1886) no. 154 (probably isotype material) (LD, s, 

WELC). - MERR. Lich. Exs. Fasc. V (191 1) no. 121, as Sf. pityrizans (c, DUKE, FH, L, US). - ZAHLBR. 

Lich. Rarior. Exs. no. 283 (1931) (s). 

Icon.: PLITT 1935, fig. 3 (field photograph). 

Mat. chim.: atranorin, stictic acid, norstictic acid. 

Distrib. : Greater Antilles group of the West Indies (Jamaica, Dominican Republic) and Venezuela; 

the variety corallizans apparently endemic to Central America (Costa Rica). Unverified record: 

Japan (MULLER ARC., 1891). DODGE (1929) and SAT^ (1943, 1965) have suggested that this record 

should refer to St. nabewariense ZAHLBR. (=St. vesuvianum var. nodulosum (WALLR.) LAMB). Although 

we have not seen the specimen in question (leg. MIYOSHI, no. 126) we would regard this opinion 

as undoubtedly correct. 

Remarks: very widespread and abundant in Jamaica at altitudes of 11W2250 m s.m. 

(PLITT, 1935). 

Var. corallizans LAMB (n. var.) 

Diagn.: Differt a var. typica phyllocladiis apicalibus breviter digitato-coralloideis, haud 

impresso-maculatis, et apotheciis nonnihil minoribus. 

Typus: Costa Rica, prov. Heredia, Cerro Central de Zurqui, altit. 1520-1600 m s. m., leg. DODGE, 

VALERIO, THOMAS and VALERIO, 1929 (no. 6119) (s, holotypus; CAN, UPS, isotypes). 

Remarks: seen from a number of localities in Costa Rica at altitudes of 1500 to 2500 m s.m. 

44. Stereocaulonjibrillosum LAMB (n. sp.) 

Diagn. : Pseudopodetia congregata, stipato-caespitosa, arcte affixa, simplicia aut tantum 

parcissime ramosa, erecta, decorticata, glabra, albida vel pallide ochracea, apicibus cornutoattenuata. 

Phyllocladia numerosa, conferta partimve magis dispersa, albicantia, initio 

minute granuliformia, mox elongata et cylindrico-coralloidea, ad 2-3 mm longa, 0.15-0.20 rnrn 

crassa, teretia, patento-divaricata, breviora simplicia, longiora semel aut compluries ramosa, 

apices pseudopodetiorum versus gradatim breviora et postremo granulosa vel papillata. 

Cephalodia nurnerosa, lateraliter sessilia, subglobosa, ad 1 mm lata vel paullo majora, olivaceoviridia, 

fusca aut nigricantia, sat pellucida, majora subbotryosa. Apothecia sparsa, lateralia, 

sessilia et brevissime pedicellata, parva, ad 1 mm lata, primum plana et margine proprio 

fusco circumdata, deinque leviter convexa et irnmarginata, disco fusconigricanti. Hypothecium 

haud pigrnentiferum. Hymenium 70-75 ,u altum, ascis cylindrico-clavatis (sporae 

maturae haud visae). Atranorinum et acidum sticticum continet (KSflavescens, PDS 

aurantiaco-rubescens). 

Typus: Madagascar, N. E. region, Marojejy massif, E. summit, altit. 2100 m s. m., on rocks, leg. 

H. DES ABBAYES, 1956 (no. 3081) (FH, holotypus; REN, isotypus). 

Mat. chim. : atranorin and stictic acid demonstrated in the holotype. 

Distrib.: Madagascar. Known only from the type material. 

Remarks: belongs to the furfuraceum-nigromaculatum assemblage, and closely related 

to St. anomalum LAMB, from which it differs in the longer, well branched phyllocladia resembling 

those of St. exutum NYL.; phyllocladia of the flattened, dorsiventral, squamulose-


subfoliose type are completely absent. Cephalodial algae Stigonema. 

45. Stereocaulon furfuraceum DUVIGN. 

Lejeunia, Mem. no. 14: 73 (1956). 

Typus: Africa, Rwanda Burundi, Karisimbi Volcano, altit. 3100m s. m., "sur pierre en la zone 

?I Hagenia, Senecio, Lobelia et Bruykres arborescentes", leg. DE WITTE, 1935 (no. 39A) (BRLU, holotypus). 

Exsicc.: V~ZDA, Lich. Sel. Exs. Fasc. LIX (1977) no. 1463 (BRNU). 

Icon. : DUVIGN. 1956, figs. 18, 19. 

Mat. chim.: atranorin, stictic acid and norstictic acid. 

Distrib. : central Africa (Uganda, Zaire, Kenya, Rwanda Burundi). 

Remarks: has a general resemblance to St. cornutum MULL. ARG., St. scutelligerum TH. 

FR. and St. virgatum ACH. Distinction from St. nigromaculatum DUVIGN. may be difficult. 

46. Stereocaulon graminosum SCHAER. 

ex MORITZI, Systematisches Verzeichniss der von H. Zollinger . . . auf Java gesammelten Pflanzen: 

127 (1845-46). - Stereocaulon turgescens NYL. Synopsis Methodica Lichenum, 1 (2): 248 (1860) 

(nom. illegit.). 

Typus: Java, "in graminosis Pangerango 9000' ", leg. ZOLLINGER (date ?) (no. 1946) (G-SCHAER, 

holotypus, not seen by us; L, isotypus). This is also the lectotype material of St. turgescens NYL. 

(lectotypif. nov.). 

Exsicc. : ZAHLBR. Lich. Rarior. Exs. no. 373 (1936) (s). 

Mat. chim. : atranorin and stictic acid (the latter sometimes sparingly present or absent). 

Distrib. : Indonesia (Sumatra, Java, Bali, Lombok), North Borneo, Philippines. Unverified records: 

Ecuador (LEIGHTON, 1866, as St. turgescens); acc. to NYLANDER (1874) this refers to St. denudatum 

(=St. vesuvianum var. nodulosum (WALLR.) LAMB); W. Africa (NYLANDER, 1862, as St. turgescens). 

Remarks: closely related to St. vesuvianum PERS. but differing in the small size of the 

crowded, often substipitate, verruculose to finally more or less peltate phyllocladia. In this 

respect it more resembles St. obesum TH. FR., which is usually smaller in stature and has a 

different chemical constitution. 

47. Stereocaulon lavicola MAGN. 

ex MAGN. et ZAHLBR. Ark. Bot. 31A (6): 38 (1944). Emend. LAMB (emend. nov.). 

Typus: U.S.A., Hawaii, Molokai, Upper Maunahui camp, altit. ca. 1050 m s. m., on lava, leg. 0. 

SELLING, 1938 (no. 5848 pr. p.) (GB, lectotypus, lectotypif. nov.; see Remarks below). 

Mat. chim. : atranorin and norstictic acid (Fox, FOLLMANN and HUNECK, 1971). 

Distrib.: Hawaiian Islands. MAGNUSSON loc. cit. mentions also a specimen from Oahu (paratype) 

which we have not seen. 

Remarks: the specimen designated as "part of type" in GB (SELLING no. 5848) consists 

of 2 pieces, which are different; one (the larger, on lava) is St. octomerellum MULL. ARG.; 

the other (on soil) is the species here under consideration, and we have lectotypified St. 

lavicola (emend.) on it. The greater part of MAGNUSSON'S diagnosis seems to refer to this 

specimen, the red reaction with K (due to norstictic acid) being particularly mentioned, but 

some details in his description obviously refer to the specimen of St. octomerellum, for 

instance as regards the apothecia and spores. The lectotype of St. lavicola forms a continuous, 

crustose patch about 3 cm across, consisting of densely crowded phyllocladia and 

ends of pseudopodetia, the latter penetrating for a short distance into the soil as root-like


extensions, their upper parts congestedly and irregularly branched, corticated, glabrous, 

ochraceous-whitish. Phyllocladia borne on the pseudopodetial apices, grain-like to shortly 

and indistinctly papillose, not coralloid, crowded and sometimes confluent, without darker 

centers. No soredia. No cephalodia present. Only a few more or less immature apoth- 

ecia present, sessile among the phyllocladia, brown-blackish, plane, up to 0.6 mm diam. 

Hypothecium faintly pigmented (yellow-brown or brownish). Hymenium low (40-50 p), 

with asci 35-40x 10-13 p. Spores 6-8nae, straight, cylindric-fusiform, small, 17-26 x2.5 p, 

only 1-septate ones seen. The material is not very good, but undoubtedly represents a dis- 

tinct species and could be recognized if more and better material becomes available. The 

small apothecia have an internal structure similar to that of species of the vesuvianum-group, 

so that St. lavicola would appear to belong to sect. Denudata JATTA, in spite of the absence 

of darker centers in the phyllocladia. This disposition of the species must however remain 

provisional pending the discovery of better material. 

48. Stereocaulon leprocephalum VAIN. 

Ark. Bot. 8 (4): 35 (1909); MAGN. Goteborgs K. Vetensk.- o. Vitterh.-Samh. Handl. ser. 4, 30 (7): 71 

(1926). 

Typus: U.S.S.R., N. E. Siberia, Chukchi Nat. Okrug, Jinretlen Peninsula, lat. 67"05' N., long. 

173"15' E., on rock, leg. E. ALMOUIST, Exped. Vega, 1878-79 (TUR-VAIN, no. 3899, holotypus; s, isotypus). 


Distrib.: arctic, probably circumpolar; at present known only from N. E. Siberia and E. arctic 

Canada (N. W. Territ., Baffin Island). 

Remarks: first mentioned in the literature as "Stereocaulon evolutum" by ALMQUIST 

(1879). VAINIO (1909) wrongly ascribed to the present species a specimen from Sri Lanka 

(Ceylon) recorded by NYLANDER (1900) as "Stereocaulon condyloideum", which is actually 

St. austroindicum LAMB. The finding of fertile material in Canada by HALE allows the species 

to be placed in the sect. Denudata JATTA, as the small lateral apothecia of lecideine appearance 

are characteristic. Morphologically and chemically, St. leprocephalum seems to be closely 

related to St. arenarium (SAV.) LAMB. 

49. Stereocaulon nigromaculatum DUVIGN. 

Lejeunia, MBm. no. 14: 64 (1956); LAMB ex FREY, Bot. Jahrb. Syst. 86: 220 (1967). 

Typus: Africa, Uganda, Ruwenzori, Camp Emin, altit. 3900 m s. m.,3O "sur roches nues avec 

Umbilicaria Haumaniana", leg. HAUMAN, 1932 (no. 932) (BRLU, holotypus; FH, isotypus). 

Facultative synonyms: Stereocaulon ruwenzoriense DUVIGN. Lejeunia, MBm. no. 14: 62 (1956). - 

Stereocaulon microthuja DUVIGN. Lejeunia, M6m. no. 14: 56 (1956), with fac. abietina DWIGN. 

Exsicc.: V~ZDA, Lich. Sel. Exs. Fasc. LVI (1976) no. 1399 (BRNU). 

Icon. : DUVIGN. 1956, figs. 1, 5, 7, 14, 15, 16, 17 ; figs. 12, 13, as St. microthuja. 

Mat. chim.: atranorin, stictic acid, usually also norstictic acid, occasionally ursolic acid 

(a triterpene), in addition to several unidentified substances. 

Distrib. : central Africa (Uganda, Rwanda Burundi, Zaire, Tanzania). 

Remarks: St. ruwenzoriense and St. microthuja represent a dwarfed or juvenile condition 

of St. nigromaculatum. 

30 2900 m acc. to descr. orig., but 3900 m acc. to the label.


Fo. fastigiato-pulvinatum LA^ (n. f.) 

Syn. Stereocaulon microthuja fac. fastigiato-pulvinata DWIGN. Lejeunia, M6m. no. 14: 61 (1956) (nom. 

inval., in lingua Latina haud descriptum). 

Diagn. : Habitu dense fastigiato-ramosum et pulvinatum, phyllocladiis apicalibus crustam 

superficialem formantibus. 

Typus: Africa, Uganda, Ruwenzori, Mt. Stanley, altit. 4350 m s. m., leg. HAUMAN, 1932 (no. 927) 

(BRLU, holotypus). 

Icon. : DUVIGN, 1956, fig. 12A (I), as St. microthuja fac. p~lvinata.'~ 

Remarks: analogous to the f. umbonatum (WALLR.) LAMB of St. vesuvianum var. nodulosum. 

Seen from 2 localities in Uganda, between 3500 and 4350 m altitude. 

50. Stereocaulon obesum TH. FR. 

De Stereoc. et Pilophor. Comment.: 28 (1857); Monogr. Stereoc. et Pilophor.: 49 (1858). 

Typus: Costa Rica, prov. Cartago, Volcan Irazu, altit. 11000 feet (3400 m s. m.), "pumici innascens", 

leg. A. ~RSTED (date ?) (UPS-TH FR, holotypus). 

(Exs~cc.'~) 

Icon.: TH. FRIES 1858, P1. X, fig. 1. 

Mat. chim. : atranorin and lobaric acid; frequently in a deficient phase with atranorin only. 

Distrib. : Central America (Guatemala to Panama) and South America (Ecuador). Unverified record: 

Greenland (KREMPELHUBER, 1868), almost certainly not this species. Erroneous record: Argentina 

(RASANEN, 1939), =St. vesuvianum PERS. 

Remarks: morphologically resembles St. graminosum SCHAER., but less robust. Usually 

esorediate, but occasionally some of the pseudopodetia may be dissolute into effuse farinose 

soredia at the tips. 

5 1. Stereocaulon pityrizans NYL. 

Ann. Sci. Nat. Bot. ser. 4, 11: 209 (1859); Synopsis Methodica Lichenum, 1 (2): 234 (1860); DODGE, 

Ann. Cryptog. Exot. 2 (2): 135 (1929). 

Typus: Peru, prov. Carabaya, "in schistosis argillaceis", leg. WEDDELL, 1847 (H-~n, no. 40143, 

holotypus; PC, isotypus). 

Mat. chim.: atranorin, stictic acid, norstictic acid. 

Distrib.: South and Central America: Peru, Colombia, Costa Rica. 

Remarks: when fertile, can be easily recognized by the lateral or terminal apothecia of 

lecanorine appearance, with conspicuous, light-colored pseudothalline margin. This misled 

ZAHLBRUCKNER (1926-27) to include the species in subgen. Lecanocaulon. Occasionally the 

apothecia may proliferate to form composite heads of small, immarginate, secondary apothecia. 

Phyllocladia typically minute and grain-like, 0.1-0.3 mrn diam., many with distinct darker spot. 

52. Stereocaulon scutelligerum TH. FR. 

Flora, 44: 412 (1861) (ut nom. nov.); DODGE, Ann. Cryptog. Exot. 2 (2): 141 (1929). 

Typus: R6union Island Porbonia), no exact locality indicated, leg. BORY DE SAINT-VINCENT 

(date ?) (UPS-TH FR, holotypus; PC, isotypus). 

Facultative synonyms: Stereocaulon vulcani subsp. richardianum MONT. ex TH. FR. Monogr. 

'O In the legend to fig. 12, the subfigures 1 and 2 have been accidentally transposed. 

Tbe material distributed as "Stereocaulon obesum" in Lich. Austroamer. ex Herb. RegneN. no. 391 

is not this species, but St. verruciferum NYL. var. surreptans (LAMB) LAMB.


Stereoc. et Pilophor.: 44 (1858). - Stereocaulon richardianum (MoNT.) ZAHLBR. Catalogus Lichenum 

Universalis, 4: 666 (1927).42 

Exsicc.: ABB. Lich. Madagasc. et Borbon. Sel. Exs. Fasc. 11 (1966) no. 36 (FH, REN). 

Icon.: TH. FRIES 1858, PI. IX, fig. 3, as St. vulcani. - NYL. 1860, PI. VII, fig. 7 a-c, as St. salazinum. 

Mat. chim. : atranorin and norstictic acid. 

Distrib. : Rkunion, Mauritius, Madagascar. 

Remarks: great nomenclatorial confusion prevailed until TH. FRIES designated this 

species with the new name St. scutelligerum, on account of the fact that part of BORY'S original 

material of Lichen salazinus was a mixture of 2 species; see remarks under St. salazinum (p. 

302). The confusion was for a time further compounded by TH. FRIES'S misapplication 

(1857, 1858) of the name St. vulcani (BORY) to the present species; the true St. vulcani is a 

different species (p. 257). The variability of St. scutelligerum appears to be slight; St. vulcani 

subsp. richardianum (MoNT.) TH. FR. refers to an aged and denuded growth-state with entirely 

decorticated, smooth and glabrous pseudopodetia, the phyllocladia almost all grain-like 

or verruculose. Cephalodia have not been found in St. scutelligerurn, and probably do not 

occur. The closest relationship is obviously with St. virgatum AcH., a West Indian and 

Central American species, and with St. anomalum LAMB (Madagascar and central Africa). 

53. Stereocaulon subdenudatum HAV. 

Lich. Exs. Norveg. no. 658 (1947) (nomen nudum); Arbok Univ. Bergen, Naturvitensk. Rekke, no. 12: 

13, 20 (1954) (cum descript.). 

Typus: Norway, Hordaland, par. Granvin, Smareggfjellet, between Baorhaug and Plantefeltet; 

3 localities mentioned, one of them near Nyadaemmo; leg. J. HAVAAS, 1940. As lectotype material 

we select the specimen from Nyadaemmo distributed in HAV. Lich. Norveg. Occid. no. 232 (FH, UPS- 

MAGN, isolectotypes; lectotypif. nov.). 

Exsicc.: HAV. Lich. Exs. Norveg. Fasc. XXI (1947) no. 658 (BG). - HAV. Lich. Norveg. Occid. 

Exs. Fasc. X (1948) no. 232 (FH, UPS-MAGN, isolectotype material). 

Mat. chim.: atranorin, stictic acid, norstictic acid, constictic acid, and some unidentified 

substances. 

Distrib.: Norway (Hordaland, Nordland); Scotland (Argyllshire, Perthshire, Angus). 

Remarks: distinguished by HAVAAS from St. denudatum (St. vesuvianum var. nodulosum 

(WALLR.) LAMB) on the basis of form of phyllocladia and a different shade of color in the 

living condition. G. DEGEL~US studied living populations on Vega Island, Norway, Nordland, 

in 1974, and informed us (in litt.) that this species, growing there together with St. vesuvianum 

var. nodulosum, is distinctly different in color in the living state. This color is a slightly 

brownish gray, not glaucous-gray or glaucous-whitish as in St. vesuvianum var. nodulosum. 

The difference is however only obvious in the fresh state. Morphologically, St. subdenudatum 

closely resembles St. vesuvianum var. nodulosum f. umbricolum (FREY) LAMB. The species 

requires further investigation. St. denudatulum NYL., here regarded as a doubtful species 

(p. 322), may possibly be synonymous. 

54. Stereocaulon symphycheilum LAMB 

Bot. Not. 114 (3): 271 (1961): POELT, Bestimmung~~chliisseI Europaischer Flechten: 633 (1969); MOT. 

Flora Polska, 3 (2): 392 (1964); DAHL et KROG, Macrolichens of Denmark, Finland, Norway and Sweden: 

131 (1973). 

42 Probably a lapsus rather than an intentional new combination.


Typus: Sweden, Lycksele Lappmark, par. Tarna, Hemavan (Bjorkfors), Syterbacken, reg. subalp., 

altit. 450-480 m s. m., on rocks beside river, leg. V. AHMADJIAN, 1959 (no. 26) (FH, holotypus; UPS, 

isotypus). 

Icon. : LAMB 1961, figs. 3a, b, c, 4a, b. 


Distrib. : Fennoscandia (Sweden, Norway, Finland), Canada (Quebec), U. S. A. (Alaska), U.S.S.R. 

(Kamtchatka). 

55. Stereocaulon vanoyei DUVIGN. 

Biol. Jaarb. "Dodonaea", 8: 162 (1941) ("van Oyei"); ibid., 9: 82, 86 (1942). 

Typus: Iceland, Krisuvik, "ad lavam inter muscos", leg. P. VAN OYE, 1938 (no. 44a) (BRLU, 

holotypus; FH, isotypus). 

Icon. : DUVIGN. 1941, fig. 6. 

Mat. chim. : atranorin, norstictic acid, connorstictic acid, and an unidentified substance 

with also occurs with norstictic acid and connorstictic acid in Parmelia dierythra HALE.^' 

Distrib. : Iceland. Known from 4 collections. 

Remarks: pseudopodetia prostrate, dorsiventral; phyllocladia up to 2.5 rnrn diam., 

whitish, convex, shield-shaped, confluent, unicolorous or sometimes with several minute, 

faintly indicated, darker spots on the surface. Apothecia sessile on the phyllocladia, small, 

brown-blackish; hypothecium faintly brown; spores 3-septate, ca. 45 x 3 p. Morphologically 

resembles St. vesuvianum var. nodulosum f. depressum (MAGN.) LAMB and St. glaucescens TUCK. 

56. Stereocaulon verruciferum NYL. 

Synopsis Methodica Lichenum, 1 (2): 248 (1860); DODGE, Ann. Cryptog. Exot. 2 (2): 118 (1929); Ms. 

Revista Univ. (Santiago), 22: 200 (1937). 

Typus: Colombia, no exact locality indicated, leg. J ~ESON (date ?); also mentioned from "America 

aequinoctiali", leg. BONPLAND (date ?). We select the former specimen, H-NYL no. 39813 pr. p., 

as lectotype (lectotypif. nov.). 

Facultative synonym: Stereocaulon speciosum LAMB, Farlowia, 4: 458 (1955). 

Exsicc.: FOLLM. Lich. EXS. Sel. Mus. Bot. Berol. Fasc. I (1968) no. 15, as St. speciosum (FH). 

Icon. : LAMB 1955, fig. 24; 1959, P1. VIII, fig. 29; as St. speciosum. 

Mat. chim. : atranorin, stictic acid, usually also norstictic acid. 

Distrib. : Argentina and Chile (Cordilleran Great Lakes region), Ecuador, Colombia. Unverified 

record: Peru (LINDAU, 1909). 

Remarks: has been confused with St. obesum TH. FR. (TH. FR. 1861; Lich. Austroamer. 

ex Herb. Regnell. no. 391), which is a much smaller and chemically different species. 

Var. surreptans (LAMB) LAMB (n. comb.) 

Basionym: Stereocaulon speciosum var. surreptans LAMB, Farlowia, 4: 459 (1955). 

Typus: Argentina, Rio Negro, Cerro Rigi near Lago Frias, altit. ca. 1650 m s. m., leg. I. M. LAMB, 

1950 (no. 6098) (CAN, holotypus). 

Exsicc.: Lich. Austroamer. ex Herb. Regnell. Fasc. XVI (1943) no. 391, as St. obesum (CAN, LD, 

LIL, S, WIS). - V~ZDA, Lich. Sel. Exs. Fasc. XXII (1967) no. 545, as St. "speciorum" var. surreptans 

(pH). 

43 ACC. to C. CULBERSON in litt. 1974.


57. Stereocaulon virgatum ACH. 

ex SPRENG. Caroli Linnaei . . . Systema Vegetabilium, Editio Decima Sexta, 4 (1): 275 (1827); MULL. 

ARG. Flora, 70: 286 (1887); VAIN. Ann. Acad. Sci. Fenn. ser. A, 6 (7): 26 (1915). 

Typus: Leeward Islands, Guadeloupe, leg. ?, date ? (H-ACH, holotypus). 

Facultative synonyms: Stereocaulon virgatum f. achariana VAIN. J. Bot. 34: 34 (1896). - Stereocaulon 

virgatum f. applanata VAIN. loc. cit. - Stereocaulon virgatum f. primaria VAIN. loc. cit. - Stereocaulon 

giltayi DUVIGN. Biol. Jaarb. "Dodonaea", 9: 87 (1942). - Stereocaulon sphaerophoroides TUCK. 

An Enumeration of North American Lichenes: 52, footnote (1845) (non auctt. plur.). 

Exsicc.: Lich. Exs. Univ. Colorado Mus. Fasc. IX (1971) no. 337 (LD). 

Icon.: DUVIGN. 1942, fig. 3A, B, as St. giltayi. 

Mat. chim. : atranorin, stictic acid and norstictic acid, the latter two substances occurring 

in varied concentration, either in approximately equal proportions or with one or other of 

them strongly preponderant or perhaps in some cases present to the total exclusion of the 

other. The macroscopic reactions are therefore variable: K+yellow then red, PD+yellow 

(norstictic acid predominating) or K+yellow, PD+orange-red (stictic acid predominating). 

VAINIO'S 2 forms achariana and primaria were distinguished on the basis of such different 

reactions. 

Distrib.: West Indies (Greater Antilles, Leeward Islands, Windward Islands) and Central America 

(Nicaragua). A specimen said to be from Venezuela, but with this origin queried by TH. FRIES, leg. 

FUNCK and SCHLIM (date ?) is present in UPS-TH FR. St. giltayi DUVIGN. was described as from Scotland, 

certainly as the result of an error. 

Remarks: St. virgatum is closely related to St. scutelligerum TH. FR., and like the latter 

is usually destitute of cephalodia. Cephalodia have however been seen in a few specimens 

from Dominica and Guadeloupe. They are of the anatomically undifferentiated, "spherical" 

to "botryose" type normally associated with species of the section Denudata. TH. FRIES 

(1857, 1858) and NYLANDER (1860) erroneously stated that St. virgatum is a synonym of 

St. furcatum FR. The latter is referable to St. ramulosum (Sw.) RAusc~., as was pointed out 

by MULLER ARG. (1887), and confirmed by RIDDLE (ex DODGE, 1929) and by the present 

author. Stereocaulon sphaerophoroides TUCK., the commonly used name for a species endemic 

to the Azores, Madeira and Canary Islands, is a synonym of St. virgatum; see discussion on 

p. 206. 

58. Stereocaulon vulcani (BORY) ACH. 

Lichenographia Universalis: 583 (1810). - Lichen vulcani BORY, Voyage dans les Quatre Principales 

Iles des Mers d'Afrique, 3: 147 (1804). - Stereocaulon denudatum var. vulcani (BORY) NYL. Synopsis 

Methodica Lichenum, 1 (2): 248 (1860). 

Typus: R6union Island (Borbonia), no exact locality given, leg. BORY DE SAINT-VINCENT (date ?). 

There are several mounts in PC-HUE (ex herb. RICHARD) which qualify as type material44. As lectotype 

we have selected the best of these (herb. HUE no. 257 pr. p.), labelled "Herb. Richard Stereocaulon 

Vulcani Ile de Bourbon. Bory" (lectotypus, lectotypif. nov.). 

Facultative synonyms: Stereocaulon flavireagens GYELN. Ann. Cryptog. Exot. 4: 173 (1931). - 

Stereocaulonflavireagens f. cinerascens MAGN. ex MAGN. et ZAHLBR. Ark. Bot. 31A (6): 43 (1944). - 

Stereocaulonflavireagens f. densum MAGN. op. cit. p. 42. 

Exsicc.:" ABB. Lich. Madagasc. et. Borbon. Sel. Exs. Fasc. I1 (1966) no. 38 (Ch. str. I) (FH). - 

HEPP, Flechten Europas Band I, Heft 1 (1853) no. 2 pr. min. p., as St. denudatum B. vesuvianum (Ch. str. 

44 DODGE (1929, p. 119) erroneously states the type of Lichen vulcani to be from Mauritius. 

46 V~ZDA, Lich. Sel. Exs. no. 415, as St.flavireagens, is not this species, but St. rubiginosum PERS.


11) (FH-TUCK, mixed with St. vesuvianum PERS. But in FH and PC it is entirely St. vesuvianum). - 

SCHAER. Lich. Helvet. Exs. Fasc. XXVII (date ?) no. 652, as St. denudaturn B. vesuvianurn (Ch. str. 11) 

(UPS). 

Mat. chim. : 2 chemical strains exist : 

Strain I, with atranorin, stictic acid, and smaller amounts of norstictic acid (Kfpersistent 

yellow, PDforange-red). The typical strain. 

Strain 11, with atranorin, norstictic acid, possibly also salazinic acid in low concentration; 

no stictic acid present (Kfdeep yellow then usually turning red, PDfpersistent deep yellow). 

The "flavireagens"-strain. 

Both strains occur in Hawaii, the Azores and Canary Islands, sometimes growing closely 

associated. The chemical situation is similar to that found in St. massartianum HUE (see 

p. 273). 

Distrib.: Ch. str. I: Rtunion, Africa (Zaire, Rwanda-Burundi), Canary Islands, Azores, Hawaii. 

Ch. str. 11: Canary Islands, Azores, Hawaii. 

Remarks: the species is variable as regards form and color; St. javireagens ff. cinerascens 

and densum MAGN. are insignificant variations. TH. FRIES, up to 1861, erroneously applied 

the name vulcani to St. scutelligerum TH. FR. The heterogeneous type material of St. cereolinum 

ACH. (see Append. 2, p. 327) consists, according to TH. FRIES (1871, p. 56), partly of 

St. vulcani. St. vulcani, after various algae, is one of the earliest pioneers of plant life on 

recent lava flows. Colonization studies in Hawaii have been made by JACKSON and KELLER 

(1970a), JACKSON (1971) and SMATHERS and MUELLER-DOMBOIS (1974). JACKSON (1969) and 

JACKSON and KELLER (1970) found that in Hawaii St. vulcani greatly accelerates the chemical 

weathering of recent basalt lava flows. The weathered rock underneath the thallus was 

found to be a form of ferric oxide chemically different from the haematite occurring as weathering 

product on the lichen-free basalt lava rocks. 

Fo. maunae-loae (MAGN.) LAMB (n. comb.) 

Basionyrn: Stereocaulon Maunae Loae MAGN. et ZAHLBR. Ark. Bot. 31A (6): 40 (1944). 

Typus: U.S.A., Hawaii, Hawaii, Mauna Loa, above rest house, altit. 3400 m s. m., leg. C. SKOT- 

TSBERG, 1922 (no. 1729) (GB, holotypus; S, isotypus). 

Remarks: a compact pulvinate form of cauliflower-like habitus with the phyllocladia 

forming an apical crust. Analogous to the f. umbonatum (WALLR.) LAMB of St. vesuvianum 

var. nodulosum. Seen from Hawaii and the Azores; all specimens in the Ch. str. 11. 

59. Stereocaulon weberi LAMB (n. sp.) 

Diagn.: Pseudopodetia erecta, stipato-caespitosa, substrato arcte affixa, singulatim 

radicata (haptero commune deficienti), simplicia aut (praesertim parte superna) subdichotomice 

ramosa; longit. 4-5 cm, crassit. 0.8-1.5 mm, teretia, apices versus sensim attenuata ibique 

obtuse subulata; tota longitudine plus minusve uniformiter phyllocladiis vestita, superficie 

interjacenti glabra, albicanti, laevi opacaque. Phyllocladia minute granuliformia, vulgo 

haud ultra 0.1 rnm lata, sorediiformia, globulosa, albescentia, conferta; basin pseudopo- 

detiorurn versus nonnihil majora, ad 0.2(-0.3)rnm lata, crassiuscule peltato-squarnulosa, 

albido-glaucescentia. Thallus primarius haud visus. Cephalodia numerosa, minuta, in- 

conspicua, fusconigricantia, irregulariter pulvinata, 0.2-0.4 (-0.8) mm lata, superficie obsolete 

tuberculata, Stigonema continentia. (Apothecia desunt.) Atranorinum et acidum lobaricum 

continet (Kfflavescens, PDfsulphureo-flavescens). - St. obeso TH. FR. affine, a quo phyllo-


cladiis minimis et granuloso-sorediiformibus distat; saltem apicalia sorediis veris censenda. 

Typus: Ecuador, Galipagos Islands, Isla Santa Cruz, summit of Mt. Crocker, altit. 800 m s. m., 

locally abundant on bare rocks, leg. W. A. WEBER and J. LANIER, 1976 (FH, holotypus; COLO, isotypus. 

Isotype material distributed in Lich. Exs. Univ. Colorado Mus. no. 494). 

Exsicc. : Lich. Exs. Univ. Colorado Mus. Fasc. XI11 (1976) no. 494, as St. microcarpum (EH) (isotype 

material). 

Mat. chim.: atranorin, lobaric acid, possibly a trace of norlobaric acid, and an unidentified 

substance. 

Distrib. : Galipagos Islands. Known only from the type collection. 

Remarks: closely related to St. obesum TH. FR., and with similar chemical constitution, 

but differing in its uniformly smaller, granulose-sorediiform phyllocladia. In St. obesum 

soredia, when present, are restricted to the extreme tips, and the phyllocladia, although small, 

are not sorediiform. 

Subsect. B. Phyllocaulon (TUCK.) LAMB 

60. Stereocaulon wrightii TUCK. 

Amer. J. Sci. Arts, ser. 2, 28: 202 (1859); A Synopsis of the North American Lichens, Part I: 234 

(1882); RIDD. Bot. Gaz. 50: 302 (1910); MAGN. Gijteborgs K. Vetensk.- o. Vitterh.-Samh. Handl. ser. 

430 (7): 77 (1926); DODGE, Ann. Cryptog. Exot. 2 (2): 149 (1929); YOSHIM. Lichen Flora of Japan in 

Colour: 179 (1974). - Stereocladium wrightii (TUCK.) NYL. ex HUE, Nouv. Arch. Mus. Hist. Nat. ser. 

3,2: 245 (sep. 37) (1890). - Phyllocaulon wrightii (TUCK.) VAIN. Ark. Bot. 8 (4): 36 (1909). 

Typus: U.S.S.R., E. Siberia, Arakamtchetchene Island on E. side of Bering Straits, leg. C. WRIGHT, 

1858 (FH-TUCK, holotypus; US, isotypus). 

Icon.: see under the following species, St. apocalypticum, which is morphologically indistinguishable. 

Mat. chim.: atranorin and stictic acid. 

Distrib. : U.S.S.R. (Bering Straits region, Extreme Orient) and Japan (Honshu). Most records in 

the literature refer to the following species (St. apocalypticum). 

Remarks: rarer than St. apocalypticum NYL. Although they are fundamentally different 

in chemistry, there is no detectable morphological or anatomical difference between St. 

wrightii and St. apocalypticum, and at least in Japan they may occur together (cfr. SAT^, 1965a). 

6 1. Stereocaulon apocalypticum NYL. 

ex MIDDENDOREF, Reise in den aussersten Norden und Osten Sibiriens, 4, Anhang VI: Iv, footnote 

(1867); YOSHIM. Lichen Flora of Japan in Colour: 178 (1974). - Stereocladium apocalypticum (NYL.) 

NYL. Flora, 58: 302 (1875). 

Typus: U.S.S.R., E. Siberia, Stanovoi Range (border of Amur Oblast and Yakut Autonomous 

SSR), Ket-Kat Mts., leg. MIDDENDORFF, 1844 (H-NYL, no. 40146, holotypus). 

Facultative synonym: Stereocaulon foliiforme HUE, Bull. SOC. Bot. France, 54: 414 (1907). 

Exsicc.: ASAH. Lich. Japon. Exs. Fasc. VI (1959) no. 291 (UPS, US). -ELENK. Lich. Flor. Ross. 

Fasc. IV (1910) no. 163, as Stereocladium wrightii (GB, PR). - KUROK. Lich. Rarior. et Crit. Exs. Fasc. 

I (1966) no. 47 (FH, LD). - SAV. Lichenotheca Ross. Decas XVII (1967) no. 165, as Stereocladium 

apocalypticum (LE). - ZAHLBR. Lich. Rarior. Exs. no. 374 (19361, as St. wrightii (EH, PR, s). 

ASAH. 1936, figs. 48, 49, 50, as St. wrightii. - ELENK. 1909, PI. I, figs. 10, 10a, as Stereocladium 

apocalypticum. - HUE 1907, figs. 48, 50, as St. foliiforme. - R ~D. 1910, fig. 3, as St. wrightii. 

1974, fig. 77f, g; PI. 33, fig. 334. - ZAHLBR. 1927, 

- SAT^ 1941, figs. 20,21, as St. wrightii. - YOSHIM. 

48 Some of these illustrations may refer to the morphologically indistinguishable St. wrightii TUCK.


PI. XI, fig. 2, as St. folii/orme. 


Distrib.: U.S.S.R. (S. E. Siberia, Amur, Yakutia, Taimyr, Transbaikalia, Kamtchatka, Extreme 

Orient, Sachalin), Manchuria, Japan (Hokkaido, Honshu), U.S.A. (Alaska). 

Remarks: according to ASAHINA (1936, as St. wrightii) it is common in alpine and subalpine 

districts of central and northern Japan; distribution map in SAT^ (1965a). Sun and 

shade forms occur, differing in compactness of tufts and breadth of foliar expansions (ASAHINA, 

loc. cit.). Apothecia are rare; described by SAVICZ (1923, p. 175) and ASAHINA (1936, p. 806). 

Subsect. C. Botryoideum (G. T. JOHNS. em. LAMB) LAMB 

62. Stereocaulon exutum NYL. 

Lichenes Japoniae : 18 (1 890); DODGE, Ann. Cryptog. Exot. 2 (2) : 120 (1929) ; SAT^ in NAKAI et HONDA, 

Nova Flora Japonica, Cladoniales (I): 67 (1941); ASAH. J. Jap. Bot. 36 (7): 225 (1961); YOSHIM. Lichen 

Flora of Japan in Colour: 170 (1974). 

Typus: Japan, Honshu, Shizuoka Pref., Fujiyama, Itchigome, altit. 1650 m s. m., "supra quisquilia 

vegetabilia", leg. E. ALMQUIST, 1879 (H-NYL, no. 39989, holotypus; s, isotypus). 

Facultative synonyms: Stereocaulon subramulosum MULL. ARC. Flora, 74: 108 (1891). - Stereocaulon 

subramulosum f. approximans HUE, Nouv. Arch. Mus. Hist. Nat. ser. 3,lO: 248 (sep. 36) (1898). 

- Stereocaulon exutum f. approximans (HUE) DODGE, Ann. Cryptog. Exot. 2 (2): 121 (1929). - 

Stereocaulon subramulosum f. complanatum HUE, NOUV. Arch. Mus. Hist. Nat. ser. 3, 10: 248 (sep. 36) 

(1898). - Stereocaulon exutum f. complanatum (HUE) DODGE, Ann. Cryptog. Exot. 2 (2): 121 (1929). 

E~sicc.'~: KUROK. Lich. Rar. et Crit. Exs. Fasc. 111 (1971) no. 140 (FH, LD). 

Icon.: ASAH. 1961b, fig. 2. - DUVIGN. 1956, fig. 11 (section of phyllocladium). - SAT^ 1941, fig. 

24. - YAS. 1925, PI. XIII, fig. 2, as St. subramulosum. - YOSHIM. 1974, fig. 76a; PI. 32, fig. 307. 


Distrib.: Japan (Honshu, Shikoku, Kiushiu, Hokkaido), Korea, U. S. S. R. (Extreme Orient), 

Australia (N. S. Wales). 

Remarks: the graceful, well branched phyllocladia of this species are usually quite teretecoralloid 

and look like disharmonic pseudopodetial branchlets, but DUVIGNEAUD (1956) has 

shown that their anatomical construction is distinctly dorsiventral. The formae approximans 

and complanatum HUE appear to be insignificant variations. The hypothecium of St. exutum 

is colorless or often stained yellow by some unidentified substance, never brown-pigmented. 

The cephalodia are distinctly verruculose-botryose, with Stigonema in all specimens examined. 

The single known Australian specimen (G-~LL, mentioned also by DODGE, 1929, p. 120) 

is from the Snowy Mountains (Muniong Range of N. S. Wales), leg. C. MOORE, prior to 1895, 

and is perfectly typical. The Russian specimen (MSK) is from the Pacific seaboard, Primorskii 

Krai, near Shkotovo, leg. V. VOROSCHILOV, 1952. ASAHINA (1961b) states St. exutum 

to be "very common in Hondo, Shikoku and Kiusiu." 

63. Stereocaulon commixtum (ASAH.) ASAH. 

J. Jap. Bot. 44: 263 (1969); YOSHIM. Lichen Flora of Japan in Colour: 173 (1974). - Stereocaulon 

aponicurq var. commixtum ASAH. J. Jap. Bot. 35: 290 (1960). 

Typus: Japan, Honshu, prov. Owari, along Kisogawa River, on rock, leg. Y. ASAHINA, 1938 (no. 

1200) (ASAH, holotypus). 

Icon.: ASAH. 1960, fig. 2A, as St. japonicum var. commixtum. - ASAH. 1969, figs. 3, 4. - YOSHIM. 

47 ASAH. Lich. Japon. Exs. no. 43, as St. exutum, is the closely related species St. nigrum HUE.


1974, fig. 76g; PI. 32, fig. 316. 

Mat. chim.: atranorin, stictic acid, norstictic acid, miriquidic acid, and 7 unidentified 

substances, acc. to chromatography by C. CULBERSON on a fragment of the holotype. ASAHINA 

(1960, 1969) distinguished this taxon from St. japonicum TH. FR. on the basis of a morphological 

difference (longer phyllocladia) and the presence, in addition to stictic and norstictic acids, 

of an unidentified substance which he characterized crystallographically and stated to be the 

same as an unidentified substance found by him in St. curtatum NYL. This "materia propria" 

can now be identified as miriquidic acid, which occurs also in St. curtatum. 

Di-strib.: Japan. ASAHINA records this taxon from 9 localities in Honshu and Shikoku. 

Remarks: the autonomy of this species is questionable. Its morphology seems to fall 

within the normal range of variation for St. japonicum TH. FR. The miriquidic acid which 

occurs in addition to stictic and norstictic acids (the normal constituents for St. japonicum) 

may be an accessory substance without taxonomic significance. 

64. Stereocaulon curtatoides ASAH. 

J. Jap. Bot. 44: 267 (1969); YOSHIM. Lichen Flora of Japan in Colour: 172 (1974). 

Typus: Japan, Kiushiu, Yakushima, Hanano-ego, on granitic rock, leg. F. FUJIKAWA, 1933 (TNS, 

holotypus; FH, isotypus). 

Icon. : ASAH. 1969, fig. 7. - YOSHIM. 1974, fig. 76c; PI. 32, fig. 31 1. 

Mat. chim. : atranorin, stictic acid, norstictic acid, miriquidic acid. 

Disrrib. : Japan (Kiushiu). 

Remarks: similar in appearance to St. curtatum NYL., but with short, broad, 3-septate 

spores measuring 22-30 x 6.5-7.5 (-8.0) p and containing stictic and norstictic acids in addition 

to miriquidic acid. 

65. Stereocaulon curtatum NYL. 

Lichenes Japoniae: 18 (1890); SAT^ in NAKAI et HONDA, Nova Flora Japonica, Cladoniales (I): 75 

(1941); ASAH. J. Jap. Bot. 36 (7): 228 (1961); YOSHIM. Lichen Flora of Japan in Colour: 172 (1974). 

Typus: Japan, Honshu, Shizuoka Pref., Fujiyama, Itchigome, "in limite arborum", on rocks, 

leg. E. ALMQUIST, 1879 (H-NYL, no. 40134, holotypus; s, isotypus). 

Exsicc.: ASAH. Lich. Japon. Exs Fasc. I1 (1955) no. 93 (UPS). 

Icon. : ASAH. 1961 b, figs. 4,5. - SAT^ 1941, fig. 26. - YOSHIM. 1974, figs. 74b, 76d; PI. 32, fig. 312. 

Mat. chim. : atranorin and miriquidic acid. 

Distrib.: Japan (Honshu). Not a common species. 

Remarks: miriquidic acid was not known at the time of ASAHINA'S description (1961b) 

and he characterized it crystallographically as an unidentified substance. HUE, 1890-92 

(1892), p. 137, and some subsequent authors have confused this species with St. octomerum 

MULL. ARG., which has the same general habitus, but differs in the usually taller stature, 

narrower spores and chemical constitution. The spores of St. curtatum are broadly fusiform, 

3M0 x 6-7 (-9) p, (5-) 7-septate, and are unique in the genus in having occasionally 1-4 

longitudinal septa, thus suggesting a transition to the genus Argopsis TH. FR. The primary 

thallus is more or less persistent, an additional distinction from St. octomerum. Pycnoconidia 

bacilliform, straight or almost so, 4.M.5 x ca. 0.7 p. 

66. Stereocaulon dendroides ASAH. 

J. Jap. Bot. 36 (7): 231 (1961); YOSHIM. Lichen Flora of Japan in Colour: 172 (1974).


Typus: Japan, Shikoku, Mt. Kenzan, leg. F. FUJIKAWA, 1934 (no. 1207) (ASAH, holotypus; CAN, 

isotypus). 

Faclrltative synonym: Stereocaulon octomerum f. robustior ASAH. J. Jap. Bot. 36 (7): 230 (1961). 

Icon. : ASAH. 1961 b, figs. 7, 8. - YOSHIM. 1974, PI. 32, fig. 3 10. 

Mat. chim.: atranorin and porphyrilic acid (dendroidin). 

Distrib. : Japan (Honshu, Shikoku, Kiushiu). 

Remarks: we have been unable to find any reliable morphological or anatomical differentiation 

between this species and St. octomerum M ~ L ARC. . The substance dendroidin, 

described by ASAHINA (1961b) from this species, was proved to be identical with porphyrilic 

acid by Fox, MAASS and LAMB (1969). 

67. Stereocaulon etigoense (ASAH.) LAMB 

ex YOSHIM. Misc. Bryol. Lichenol. 6 (8): 135 (1974) ("echigoense"); YOSHIM. Lichen Flora of Japan in 

Colour: 173, 282 (1974). -. Stereocaulon japonicum subsp. etigoense ASAH. J. Jap. Bot. 35: 291 (1960). 

Typus: ASAHINA listed 7 specimens from Japan, Honshu, in the original description, without designating 

a holotype. The specimen in his herbarium marked by him as "Typus" is not mentioned 

among these, but this does not invalidate its status as type material since it was one of the specimens 

used in compiling the description. It is from Prov. Etigo, Minami-uonuma-gun, Yuzawa Hot Spring, 

leg. M. TOGASHI, 1954 (no. 54719), and may be designated as lectotype (ASAH, lectotypus, lectotypif. 

nov.). Several of the other (paratype) specimens mentioned in the protologue have also been studied by 

US. 

Icon.: ASAH. 1960, fig. 2B, as St. japonicum subsp. etigoense. - YOSHIM. 1974, P1.32, fig. 315. 

Mat. chim. : atranorin, colensoinic acid, and traces of 4 unidentified substances. 

Distrib.: Japan. Appears to have a well-delimited distribution in the provinces of Etigo, Etizen, 

Shinano and Hida in N. central Honshu. 

Remarks: morphologically hardly distinguishable from St. japonicum TH. FR., although 

some specimens are more robust than is usual in that species and approach St. octomerum 

MULL. ARC. in appearance. Colensoinic acid has been demonstrated by chromatography 

in 6 specimens, including the lectotype. 

68. Stereocaulon exile ASAH. 

J. Jap. Bot. 35: 293 (1960); YOSHIM. Lichen Flora of Japan in Colour: 174 (1974). 

Typus: Japan, Honshu, Prov. Iwashiro, Iizuka, Moniwa, altit. 200 m s. m., leg. T. HIGUTI, 1957 

(no. 6283) (ASAH, holotypus). Paratype specimens from other localities in Honshu and Kiushiu are 

also mentioned in the protologue. 

Facultative synonym (invalid): Stereocaulon tenerrimum LAMB ex ASAH. J. Jap. Bot. 43: 98 (1968) 

(nomen nudum). 

Icon. : ASAH. 1960, fig. 5. - YOSHIM. 1974, fig. 75c; PI. 32, fig. 318. 

Mat. chim. : atranorin, stictic acid, norstictic acid. 

Distrib. : Japan (Honshu and Kiushiu), apparently rather rare. 

Remarks: characterized chiefly by the slender, smooth, glabrous pseudopodetia irre- 

gularly clothed with sorediiform phyllocladial granules, and habitually somewhat resembling 

states of St. nanodes TUCK. (f. carinthiacum) or sorediate St. vesuvianum var. nodulosum 

(WALLR.) LAMB. The minuteness of the phyllocladial granules, which are for the most part 

deliquescent into finely granulose soredia, renders the placing of the species in the subsection 

Botryoideum somewhat doubtful. The cephalodia are however distinctly of the botryose 

type. Usually fertile; hypothecium faintly to moderately brown-pigmented; hymenium 55- 

75 p high; spores (I-) 3 (-4)-septate, 25-33 x 3.0 (-3.5) p.


69. Stereocaulon hokkaidense ASAH. et LAMB 

ex ASAH. J. Jap. Bot. 36: 46 (1961); ASAH. Misc. Bryol. Lichenol. 2 (5): 61 (1961); YOSHIM. Lichen Flora 

of Japan in Colour : 179 (1974). 

Typus: Japan, Hokkaido, Shiretoko Peninsula, on rock (or thin detritus over rock), leg. FUKUSHIMA 

1959 (ASAH no. 5902, holotypus; FH, isotypus). 

Icon. : ASAH. 1961, figs. 1, 2; 1961a, fig. 4. 


Distrib. : Japan (Hokkaido). Known only from the type material. 

Remarks: closely related to St. saviczii DU RIETZ, which has similar habitus, large apothecia 

and broad spores, but which differs in having more or less obvious dark spots or centers 

on the phyllocladia. The phyllocladia of St. hokkaidense do however sometimes show an 

irregular and indistinct mottling with minute darker spots, which may be interpreted as an 

intermediate condition; they are persistently grain-like to verrucose, not becoming dactylaeform 

or coralloid. 

70. Stereocaulon japonicum TH. FR. 

De Stereoc. et Pilophor. Comment.: 18 (1857); Monogr. Stereoc. et Pilophor.: 343 (sep. 39) (1858); 

NYL. Synopsis Methodica Lichenum, 1 (2): 246 (1860); HUE, NOUV. Arch. Mus. Hist. Nat. ser. 3, 10: 

253 (sep. 41) (1898); SAT^ in NAKAI et HONDA, Nova Flora Japonica, Cladoniales (I): 72 (1941); 

ASAH. J. Jap. Bot. 35: 289 (1960); YOSHIM. Lichen Flora of Japan in Colour: 173 (1974). - Stereocaulon 

coralloides var. japonicum (TH. FR.) NYL. MCm. Soc. Sci. Nat. Cherbourg, 5: 96 (1857). 

Typus: Japan, exact locality not indicated, leg. C. P. THUNBERG (date ?) (UPS-THUNB, holotypus). 

Facultative synonyms: Stereocaulon subramulosum var. humile MULL. ARG. Flora, 74: 108 (1891). 

- Stereocaulon exutum var. humile (MULL. ARG.) DODGE, Ann. Cryptog. Exot. 2 (2): 121 (1929). - 

Stereocaulon uvuliferum MULL. ARG. Flora, 74: 109 (1891). - Stereocaulon armatulum ZAHLBR. 

Feddes Repert. 33: 47 (1933). 

Exsicc.: KUROK. Lich. Rarior. et Crit. Exs. Fasc. IV (1973) no. 193 (LD). - YOSHIM. Lich. Japon. 

Exs. ser. 2 (1972) no. 47 (NICH). 

Icon. : TH. FR. 1858, PI. IX, fig. 2. - SAT^ 1941, fig. 25. - ASAH. 1959, PI. 66, fig. 68 ; 1960, fig. 1A ; 

1961b, fig. 1. - YOSHIM. 1974, fig. 76f; PI. 32, fig. 314. 

Mat. chim.: atranorin, stictic acid, and lower concentration of norstictic acid. 

Distrib.: Japan (Honshu, Shikoku, Kiushiu, Ryukyu Islands, the var. aogasimense also in the Izu- 

Shichita Islands), Formosa, China (Chekiang Prov.), Hong Kong, Philippines. 

Remarks: a common and rather polymorphic species. 

Var. subfastigiatum ASAH. 

Bull. Nat. Sci. Mus. (Tokyo), no. 45: 378 (1959) (nomen nudum); J. Jap. Bot. 35: 289 (1960) (cum descript.). 

Typus: Japan, Honshu, Prov. Owari, Jnuyama, leg. Y. ASAHINA, 1938 (no. 1201) (ASAH, holotypus; 

CAN, isotypus). 

Icon.: ASAH. 1959, PI. 61, fig. 21; 1960, fig. 1B. 

Remarks: characterized by the strongly fastigiate-caespitose habitus with harmonic 

phyllocladial branchlets in the lower parts; the upper true phyllocladia are typically developed. 

Rare; seen only from 2 localities in Honshu and Kiushiu. 

Var. tokioense LAMB 

ex ASAH. J. Jap. Bot. 43: 98 (1968) (nomen nudum); ex ASAH. J. Jap. Bot. 44: 265 (1969) (cum descript.).


Typus: Japan, Honshu, Tokyo, on stone wall of the University, leg. Y. ASAHINA. 1943 (no. 650) 

(CAN, holotypus; ASAH, isotypus). 

Icon.: ASAH. 1969, figs. 5, 6. 

Remarks: primary thallus persistent and well developed, with very short, granuliform 

phyllocladia and very dwarf crowded pseudopodetia only 3-5 (-12) rnrn in length. Specimens 

seen from Honshu and Kiushiu. 

Var. aogasimense (ASAH.) LAMB 

ex ASAH. J. Jap. Bot. 44: 266 (1969). - Stereocaulon aogasimense ASAH. J. Jap. Bot. 30: 223 (1955). 

Typus: Japan, Izu-Shichita Islands, Aogashima, on volcanic rock, leg. M. MIZUSHIMA, 1954 

(ASAH no. 541 16, holotypus; FH, isotypus). 

Icon.: ASAH. 1955, p. 224. 

Remarks: distinguished by the minute size of the phyllocladia, which are cylindriccoralloid, 

not over 0.5 rnrn long and less than 0.1 rnrn in thickness. Primary thallus persistent. 

Analogous to the var. subfurfurascens of St. verruculigerum HUE. Known only from the 

Ryukyu and Izu-Shichita Islands. I. YOSHIMURA, who collected material of this variety on 

Okinawa Island in 1973, informed us in litt. that the pseudopodetia are often pendulous in 

nature, which may be a further distinguishing feature. 

71. Stereocaulon nigrum HUE 

Nouv. Arch. Mus. Hist. Nat. ser. 3, 10: 248 (sep. 36) (1898); G. T. JOHNS. Bryologist, 44: 8 (1941); 

ASAH. J. Jap. Bot. 36 (7): 227 (1961); YOSHIM. Lichen Flora of Japan in Colour: 170 (1974). - Stereocaulon 

exutum var. nigrum (HUE) SATB in NAKAI et HONDA, Nova Flora Japonica, Cladoniales (I): 71 

(1941). 

Typus: Japan, Kiushiu, Unzen, leg. R. F. FAURIE, 1895 (no. 15389) (PC-HUE, holotypus). 

Facultative synonym (invalid): Stereocaulon exutoides LAMB ex SATB, Misc. Rep. Res. Inst. Nat. 

Resources (Tokyo), no. 17/18: 172 (1950) (nomen nudum). 

Exsicc.: ASAH. Lich. Japon. Exs. Fasc. I (1954) no. 43, as St. exutum (UPS, us). 

Icon.: ASAH. 1961b, fig. 3. - YOSHIM. 1974, PI. 32, fig. 308. 


Distrib. : Japan (Honshu, Shikoku, Kiushiu). 

Remarks: by some authors confused with St. exutum NYL. It is very similar to that 

species, but differs in having a distinctly brown-pigmented hypothecium. The black color 

of the apothecia emphasized by HUE is an inconstant character; they vary from pale brown 

to dark brown or blackish. 

72. Stereocaulon octomerellum MULL. ARC. 

Nuovo Giorn. Bot. Ital. 24: 190 (1892); G. T. JOHNS. Bryologist, 44: 9 (1941); SAT^ in NAKAI et HONDA, 

Nova Flora Japonica, Cladoniales (I): 66 (1941); ASAH. J. Jap. Bot. 45: 68 (1970); YOSHIM. Lichen Flora 

of Japan in Colour : 172 (1 974). 

Typus: Japan, Honshu, Nikko, "saxicola ut videtur", leg. YATABE, 1890 (no. 173) (G-MOLL, holotypus). 

Facultative synonyms: Stereocaulon octomerelloides ASAH. J. Jap. Bot. 45: 68 (1970). - Stereocaulon 

sandwicense MAGN. Ark. Bot. 31A (6): 39 (1944). - Stereocaulon octomerellum var. sandwicense 

(MAGN.) LAMB, Canad. J. Bot. 29: 582 (1951).


Exsicc. ASAH. Lich. Japon. Exs. Fasc. VI (1959) no. 296 (UPS, us). - YOSHIM. Lich. Japon. Exs. 

ser. 2 (1972) no. 48 (NICH). 

Icon. : SAT6 1941, fig. 23. - ASAH. 1970, fig. 3. - YOSHIM. 1974, fig. 76e; PI. 32, fig. 3 13. - MAGN. 

et ZAHLBR. 1945, PI. VII, fig. 1, as Sf. sandwicense. 


Distrib. : Japan (Honshu, Hokkaido, Shikoku, Kiushiu); Hawaii. ASAHINA, 1970, states that many 

specimens of Sf. octornerellurn have been collected in Honshu and Hokkaido, but only a few in Shikoku 

and Kiushiu. 

Remarks: this tiny species is saxicolous (very rarely terricolous), firmly attached to rock, 

and has a distinct persistent primary thallus consisting of minute, scattered or crowded verruculae 

which are not of coralloid form except when giving rise to pseudopodetial initials. 

The phyllocladia on the pseudopodetia vary from papillose-verruculose to cylindric-coralloid. 

It seems to be always fertile. St. octomerelloides ASAH. was distinguished on the basis of 

verruculose phyllocladia on the pseudopodetia, "more delicate appearance and less separated 

spores", but these characteristics show all intergradations to the typical condition. St. 

sandwicense, formerly considered by us as a distinct variety (LAMB, 1951, LAMB ex FREY 1967), 

also intergrades in all characters formerly considered to separate it, and can no longer be 

regarded as distinct. St. octomerellum is therefore a somewhat variable species in its morphology 

and anatomy. The algae of the cephalodia are usually Stigonema, but in the lectotype 

specimen of St. sandwicense from Hawaii (s) they are Calothrix parietina (NAEG.) THUR. 

(identified by Dr. F. DROUET), a Cyanophyceous genus not previously reported in Stereocaulon 

(cfr. AHMADJIAN, 1967, p. 149/150). 

73. Stereocaulon octomerum MULL. ARC. 

Flora, 74: 109 (1891); DODGE, Ann. Cryptog. Exot. 2 (2): 115 (1929) (as syn. of St. curtaturn NYL.); 

ASAH. J. Jap. Bot. 36 (7): 230 (1961); YOSHIM. Lichen Flora of Japan in Colour : 170 (1974). 

Typus: Japan, Honshu, summit of Mt. Ontake, leg. M. MlYos~r, 1890 (no. 113) (G-MULL, holotypus). 

Icon.: ASAH. 1961b, fig. 6; 1970, fig. 1A. - YOSHIM. 1974, fig. 76b; P1. 32, fig. 309. 


Distrib. : Japan (Honshu, Shikoku, Hokkaido), Formosa, U.S.A. (Alaska). 

Remarks: has been confused with St. curtatum NYL., but differs from the latter not only 

chemically but also in its taller growth, soon evanescent primary thallus, narrower spores etc. 

HUE'S description of "Stereocaulon octomerum", 1898-1901 (1898), p. 249 (sep. 37) obviously 

refers to St. curtatum. Although the spores of St. octomerum are normally 7-septate, there 

is a good deal of variation; many 3-, 4- or 5-septate spores also occur, and occasionally a 

specimen is found with only 34-septate spores. The single Alaskan specimen seen was from 

the Kenai Peninsula, leg. H. KROG, 1957 (0); see KROG, 1968, p. 88. 

74. Stereocaulon pendulum ASAH. 

ex SAT^, J. Jap. Bot. 17: 247 (1941); YOSHIM. Lichen Flora of Japan in Colour: 174 (1974). 

Typus: Japan, Honshu, Prov. Kozuke, Mt. Akagi, "ad saxa non calcarea", leg. K. TSUNODA, 

1923 (TI, holotypus, not seen by us). Paratypes from Prov. Musashi and Prov. Shinano, sent by Dr. 

ASAHINA, were however examined. 

48 MAGNUSSON in MAGN. et ZAHLBR. (1944) stated that the exsiccat ZAHLBR. Lich. Rarior. Exs. no. 

58, as St. condensaturn, should be referable to St. sandwicense MAGN. But according to our examination 

of the examples in PC and UPS, it is St. condensaturn HOFFM. as originally named.


Exsicc.: ASAH. Lich. Japon. Exs. Fasc. VI (1959) no. 297 (UPS, us). 

Icon.: SAT^ 1941a, fig. 11. -YOSHIM. 1974, PI. 32, fig. 320. 


Distrib. : Japan (Honshu). 

Remarks: apparently a rare species, characterized by the unique pendulous habitus. 

The phyllocladia are for the most part granular or papillate, very small, but in the lowermost 

parts of the pseudopodetia are larger and distinctly cylindric-coralloid. The apothecia (not 

present in the holotype specimen) are terminal, sometimes on short side-branchlets, and up 

to 1.5 rnm dim.; hypothecium pigmented (reddish- or yellowish-brown); hymenium 60-80 p 

highppores cylindric-fusiform, (I-) 3 (-5)-septate, 25-35 (-40) x4.04.5 p. 

75. Stereocaulon prostratum ZAHLBR. 

Bot. Mag. (Tokyo), 41: 340 (1927); SAT^ in NAKAI et HONDA, Nova Flora Japonica, Cladoniales (I): 

76 (1941); Sci. Rep. Tohoku Imp. Univ., ser. 4, Biol. 29 (314): 222 (1963); YOSHIM. Lichen Flora of 

Japan in Colour : 174 (1 974). 

Typus: Japan, Honshu, Prov. Mutsu, Mt. Hakkoda, leg. Y. ASAHINA, 1924 (no. 14748) (w, holotypus). 

Icon. : ZAHLBR. 1927, PI. XI, fig. 7. - YOSHIM. 1974, PI. 32, fig. 319. 


Distrib. : Japan (Honshu, Hokkaido). 

Remarks: rare; seen only from about 5 localities. It grows on soil among mosses or 

directly on rock, forming almost crustose, appressed patches. Primary thallus more or less 

persistent, consisting of coralloid phyllocladia. Pseudopodetia more or less ligneous, glabrous, 

copiously branched. Phyllocladia densely crowded, often in branched clusters, up to 

0.8 mm long.80 Hypothecium pigmented, yellow-brown to brown (not "decolor" as stated 

by ZAHLBRUCKNER). Hymenium 70-90 p high. Spores 3-septate, 20-28 x 3.5-4.5 p. There 

is some resemblance, as ZAHLBRUCKNER states, to St. curtatum NYL., but the latter differs in 

its chemistry, 7-septate spores, different habitus, and coarser and less crowded phyllocladia. 

76. Stereocaulon saviczii Du RIETZ 

Ark. Bot. 22A (1 3) : 13 (1 929) ; ASAH. J. Jap. Bot. 45 : 66 (1 970) ; YOSHIM. Lichen Flora of Japan in Colour 

179 (1 974). 

Typus: U.S.S.R., Kamtchatka, Awatchinskaja Sopka volcano, altit. 700 m s. m., on rock, leg. E. 

HULTEN, 1920 (no. 686a) (ups, holotypus; FH, isotypus). 

Exsicc.: KUROK. Lich. Rarior. et Crit. Exs. Fasc. IV (1973) no. 194 (LD). 

Icon.: Du RIETZ 1929, PI. I, fig. 2. - ASAH. 1970, figs. lB, 2. - YOSHIM. 1974, fig. 77h; PI. 33, 

fig. 335. 


Distrib. : U.S.S.R. (Kamtchatka), U.S.A. (Alaska), Japan (Hokkaido and Honshu). 

Remarks: closely related to St. hokkaidense ASAH. et LAMB. This pair of species seem 

to be the most primitive representatives of subsect. Botryoideum (see discussion on p. 194). 

The phyllocladia of St. saviczii are subglobose-verrucose to cylindric-coralloid, with darker, 

spot-like, depressed centers (wrongly interpreted as perforated by Du RIETZ) or irregularly 

mottled with darker spots or patches. The spores are (I-) 3-4 (-5)-septate, (1%) 17-37 x 

(3) (-6.5) p. Several collections seen from Kamtchatka, and two from Alaska: Nunivak 

4B ZAHLBRUCKNER 

loc. cit. gives the number as 149. 

80 By a typographical error given as 8 mm in the original description.

I. MACKENZIE LAMB: A Conspectus of Srereocaulon 

Island (cfr. KROG, 1968, p. 89) and Iliamna Bay, leg. R. THOMAS (WIS). 

77. Stereocaulon verruculigerum HUE 

Bull. Soc. Bot. France, 54: 417 (1907); DODGE, Ann. Cryptog. Exot. 2 (2): 148 (1929); ASAH. J. Jap. Bot. 

35: 294 (1960); YOSHIM. Lichen Flora of Japan in Colour: 174 (1974). 

Typus: Java, exact locality not stated, "supra terram", leg. Bors, 1903 (PC-HUE, holotypus, not seen 

by us). 

Faculrarive synonyms: Stereocaulon verruculigerum var. formosanum ASAH. J. Jap. Bot. 35: 295 

(1960). - Stereocaulon formosanum ASAH. loc. cir. p. 294 (nom. inval.). 

Icon.: ASAH. 1960, fig. 6. - YOSHIM. 1974, PI. 32, fig. 317. 


Distrib. : Java, Sumatra, Celebes, Formosa, Japan (Honshu). 

Remarks: morphologically indistinguishable from St. japonicum TH. FR. but with a different 

chemistry and a more southerly Asian distribution pattern. It is very common in 

Java, growing on rocks, stones and soil, sometimes on roof-tiles. 

Var. subfurfurascens LAMB (n. var.) 

Diagn.: A specie typica differt phyllocladiis minutissimis, valde congestis, furfuraceocoralloideis 

(fere isidioideis), pseudopodetio applanatis et intricatis (haud divaricatis) et 

crustam granulatam formantibus. 

Typus: Sumatra, near Aer Batumbak along road between Lubuk Selasih and Simpang, altit. ca. 

1300 m s. m., on rock, leg. P. GROENHART, 1953 (no. 9274) (L, holotypus; FH, isotypus). 

Remarks: known from 3 collections made in or in the vicinity of the type locality in 

Sumatra. Pseudopodetia more or less erect, 2.0-3.5 an long. Analogous to the var. aogasimense 

(ASAH.) LAMB of St. japonicum TH. FR. 

Sect. 4. Lobophoron (DUVIGN.) LAMB 

78. Stereocaulon foliolosum NYL. 

Synopsis Methodica Lichenum, 1 (2): 240 (1860); HUE, Bull. Soc. Bot. France, 54: 416 (1907); DODGE, 

Ann. Crypteg. Exot. 2 (2): 122 (1929); LAMB, Ergebn. Forsch.-Unternehmen Nepal Himalaya, 1 (4): 

351 (1966); J. Jap. Bot. 43: 296 (1968). 

Typus: 2 collections cited from India, Himalayas: one unlocalized, leg. JACQUEMONT (date ?), 

the other from Kumaon (Kumaun, Uttar Pradesh), altit. 12000 feet, leg. STRACHEY and W~NTERBOTTOM 

(date ?); no holotype designated. As lectotype DODGE (1929, p. 122) designated the specimen collected 

by JACQUEMONT (no. 743) (PC, lectotypus; H-NYL, no. 40092, isolectotypus). The second (paratype) 

specimen also seen by us (H-~n, no. 39771). 

Icon. : LAMB 1968, figs. ID, 2C. 


Distrib.: Himalayan region. India (Uttar Pradesh, Punjab, N. W. Bengal), Nepal, Sikkim. The 

var. botryophorum also in central Africa (Rwanda), and the var. srrictum also in China (Yiiman). 

Remarks: in the Himalayas it grows at altitudes between 1800 and 4000 m s.m., on rocks 

and stones, more rarely on soil. 

Var. botryophorum (MULL. ARG.) LAMB (n. comb.) 

Basionym: Stereocaulon botryophorum MULL. ARG. Flora, 74: 371 (1891). 

Typus: India, Uttar Pradesh, above Ihala, altit. 12-13000 feet, leg. DUTHIE (prior to 1886) (G-M~~LL,


holotypus; K, isotypus). 

Icon. : LAMB 1951, fig. 2D (ceph. cortex); 1968, figs. IC, 2C; as St. botryophorum. 

Remarks: it now seems clear that this taxon should be placed with St. foliolosum, as 

suspected by LAMB, 1966, p. 351. The phyllocladia remain in an indeterminate, subsquamulose 

to subcoralloid condition, not becoming distinctly foliose. If sterile, a certain distinction 

from St. himalayense AWAS. et LAMB does not generally seem possible, but in the case of the 

DUTHIE type, although it has no apothecia, the tall and robust growth refers it to St. jolio~osum 

rather than to St. himalayense. 

Var. strictum (BAB.) LAMB 

Canad. J. Bot. 29: 582 (1951); J. Jap. Bot. 43: 297 (1968). - Stereocaulon ramulosum var. strictum 

BAB. Hooker's J. Bot. Kew Gard. Misc. 4: 250 (1852). - Stereocaulon rnixturn [subsp.]** strictum 

(BAB.) NYL. Synopsis Methodica Lichenum, 1 (2): 239 (1860). - Stereocaulon strictum (BAB.) MULL. 

ARG. Bull. Herb. Boissier, 3 (4): 194 (1895) (nom. illegit.; non St. strictum TH. FR. ex anno 1857). - 

Stereocaulon macrocephalum var. strictum (BAB.) DODGE, Ann. Cryptog. Exot. 2 (2): 126 (1929). 

Typus: 2 collections from N. India, Uttar Pradesh, Kurnaun Div., cited in the protologue: Madhari, 

altit. 8700 feet, and Sagtia deo Pass, altit. 11500 feet, both leg. STRACHEY and WINTERBOTTOM 

(date ?), without designation of holotype. DODGE (1929, p. 126) lectotypified the taxon on the former 

specimen. This specimen (EM-BAB, lectotypus, is a mixture of the present variety and St. piluliferum 

TH. FR., and the isolectotype in K consists of the same mixture. The other (paratype) specimen from 

Sagtia deo Pass is also mainly the present variety, but with an intermixture of St. pomiferurn DuVIGN. 

Icon.: NYL. 1860, PI. VII, fig. 13, as St. mixtum subsp. strictrim (spores). 

Remarks: in this variety the foliose phyllocladia show a transition to coralloid form in 

the upper parts of the pseudopodetia. AGHORAMURTHY, SARMA and SESHADRI (1961) reported 

a Himalayan specimen ascribed to St. foliolosum var. "Stictum" [sic] to contain atranorin, 

stictic acid, an undetermined substance (probably an aliphatic hydrocarbon) and 

D-arabitol; if stictic acid were indeed present, the specimen must have been incorrectly determined. 

79. Stereocaulon coniophyllum LAMB 

Bot. Not. 114 (3): 266 (1961); J. Jap. Bot. 43: 294 (1968); Oz. et CLAUZ. Les Lichens: 470 (1970); 

YOSHIM. Lichen Flora of Japan in Colour: 175 (1974). 

Typus: Norway, Hordaland, Eidfjord, Hardanger, Veringfoss, reg. subalpina, altit. 450-630 m s. 

m., leg. J. HAVAAS, 1899 (FH, holotypus; BG, isotypus). 

Icon.: LAMB 1961, figs. 1, 2; 1968, figs. lA, 2A. - HENSSEN et JAHNS 1974, fig. 3.21~ (cephalodium 

in section). - YOSHIM. 1974, fig. 75e. 


Distrib. : Europe, N. America, Himalayan region, Africa, Japan. Sweden, Norway, Finland, Austria, 

Canada (N. W. Territ.), U. S. A. (Alaska), N. India, Nepal, Sikkim, Africa (Uganda), Japan. 

Remarks: the very wide distributional range suggests that this may be a very ancient 

species of Afro-Asiatic origin, perhaps persisting at the present day only in relict stations of 

a formerly continuous area in the northern hemisphere. Its ecology in Norway was studied 

by VEVLE (1975); it is there restricted to the spray zones of larger waterfalls, the effect of the 

spray seeming to be twofold, i.e. keeping the plant continuously moist and stabilizing the 

temperature climate. VEVLE describes a new association, the Stereocauletum coniophyllae, 

which includes also Vestergrenopsis isidiata (DEGEL.) DAHL and Placopsis gelida (L.) LINDS.


80. Stereocaulon himalayense AWAS. et LAMB (n. sp.) 

Diagn.: Pseudopodetia caespitosa, erecta, substrato arcte affixa, ad 2.5(-4.0) cm longa 

(sed saepe depauperata et breviora), subsimplicia vel parum aut sat congeste ramosa, nonnihil 

rigida, glabra et denudata, albida aut subochraceo-pallida, phyllocladiis granuliformibus vel 

brevissime dactyloideis inaequaliter vestita, basi vulgo e crusta phyllocladiorum thalli primarii 

subpersistenti surgentia. Soredia nulla. Cephalodia protosacculata, intus solida, subglobosa 

aut pulvinata, glebosa et tandem e globulis aggregatis composita. Apothecia terminalia, 

primum urceolata et sat crasse marginata, dein plana convexave, margine reflex0 et evanescenti, 

disco fusconigro. Hypothecium incoloratum. Hymenium pro ratione humile (7675 p, raro 

ad 90 p altum), ascis vulgo 5665 p longis. Sporae 5-8nae, rectiusculae, pauciseptatae, septis 

(I-) 3-4, rarissime 5-7, cylindrico-fusiformes, longit. (26) 25-40 (-49, crassit. 3.5-4.5 (-5.0) p. 

Atranorinum et acidum lobaricum continens (phyll. Ktflavesc., PD+pallide flavesc.). - 

Habitu St. foliolosum var. botryophorum exacte referens, a quo statu sterili vix distinguendum, 

differentiis enim e fructificatione, viz. altitudine hymenii et septatione sporarum, desumptis. 

Typus: Nepal, Khumbhakarna Himal, Dhankuta Distr., Upper Barun Valley, altit. 4800 m s. m., 

leg. WRABER, 1972 (AWAS, holotypus; FH, GZU, isotypes). 


Distrib.: Himalayan region: N. India (Uttar Pradesh) and Nepal, at very high altitudes (3500- 

5400 m s. m.). 

Remarks: morphologically and habitually similar to St. foliolosum var. botryophorum 

(M~~LL. ARG.) LAMB, but differing in the lower hymenium and the shorter, pauciseptate spores. 

This obvious reduction in the dimensions of the reproductive structures, unlike the degree 

of vegetative development, is unlikely to be produced by environmental conditions, but is 

most probably genetically determined, and we noted in the present case that the asci are 

frequently well developed, many with mature spores of quite healthy appearance. Plants 

growing at the highest altitudes may be very stunted, of crustose habitus. Grows on stones 

and boulders of glacial moraines, sometimes associated with St. coniophyllum LAMB and 

Hypogymnia sp. Known from 9 different collections. 

8 1. Stereocaulon humbertii DUVIGN. 

Lejeunia, Mkm. no. 14: 132 (1956); LAMB, J. Jap. Bot. 43: 295 (1968). 

Typus: Africa, Zaire, Kivu, Mt. Mikeno, altit. 4250 m s. m., leg. H. HUMBERT, 1929 (BRLU, holotypus). 

Icon.: DWIGN. 1956, fig. 41. -LAMB 1968, figs. lB, 2B. 

Mat. chim. : atranorin and lobaric acid, or quite often in a deficient phase with atranorin 

only demonstrable. 

Distrib.: central Africa (Uganda, Zaire, Kenya), at altitudes of 35U550 m s. m. 

Remarks: habitually resembles St. botryosum ACH. em. FREY, apart from the distinctive 

cephalodia. Occasionally fertile; apothecia very large, 4-5 (-8) rnrn diam., terminal, flat or 

variously undulate (not becoming strongly convex), with subpersistent dark proper margin; 

finally becoming fissured and dividing into secondary discs. Hypothecium colorless. Hymenium 

7@90 ,u high. Spores (3-) 4-5 (-6)-septate, 45-60 x (3%) 4.0-4.5 p. 

Subgen. 11. Pilophoropsis (LAMB) LAMB 

82. Stereocaulon curtum (RA~.) LAMB 

Canad. J. Bot. 29: 565, 581 (1951). - Stereocaulon macrocarpurn ("macrocarpon") var. curta RAs.


Revista Univ. (Santiago), 22: 200 (1937). 

Typus: Chile, Valdivia, Volciin Villarrica, altit. 1300m s. m., leg. A. HOLLERMAYER, 1933 (H- 

RAS, holotypus). 

Icon.: LAMB 1951, fig. 4. 

Mat. chim.: atranorin, lobaric acid, and traces of unidentified substances. 

Distrib. : apparently endemic to a rather restricted region of the South American Cordillera between 

latitudes 39" and 43" S. (provinces of Valdivia and Llanquihue in Chile, Rio Negro and Chubut in 

Argentina). 

Remarks: when typically developed, the stiff, thick, simple or sparingly branched pseudopodetia, 

with verrucose thallus-mantle, have much the appearance of a Pilophorus. Alpine 

specimens tend to be more compact and stunted, of a darker cinereous color, more copiously 

and closely branched, and widely decorticated in the basal and middle parts, with a greater 

development of coralloid phyllocladial branchlets. The pigmented tissue of the upper central 

cone (unique to this species) extends downwards into the lower colorless part in ill-defined 

strands; its color is not changed by KHO. In the type specimen the spores are (7-) 8-9 (-10)septate, 

48-68 x 3.54.0 p. In one specimen from Argentina some spores are simple (eseptate), 

others 1-5-septate; apparently an abnormality. 

Subgen. III. Holostelidium LAMB 

Sect. 1. Holostelidium 

Subsect. A. Holostelidium 

83. Stereocaulon sorediifrum HUE 

Nouv. Arch. Mus. Hist. Nat. ser. 3, 10: 250 (sep. 38) (1898); DODGE, Ann. Cryptog. Exot. 2 (2): 143 

(1929); SATO in NAKAI et HONDA, Nova Flora Japonica, Cladoniales (1): 85 (1941); LAMB, J. Jap. Bot. 

40: 274 (1965); YOSHIM. Lichen Flora of Japan in Colour: 175 (1974). 

Typus: Japan, Honshu, near Yokosuka ("Yokoska"), on rock, leg. SAVATIER (date ?) (PC-HUE, 

holotypus). 

Facultative synonyms: Stereocaulon exutum var. sorediata RAs. J. Jap. Bot. 16: 88 (1940). - 

Stereocaulon sorediiferum f. esorediosum ASAH. Bull. Natl. Sci. Mus. (Tokyo), no. 45: 381 (1959) (nomen 

nudum). 

Exsicc.: ASAH. Lich. Japon. Exs. Fasc. I (1954) no. 44 (UPS, us). - Krypt. Exs. Vindob. Cent. XLIII 

(1961) no. 4238 (FH, s). - YOSHIM. Lich. Japon. Exs. ser. 2 (1972) no. 49 (NICH). 

Icon. : YAS. 1925, PI. XIII, fig. la, as St. ramulosum. - S A 1941, ~ fig. 30. - ASAH. 1959, P1. 66, 

fig. 67, as St. sorediiferum f. esorediosum. - YOSHIM. 1974, figs. 74c, 75d; PI. 32, fig. 321. 

Mat. chim.: atranorin and lobaric acid. The mention of a chemically aberrant strain 

from Vietnam in LAMB, 1965, p. 274 was based on an error; the material in question was 

subsequently found to contain atranorin and lobaric acid, in spite of its spurious reddish PD 

react ion. 

Distrib.: Japan (Honshu, Shikoku, Kiushiu), Formosa, China (Szechuan), Hong Kong, Philippines, 

Vietnam. 

Remarks: cephalodia sacculate. Occasionally almost or even totally lacking in the 

production of soredia (f. esorediosum ASAH.); such plants appear morphologically like St. 

massartianum HUE, but may be distinguished by their different chemi~try.~~ 

61 But see discussion under St. massartianum, p. 273.


Var. leprosolingulatum LAMB (n. var.) 

Diagn. : Soredia non solum ad ramulos phyllocladoideos prodientia, sed etiam in apicibus 

pseudopodetiorum, ubi strias angustas linguiformes unifaciales formant. 

Typus: Taiwan (Formosa), Mt. Tsu-Tson-San, Mt. Ali, altit. 2200-2600 m s. m., on rock, leg. S. 

KUROKAWA, 1964 (no. 475) (FH, holotypus; TNS, isotypus). 

Remarks: in this variety soredia occur not only on the ends of the short divaricate phyllocladial 

branchlets, but also on the apices of the pseudopodetia, where they form narrow, 

tongue-like, often curved, apically acuminate bands on one side only (the opposite side being 

smoothly corticate). They extend downwards to a distance of up to 7 mm from the tip, and 

are whitish and finely powdery. 

84. Stereocaulon brassii LAMB (n. sp.) 

Diagn. : Pseudopodetia erecta, caespitosa, substrato arcte adhaerentia, valida et robusta, 

5-8 (-10) cm longa, inferne ca. 2 mm, in partibus superioribus 0.7-1.3 rnrn crassa, irregulariter 

et sparse vel sat copiose ramosa, rigida et quasi lignosa, glabra, pro majore parte decorticata. 

Phyllocladia cylindrico-coralloidea aut subulata. Soredia desunt. Cephalodia subglobosa, 

sacculata, scrobiculato-convoluta. Apothecia terminalia aut praesertim ad apices ramulorum 

lateralium prolata, e clavulis parvis subglobosis oriunda, tandem convexa et immarginata, 

1-2 mm lata. Hypothecium incolor aut leviter isabellino-nubilatum. Hymenium altum 

(15CL200 p). Sporae vermiformi-sinuosae, 10CL210 p longae, 4-5 p latae, multiseptatae, 

septis (7-) 9-16 (-20). Conidia filiformia, leviter arcuata. Mat. chim.: atranorinum, acidum 

sticticum et acidum norsticticum (K+flavescens, PD fvulgo aurantiaco-rubescens). - Habitu 

St. massartiano HUE persimile, sed multo magis robustum et rigidum. 

Typus: New Guinea, north-east region, Eastern Highlands Distr., Mt. Wilhelm, altit. 3560 m s. 

m., "terrestrial in alpine grassland", leg. L. J. BRASS, 1959 (no. 30042) (us, holotypus; FH, isotypus). 

Mat. chim.: atranorin, stictic acid, norstictic acid, one or more unidentified substances 

related to stictic acid, and occasionally ursolic acid (a triterpene). 

Distrib. : N. E. New Guinea and W. New Guinea (Irian), at high altitudes. 

Remarks: resembles a very robust condition of St. massartianum HUE. AS in that species, 

the apothecia often occur at the ends of short lateral side-branchlets. The phyllocladia 

towards the apices of the pseudopodetia are small, short, subulate, but not reduced to papillae 

as in St. montagneanum LAMB. The cephalodial algae are usually of a pale Nostocoid type; 

Stigonema and Scytonema were found only in specimens from W. New Guinea (Irian). One 

specimen from Irian collected by BRASS and MEYER DREES (FH) has very abundant apothecial 

clavulae, which contain numerous immersed coiled ascogonia provided with trichogynes 

protruding out above the surface, giving a whitish-pruinose macroscopic appearance. 

85. Stereocaulon claviceps TH. FR. 


DODGE, Ann. Cryptog. Exot. 2 (2): 110 (1929); LAMB, J. Jap. Bot. 43: 297 (1968). - Stereocaulon 

ramulosum [subsp.]* claviceps (TH. FR.) NYL. Synopsis Methodica Lichenum, 1 (2): 236 (1860). 

Typus: Mexico, Cumbre de Sempoaltepec, altit. 11000 feet. leg. LIEBMANN (date ?) (no. 71a) 

(UPS-TH FR, holotypus; C, FH, isotypes). 

Icon.: TH. FR. 1857a, fig. c (spore); 1858, PI. VIII, fig. 2. - NYL. 1860, PI. VII, fig. 12 (spores). 

- LAMB 1968, figs. 1 E, 2 E, F. 

Mat. chim.: atranorin, stictic acid, norstictic acid.

272 Joum. Hattori Bot. Lab. No. 43 1977 

Distrib. : Mexico, Costa Rica, central and South Africa. 

Remarks: cephalodia protosacculate. Has been commonly confused with St. pomiferum 

DUVIGN., from which it differs in being sorediate. Not a common species, and usually of 

local and scanty occurrence; seen from several localities in Mexico (Puebla, Mexico and 

Veracruz States) and from one each in Costa Rica (LAMB and METZGER, 1969, FH), Kenya 

(Sw~~scow, 1973, SWINSC) and South Africa, Natal (ESTERHUYSEN, 1953, BOL). St. claviceps 

var. yunnanense HUE belongs to St. pomiferum DUVIGN. (see p. 277). 

86. Stereocaulon macrocephalum MULL. ARG. 

Flora, 74: 371 (1891); DODGE, Ann. Cryptog. Exot. 2 (2): 125 (1929); LAMB, J. Jap. Bot. 43: 298 (1968). 

Typus: India, Uttar Pradesh, Kumaun Div., Garhwal Distr., above Bhowani, altit. 12-14000 feet, 

on rocks, leg. DUTHIE (date ?) (no. 5227) (K, holotypus; G-MULL, isotypus). 

Icon.: DUVIGN. 1956, fig. 10 B. - LAMB 1968, figs. 1 F, 2 D. 

Mat. chim. : atranorin and stictic acid. 

Distrib. : Himalayan region: N. India and Nepal. 

Remarks: cephalodia protosacculate. A rare species, seen to date only from two collections. 

It is possible that it may be conspecific with St. pomiferum DUVIGN., representing 

a vegetatively reduced state of the latter, although the habitus, the apparent absence of distinct 

clavulae in the immature apothecia, and the structure of the cephalodial cortex appear to 

afford distinctive characters. If it should prove to be the same as St. porniferum, the species 

epithet macrocephalum would have priority. ZAHLBRUCKNER, 192G27 (1927), p. 660, erroneously 

places St. macrocephalum as a synonym of St. piluliferum TH. FR. 

87. Stereocaulon massartianum HUE 

Nouv. Arch. Mus. Hist. Nat. ser. 3, 10: 252 (sep. 40) (1898); LAMB, J. Jap. Bot. 40 (9): 271 (1965). 

Typus: Java, near Lebak-Saet, altit. 2100 m s. m., leg. D. MASSART, 1895 (no. 1689) (BRLU, holotypus). 

Facultative synonyms: "Stereocaulon nesaeum" Auctt. (non Stereocaulon nesaeum NYL. Ann. Sci. 

Nat., Bot. ser. 4, 11: 236 (1859), vide infia.) - Stereocaulon rnixtum [subsp.]*** nesaeum (NYL.) NYL. 

Synopsis Methodica Lichenum, 1 (2): 240 (1860) (saltem pr. p.; non St. nesaeum NYL. ex anno 1859). 

- Stereocaulon nesaeum var. Iecideoides VAIN. Philipp. J. Sci. 4 (5) : 662 (1909). - Stereocaulon nesaeum 

var. zeorina VAIN. ibid. p. 661. - Stereocaulon chlorocarpoides ZAHLBR. Feddes Repert. 33: 48 (1933). 

- Stereocaulon massarfianum var. chlorocarpoides (ZAHLBR.) LAMB J. Jap. Bot. 40 (9): 271 (1965). - 

Stereocaulon stalagmitiferum DUVIGN. Biol. Jaarb. "Dodonaea", 9: 93 (1942). 

Exsicc.: Krypt. Exs. Vindob. Cent. XXI (1913) no. 2067, as St. mixtum (c, FH, L, LD, PR, US, WELC) 

(Strain 11). - Lich. Exs. Univ. Colorado Mus. Fasc. VIII (1971) no. 302 (co~o, FH). - VBZDA, Lich. 

Sel. Exs. Fasc. XXVIII (1968) no. 684, as St. massartianum var. chlorocarpoides (FH). 

Icon.: NYL. 1860, PI. VII, fig. 15 a, b (spores and ceph. algae), as Sf. mixtum subsp. nesaeum. - 

KREMP. 1870, PI. XIX, fig, la, b, c, d, as St. nesaeum. SAT^ 1941, fig. 29, as St. chlorocarpoides. - 

DUVIGN. 1942, fig. 5, as St. stalagmitiferum; 1956, fig. 10c; fig. 31 (apothecia and clavules), as St. 

nesaeum and St. stalagmitijerum. - LAMB 1965, fig. 1, partly as St. massartianum var. chlorocarpoides. 

Mat. chim. : there are two morphologically indistinguishable chemical strains, which may 

later possibly prove to be entitled to the rank of separate species: 

Strain I, with atranorin, stictic acid, often a small amount of norstictic acid, and possibly 

some trace of constictic acid (K+pale yellow, PD+orange-red). The typical strain. 

Strain 11, with atranorin, norstictic acid and connorstictic acid (no stictic acid present) 

(K+yellow or occasionally red, PD+persistent yellow).

I. MACKENZIE Lam: A Conspectus of Stereocaulon 273 

Crushed thallus or apothecial fragments of Strain 11, when tested with KHO under the 

microscope, regularly produce the characteristic red spicular crystals of the potassium salt 

of norstictic acid; those of Strain I similarly treated may or may not do so, depending on the 

concentration of norstictic acid present. The biosynthetic difference between the two strains 

would appear at first sight to be rather insignificant, stictic acid being probably formed from 

norstictic acid by 0-methylation in a terminal reaction, but studies with other lichen genera 

seem to indicate that the difference is often correlated with a phenotypic morphological 

difference, a different distributional pattern, and different ecological preferences, these features 

combining to reinforce the chemical evidence to such an extent that distinct species seem to 

be indicated; see W. L. CULBERSON'S remarks on the Ramalina siliquosa group (1969, p. 163). 

A segregate from St. massartianum characterized by quite different chemical constitution 

(atranorin, lobaric acid and porphyrilic acid) and different geographical distribution (Himalayan 

region) is St. togashii LAMB (p. 278). 

The macroscopic reactions on the phyllocladia in St. massartianum are not always reliable 

for distinguishing the two chemical strains, and chromatography is necessary in some doubtful 

cases. Specimens of Strain I1 with a low concentration of norstictic acid may give the reactions 

Kt-yellow, PD +yellow or -, and therefore not be distinguishable by this means from 

St. togashii. Usually in such cases, however, crushed-out carpogonial clavulae, which are 

richer in lichen substances than the phyllocladia, will be found to show a few red crystals of 

the norstictic acid salt on treatment with KHO under the microscope. 

A specimen collected by WISSEL, 1936, in West New Guinea (Irian), Mt. Carstensz, no. 

1897 (FH, L), giving a negative reaction with PD, was found by chromatography and UV 

spectrum analysis (C. CULBERSON, S. HUNECK) to contain atranorin and an unidentified 

substance which appears to be an unknown depsidone. Unfortunately not enough material 

was available for a further analysis by other methods. 

Japanese material recorded as "Stereocaulon massartianum var. chlorocarpoides, lobaric 

acid strain" by YOSHIMURA, 1974, p. 175, probably refers to an atypically esorediate condition 

of St. sorediiferum HUE. Some similar specimens, esorediate and morphologically indistinguishable 

from St. massartianum, but containing lobaric instead of stictic acid, have been 

seen in various herbaria also from Borneo, Celebes, New Guinea and Java. They may 

represent an undescribed species differing chemically from St. massartianum, rather than 

esorediate states of St. sorediiferum, as their occurrence falls outside the main geographical 

area of the latter. How they should be dealt with taxonomically is a problem which we are 

unable to resolve at present. 

Distrib.: S. E. Asia: Java, Lombok, Sumatra, Celebes, Malaya, Borneo, Philippines, New Guinea, 

Formosa; occurring at higher elevations. The distribution of the two chemical strains seems to coincide, 

except that Strain I1 has not yet been seen from Formosa, Malaya, Lombok or Sumatra. Records 

of "Stereocaulon nesaeurn" from outside this area refer to other species. A specimen in M from 

Brazil, leg. GLAZIOU, thus determined by KREMPELHUBER, is St. rarnulosurn (SW.) RAUSCH. MULLER ARG. 

(1894) recorded "St. nesaeurn" from New Zealand; the specimen not seen by us, but doubtless referable 

also to St. ramulosum. 

Remarks: cephalodia sacculate. In the older literature and in herbaria St. massartianum 

has figured under the name Stereocaulon nesaeum NYL., an epithet which is however of somewhat 

uncertain application and certainly cannot correctly refer to the present species. DODGE 

(1929) typified St. nesaeum NYL. on a Javan specimen collected by ZOLLINGER, in PC. This 

lectotypification is inadmissable, since in NYLANDER'S original description (Ann. Sci. Nat., 

Bot. ser. 4, 11: 236. 1859) the only locality mentioned is "in insulis Sandwich" (no collector's


name given). In his Synopsis (1860) he mentioned specimens from Java, the Philippines and 

Polynesia, as St. mixtum subsp. nesaeum, using the epithet in the sense of St. massartianurn, 

a circumscription which was followed by later authors. Nevertheless, the name nesaeum 

must be typified on the material listed in the original description: Sandwich Islands, = Hawaii. 

St. massartianum or related species do not seem to occur in the Hawaiian Islands, and we 

have not been able to find any material from there designated by NYLANDER as St. nesaeum 

either in his herbarium (H-NYL) or in PC. MAGNUSSON in MAGNUSSON and ZAHLBRUCKNER, 

1944, p. 47, expressed the opinion that the type material of St. nesaeum from the Hawaiian 

Islands must belong to St. ramulosum (Sw.) RAUSCH., but the question cannot be satisfactorily 

resolved unless NYLANDER'S original specimen comes to light. 

The two varieties lecideoides and zeorina distinguished by VAINIO are taxonomically 

insignificant. According to our lectotypification (p. 318) both appear to belong to Strain 11. 

Stereocaulon chlorocarpoides ZAHLBR. represents the condition of St. massartianum in which 

the apothecia are borne at the ends of apical branches, not on short lateral branchlets. Although 

this state was formerly considered by us to be a distinct variety (LAMB, 1965), it has 

since become obvious that it is a normal non-taxonomic variation. Similar states occur in 

the closely related species St. brassii and St. togashii. Stereocaulon stalagmitiferum DUVIGN. 

was separated from St. massartianum on the basis of having compound or aggregated carpogonial 

clavulae. This feature seems to be without taxonomic value. 

88. Stereocaulon montagneanum LAMB 

J. Jap. Bot. 40 (9): 272 (1965). 

Typus: Malaya, Pahang State, Cameron Highlands, altit. 1500-2500 m s. m., leg. M. TOGASHI, 

1962 (no. 62236) (FH, holotypus; ASAH, isotypus. Isotype material distributed in KUROK. Lich. Rarior. 

et Crit. Exs. no. 142). 

Facultative synonym (doubtful) : Stereocaulon coralligerum MEYER, Nebenstunden meiner Beschaefftigungen 

im Gebiete der Manzenkunde, Erster Theil: 156 (1825); see Remarks below. 

Exsicc.: KUROK. Lich. Rarior. et Crit. Exs. Fasc. I11 (1971) no. 142 (isotype material) (FH). 

Icon.: LAMB 1965, fig. 2. - G. T. JOHNS. 1954, fig. 1, as St. nesaeum. 

Mat. chim. : atranorin and stictic acid; possibly also norstictic acid. 

Distrib. : Indonesia (Java, Sumatra, Celebes), Malaya, Borneo, New Guinea, at higher elevations. 

Remarks: cephalodia sacculate. Closely related to St. massartianum HUE, but differing 

in its taller growth, conspicuously dendroid habitus, and in the reduction of the phyllocladia 

on the uppermost branches to short papillae. It has been given various unpublished herbarium 

names by several authors, the first being MONTAGNE. A doubtful published epithet 

which may possibly refer to this species is Stereocaulon coralligerum MEYER (1825), very 

incompletely described as from Chile, leg. CHAMISSO, and Brazil, leg. BEYRICH. MEYER 

describes the apothecia as being in swollen heads resembling the fruit bodies of Sphaerophorus, 

a character congruent with the present species. Furthermore, there is a specimen in UPS-TH 

FR, labelled by TH. FRIES "Stereocaulon coralligerum Mey. Specimen origin. Loc. nat. non 

indic. Ex hb. Berol. accepi. Berol. Dec. 1861. Th. Fr.", and it is a very typical fertile specimen 

of St. montagneanum. DUVIGNEAUD, 1942, p. 91, attributes the name St. coralligerum to 

various specimens seen by him from Chile under the name of St. ramulosum, but does not 

give a clear description of them. St. montagneanum is not known from S. America, where 

its occurrence is very unlikely, and it is almost certain that the specimen in UPS-TH FR mentioned 

above must have come from S. E. Asia. If this is so, it cannot be regarded as type 

material, which would have to be from either Chile or Brazil, the countries cited in the pub-


lished protologue. There is a specimen of St. montagneanum in L from Java, without name 

of collector, stated to be from the herbaria of BUSE and SPRENGEL, named "Stereocaulon 

ramulosum" and "Stereocaulon coralligerum Meyer". The available evidence points to the 

probability of St. coralligerum MEYER having been based on an Indonesian specimen of St. 

montagneanum, but because the protologue cites only specimens from Chile and Brazil, the 

typification of the epithet in the sense of the present species is not nomenclaturally possible. 

89. Stereocaulon paradoxum LAMB (n. sp.) 

Diagn.: Pseudopodetia (inter muscos et hepaticas crescentia) plus minusve decumbentia 

aut prostrata, substrato laxe vel vix affixa et partibus infimis ut videtur emorientia; longit. 

24 cm, crassit. 0.64.8 (-1.0) mm, teretia, subsimplicia vel parce irregulariter ramosa, varie 

flexuosa intricataque, pr. maj. p. bene tomentosa (tomento coacto-spongioso, pallido, ut in 

St. myriocarpo). Soredia desunt. Phyllocladia nurnerosa, aggregata, digitata aut cylindricocoralloidea, 

bene ramosa, parva exiliaque, longit. 0.5-1.5 mm, crassit. 0.10-0.25 mm, albida, 

pseudopodetia aequaliter vestientia et in eorum apicibus minute papillata sed mox digitatoelongantia. 

Cephalodia pseudopodetiis lateraliter insidentia, intus solida (protosacculata), 

laeviuscule subglobosa, pallide rufofuscescentia, parva (0.54.8 rnm lata), nec pedicellata nec 

scrobiculata, sed majora interdum fissurantia et tandem verrucoso-divisa; structura corticalis 

gelatinoso-pseudoparencymatica. (Apothecia et pycnidia adhuc ignota.) Atranorinum et 

acidum lobaricum continet (K+flavescens, PD+pallide flavescens). -Prim0 intuitu St. 

myriocarpum TH. FR. simulans, sed cephalodiis protosacculatis mox distinguendum. 

Typus: India, Uttar Pradesh, Charnoli Distr., between Waan and Bhuna, altit. 3500 m s. m., on 

soil over rock among mosses and foliose hepatics, leg. A. SINGH, 1967 (no. 91599) (FH, holotypus; 

LWG, ~SO~YPUS). 

Mat. chim.: atranorin, lobaric acid, and trace of an unidentified substance. 

Distrib.: Himalayan region. Known only from the type specimen. 

Remarks: This rather strange new species with protosacculate cephalodia superficially 

resembles St, myriocarpum TH. FR. in habitus, form of phyllocladia and tomentosity of 

pseudopodetia. As apothecia are not known, its placing in Subsect. Holostelidium can only 

be tentative. The cephalodia contain Nostocoid algae; their cortex is 20-45 p thick, pale 

sordid yellowish but not nubilated, of pseudoparenchymatous structure with isodiametric, 

thick-walled cells; lumina rounded, 34 p diam., the walls gelatinously confluent and 2-3 p 

thick. There is no indication of any palisadic structure. 

90. Stereocaulon piluliferum TH. FR. 


DODGE, Ann. Cryptog. Exot. 2 (2): 134 (1929). 

Typus: Nepal, exact locality not given, leg. WALLICH (date ?) (UPS-TH FR, holotypus; c, FH-TAYL, 

isotypes). 

Fac~ltative synonyms: Stereocaulon sinense HUE, Nouv. Arch. Mus. Hist. Nat. ser. 3, 10: 251 (sep. 

39) (1898). - Stereocaulon piluliferum var. sinense (HUE) LAMB ex ASAH. in KIHARA, Fauna & Flora of 

Nepal Himalaya, 1952-53, 1: 50 (1955) (comb. inval.). 

Exsicc.: KUROK. Lich. Rarior. et Crit. Exs. Fax. I11 (1971) no. 144 (FH, LD). - Lich. EXS. Univ. 

Colorado Mus. Fasc. IV (1966) no. 150, as St. '>iliferumM (LD). - V~ZDA, Lich. Sel. Exs. Fax. XLI 

(1971) no. 1016 (FH). 

Icon.: TH. FR. 1858, PI. VIII, fig. 3. -HUE 1898-1901 (1899), PI. 111, fig. 4, as St. sinense. 

Mat. chim. : atranorin, stictic acid, norstictic acid; in one specimen from India, Darjeeling


Distr., leg. AWASTHI, 1954, no. 3119 (AWAS, FH) porphyrilic acid is also present, apparently 

as an accessory constituent. 

Distrib.: Nepal, N. India (Uttar Pradesh, N. West Bengal), Sikkim, Assam, China (Yunnan, 

Szechuan), possibly E. Tibet (localization uncertain); Java ? (see Remarks below). Erroneous record: 

Japan M SAT^, 1941, 1965; acc. to I. YOSHIMURA in litt. based on a misidentification by DODGE of St. 

pomiferum DUVIGN.). 

Remarks: cephalodia sacculate. Closely related to St. pomiferum DUVIGN., but clearly 

distinguished by the cephalodial structure. Usually fairly small, ca. 3 cm tall, but occasionally 

up to 6 cm. Hymenium up to 19q-240) p high. Spores (80-) 150-170 p long, 10-19 (-29)septate. 

Pycnoconidia filiform, arcuate, 8-12xca. 0.4 y. HUE, 1898-1901 (1898), p. 250 

(sep. 38) gives a detailed description, as St. piluliferum, of a specimen from India, Himachal 

Pradesh, Simla, but its identity is not certain from the data given. In M-KREMP there is a 

packet labelled in NYLANDER'S writing "Stereocaulon nesaeum" from Java, Pangerango, leg. 

KURZ, 1861 (no. 706). It consists of a mixture of St. massartianum HUE and St. piluliferum; 

the latter perhaps an accidental herbarium intermixture, as suggested by duplicates of the 

same collection seen in FH, H and MEL, which do not contain it. 

9 1. Stereocaulon pomifeum DUVIGN. 

Lejeunia, Mem. no. 14: 1 19 (1956); LAMB, J. Jap. Bot. 43: 298 (1968); YOSHIM. Lichen Flora of Japan 

in Colour: 175 (1974). - Stereocaulon claviceps var. pomiferum (DUVIGN.) LAMB, Ergebn. Forsch.- 

Unternehmen Nepal Himalaya, 1 (4): 352 (1966). 

Typus: Africa, Uganda-Zaire border, Ruwenzori, Mt. Stanley, altit. 4200 m s. m., leg. HAUMAN, 

1932 (no. 931B) (BRLU, holotypus). 

Facultative synonyms: Stereocaulon mamillosum DUVIGN. Lejeunia, Mdm. no. 14: 111 (1956). - 

Stereocaulon claviceps var. yunnanense HUE, NOUV. Arch. Mus. Hist. Nat. ser. 3, 10: 251 (1898).62 - 

Stereocaulon macrocephalum var. yunnanense (HUE) DODGE, Ann. Cryptog. Exot. 2 (2): 125 (1929). - 

Stereocaulon yunnanense (HUE) LAMB ex ASAH. in KIHARA, Fauna &Flora of Nepal Himalaya, 1952- 

53, 1: 50 (1955) (comb. inval.). - Stereocaulon claviceps var. tuberatum LAMB ex FREY, Bot. Jahrb. 

Syst. 86: 219 (1967) (nomen nudum). 

Exsicc. : ASAH. Lich. Japon. Exs. Fasc. VI (1959) no. 292, as St. claviceps (ups, us). 

Icon.: DUVIGN. 1956, figs. 32, 38, 39; figs. 1, 33, 34, 35, 36, 37, as St. mamillosum. -LAMB 1968, 

figs. lG, 2E, F. - YOSHIM. 1974, PI. 32, fig. 322. 

Mat. chim. : there appear to be 2 chemical strains of this species: 

Strain I, with atranorin, stictic acid, usually also some norstictic acid, occasionally 

porphyrilic acid as accessory substance. The typical, common and widely distributed strain. 

Deficient phase I, with atranorin and porphyrilic acid. One specimen seen, from Bhutan. 

Deficient phase 11, with atranorin and an unidentified pigment. One specimen seen, 

from Nepal. 

Strain 11, with atranorin and norstictic acid (no stictic acid). Seen only from the Hawaiian 

Islands, and perhaps to be regarded as a distinct species. Also present in Hawaii are plants 

with atranorin only demonstrable, apparently a deficient phase of this strain. 

Distrib.: central Africa (Uganda, Zaire, Rwanda-Burundi, Kenya), India (Uttar Pradesh, N. 

West Bengal), Nepal, Bhutan, Sikkim, Tibet, China (Yunnan), Mexico, Venezuela, Peru, Japan, Formosa 

and (the Strain 11) U.S.A. (Hawaii). Some other plant groups show broadly the same Asiatic- 

African distribution pattern: lichens mentioned by MAGNUSSON (1940), mosses by SHARP and IWATSUKI 

(1965), algae by MAEKAWA (1966), phanerogams by MATUDA (1953) and HEDBERG (1961). 

82 Lapsu "yltnnana" in ZAHLBRUCKNER, 1926-27 (1927), p. 635.


Remarks: cephalodia protosacculate. Till recently confused with St. claviceps TH. FR. 

(a sorediate species) to which it is closely related. The habitus is occasionally corymbose- 

dendroid in African specimens, with the cephalodia mainly basal on the pseudopodetia. 

The apothecial receptacles vary from smooth to strongly tesselate-tuberculate. Hymenium 

126200 (-270) p high. Spores (1W) 140-180 (-220) x3.54.0 (4.5) p, (16) 15-22 (-32)- 

septate. St. claviceps var. yunnanense HUE represents a well developed state of St. pomiferum 

with very verrucose-tuberculate receptacles, high hymenium (up to 270 p) and spores up to 

32-septate, reaching 220 p in length. 

Fo. mikenoense (DUVIGN.) LAMB (n. comb.) 

Basionym: Stereocaulon mikemense DUVIGN. Lejeunia, Mkm. no. 14 : 126 (1 956). 

Typus: Africa, Zaire, Kivu, Mikeno Volcano, altit. 4250 m s. rn., leg. H. HUMBERT, 1929 (BRLU, 

holotypus). 

Icon.: DUVIGN. 1956, fig. 40, as St. mikenoense. 

Remarks: distinguished by the robust pseudopodetia, corymbosely branched above and 

densely clothed with strigose phyllocladia. Receptacle tesselate-verruculose. 4 specimens 

seen, from Zaire, Uganda and Rwanda-Burundi. 

92. Stereocaulon procerum GROENH. ex LAMB (n. sp.) 

Diagn.: Pseudopodetia caespitosa, substrato ut videtur sat arcte affixa, elongata, 7-13 cm 

longa, simplicia aut parce ramosa, fragilia, decorticata, superficie subtomentosa. Phyllocladia 

coralloideo-cylindrica aut subulata, in parte superiori pseudopodetiorum brevia simpliciaque, 

inferne longiora (ad 7 mm) et ramosa, superficie subtomentoso-mollescenti. Soredia 

nulla. Cephalodia sacculata, subglobosa, fusca, valde scrobiculata, intrinsecus laxe hyphosa, 

strato corticali hyalino e hyphis fistulosis, gelatinosis, paliformibus contexto. Apothecia in 

apicibus ramulorum lateralium sita, ca. 1 mm lata, e clavulis pyriformibus plus minusve persistentibus 

oriunda, mox convexa et indistincte marginata. Hypothecium leviter pigmentiferum 

(pallide fuscescens). Hymenium 166170 p altum, paraphysibus filiformibus tenuibusque. 

Sporae vermiformes, flexuosae, in asco partim spiraliter contortae, 16-19- septatae, 

116140 p longae et 4.04.5 p latae. Mat. chim.: atranorinum et acidum sticticum (Kf 

flavescens, PDf aurantiaco-rubescens). - St. massartiano HUE simile et arcte afie, sed statura 

proceriori, fragilitate, superficie subtomentoso-mollescenti, et phyllocladiis longioribus distinctum. 

Typus: Sumatra, Atjeh Gajolanden, G. Kemiri, altit. 3200-3300 m s. m., on rocks in mountainheath, 

leg. VAN STEENIS, 1937 (no. 10259) (L, holotypus; FH, isotypus). 

Mat. chim.: atranorin and stictic acid. 

Distrib.: Indonesia (Sumatra). Known from 2 collections made by VAN STEENIS at elevations 

between 3200 and 3400 m s. m. 

Remarks: closely related and similar to St. massartianum HUE, but distinguished by its 

very tall growth, obvious fragility, longer and coarser phyllocladial branchlets, and especially 

by the spongy, subtomentose, smoothly felted surface of pseudopodetiurn and phyllocladia, 

resembling in this respect the varieties exalbidum and submollescens of St. ramulosum (Sw.) 

RKusc~. St. massartianum sometimes has a slightly spongy or subtomentose pseudopodetial 

surface, but its phyllocladia are always more or less smoothly corticate. The pycnidia of 

St. procerum are immersed in small swellings around the base of developing apothecial hypophyses, 

and appear as blackish spots; conidia filiform, arcuate, 1612 x ca. 0.5 p. Dr. P.


GROENHART, who died in 1965, distinguished St. procerum as a new species and left isotype 

material and drawings with the present author. 

93. Stereocaulon togashii LAMB (n. sp.) 

Diagn.: Statura et habitu omnino ut in St. massartiano HUE: pseudopodetia stipatocaespitosa 

aut forsan partim decumbentia, ad 6.5 cm alta, simplicia aut parum ramosa, 

glabra, pr. max. p. denudata, sorediis destituta, phyllocladiis divaricatis, coralloideis, ramosis, 

cephalodiis sacculatis, scrobiculato-corrugatis, intus laxe hyphosis, apotheciis vulgo singulatim 

in apicibus ramulorum lateralium prolatis, parvis, convexo-immarginatis; hypothecium 

incoloratum; hymenium (loo-) 130-150 p altum; sporae vermiformes, 7-12-septatae, longit. 

75-120 p, crassit. 3.54.5 p. Thallus atranorinum, acidum lobaricum et acidum porphyrilicum 

geret (phyllocladiis PD leviter flavescentibus). - Materiis chimicis necnon distributione 

geographica (himalayensi) a St. massartiano diversum. 

Typus: Sikkim, Puleg-chu, altit. 2100 m s. m., leg. M. TOGASHI, 1960 (no. 6072) (FH, holotypus; 

ASAH, isotypus). 

Mat. chim.: atranorin, lobaric acid, porphyrilic acid. 

Distrib.: Himalayan region; N. India, Nepal, Sikkim, Bhutan). 

Remarks: morphological and microscopic characters are in full agreement with St. 

massartianum HUE, but the presence of lobaric and porphyrilic acids instead of stictic acid, 

combined with the distinctly separated pattern of distribution, surely entitles this taxon to 

the rank of a separate species. Cephalodial algae Scytonema in all specimens examined; 

cephalodial cortex 2540 p thick, gelatinized, colorless to sordid yellowish, clear, palisadic, 

with vertically parallel, oblong lurnina 1.0-1.5 p wide, also intermixed a number of larger, 

oval to isodiametric lumina 34 p in diameter. Pycnidia forming minute, dark brown, semiglobose 

swellings 0.10-0.15 mrn diarn. at apices of pseudopodetia; conidia filiform, curved, 

9-lox ca. 0.5 LA. 

Subsect. B. Aciculisporae Du RIETZ ex LAMB 

94. Stereocaulon rarnulosum (Sw.) RAUSCH. 

Nomenclator Botanicus, edit. 3: 328 (1797); TH. FR. De Stereoc. et Pilophor. Comment.: 11 (1857); 

Monogr. Stereoc. et Pilophor.: 329 (sep. 24) (1858); NYL. Synopsis Methodica Lichenum, 1 (2): 235 

(1860); TUCK. A Synopsis of the North American Lichens, Part I: 230 (1882); HUE, NOUV. Arch. MUS. 

Hist. Nat. ser. 3, 10: 243 (sep. 31) (1898)s3; DODGE, AM. Cryptog. Exot. 2 (2): 137 (1929); MAGN. in 

MAGN. et ZAHLBR. Ark. Bot. 31A (6): 44 (1944); MARTIN et CHILD, Lichens of New Zealand: 155 

(1972). -Lichen ramulosus Sw. Nova Genera et Species Plantarum: 147 (1788). - Stereocaulon 

ramulosum [f.] a. genuinum TH. FR. De Stereoc. et Pilophor. Comment.: 11 (1857). 

Typus: Jamaica, leg. SWARTZ (date ?) (s, holotypus; H-ACH, isotypus). 

Facultative synonyms: Stereocaulon furcaturn FR. Systema Orbis Vegetabilis, Pars I: 285 (1825) 

(see Remarks below). - Stereocaulon ramulosum var. furcaturn (FR.) NYL. Mem. Soc. Sci. Nat. Cherbourg, 

5: 96 (1857). - Stereocaulon mixtumN~~. Ann. Sci. Nat., Bot. ser. 4, 11: 210 (1859). - Stereocaulon 

proximum NYL. Ann. Sci. Nat., Bot. ser. 4,ll: 210 (1859). - Stereocaulon rarnulosum f. proximum 

(NYL.) TUCK. Proc. Amer. Acad. Arts, 7: 228 (1866). - Stereocaulon rarnuloslrrn var. compacturn 

MULL. ARC. Bull. Herb. Boissier, 4: 87 (1896). - Stereocaulon rarnulosum [f.] y. strigosum TH. FR. De 

Stereoc. et Pilophor. Comment.: 12 (1857). - Stereocaulon rarnulosum var. strigosum (TH. FR.) HUE, 

Nouv. Arch. Mus. Hist. Nat. ser. 3, 10: 244 (sep. 32) (1898). - Stereocaulon ramulosum var. microcar- 

n3 Description based on a Patagonian specimen which is probably referable to f. elegans TH. FR.


poides MULL. ARG. Flora, 72: 505 (1889). - Stereocaulon vimineum TH. FR. De Stereoc. et Pilophor. 

Comment. : 13 (1857). - Stereocaulon ramulosum var. vimineum (TH. FR.) NYL. Ann. Sci. Nat., Bot. 

ser. 4, 11: 209 (1859). - Stereocaulon ramulosum f. vimineum (TH. FR.) HUE, NOUV. Arch. Mus. Hist. 

Nat. ser. 3, 2: 245 (sep. 37) (1890). - Stereocaulon ramulosum var. sublecanoreum LAMB ex VARGAS, 

Revista Univ. (Cuzco), 37 (94): 187 (1948) (nomen nudum). 

Ex~icc.~': FOLLM. Lich. Exs. Sel. Mus. Bot. Berol. Fasc. I (1968) no. 14 (FH. In DUKE it is a mixture 

of St. ramulosum and St. vesuvianum PERS.) - Lich. Exs. Sect. Bot. Mus. Hist.-Nut. Hungar. (1969) no. 

87 (LD). - Lich. Exs. Univ. Colorado Mus. Fasc. VII (1969) no. 249, as St. ramulosum var. nudatum 

(DUKE, LD). - Lichenotheca Parva, edit. Sect. Bot. Mus. Hist.-Nut. Hungar. (1969) no. 83 (LD). - 

LINDIG, Lich. Novo-Granatensesbb no. 2501, as St. mixtum (FH-TUCK); no. 2537, as St. proximum (FH- 

TUCK). - LQIKA, Lichenotheca Univ. Fasc. IV (1886) no. 153 (WELC). - MERR. Lich. EXS. Fasc. 111 

(1910) no. 55, as St. ramulosum f. proximum (c, DUKE, FH, LD. In us it is var. pilophoroides (TUCK.) 

LAMB); Fax. V (1910) no. 110. as St. mixtum (LD, us); Ser. 11, Fasc. V (1927 ?) no. 109 (co~o, us). - 

TILDEN, South Pacific Plants (date ?) no. 160 (co~o, us). - V~ZDA, Lich. Sel. Exs. Fax. XXII (1967) 

no. 544 (FH, DUKE); Fasc. XXXVII (1970) no. 910 (FH); Fasc. LIX (1977) no. 1464 (BRNU). - ZAHLBR. 

Lich. Rarior. Exs. no. 138 (1910), as St. mixtum (FH). 

PAYER 1850, fig. 415. -NYL. 1860, P1. VII, fig. 10 (spores, pycnidium, conidia); fig. 28, 

as St. furcatum (conidia, cephalodial algae). - LINDS. 1861, figs. 27-32 (pycnidia and conidia). - 

BORNET 1873, PI. XI, figs. 1-3 (cephalodial structure); P1. XIII, figs. 8-9, as St. furcatum (cephalodial 

structure). - REINKE 1895, fig. 48, subfigs. 111, IV. - PLITT 1935, fig. 3 (field photograph). - GRASS] 

1950, fig. 6; P1. VIII, lower fig. - MARTIN and CHILD 1972, P1. 46. - HENSSEN and JAHNS 1974, figs. 

3.21b (cephalodial structure), 13.14d (fertile pseudopodetium). -LAMB 1951, figs. 1, 2a; PI. I, fig. 1, 

PI. 11, fig. 6; 1976, fig. 1 (cephalodium). 

Mat. chim.: atranorin, perlatolic acid, often anziaic acid, occasionally possibly glomelliferic 

and stenosporic acids as accessory compounds, 2 unidentified substances. HUNECK and 

FOLLMANN (1967) reported atranorin only in a Chilean specimen; probably a deficient phase. 

CAMFSIE (1968) reported atranorin, anziaic acid and a hitherto unknown dimethyl ether of 

anziaic acid in New Zealand material. A mixture of unidentified tetrahydroxy fatty acids, 

in addition to atranorin and perlatolic acid, was reported in specimens from Jamaica, New 

Zealand, Argentina and Costa Rica by Fox and HUNECK (1970) and Fox, FOLLMANN and 

HUNECK (1971). DUVIGNEAUD (1942, 1956) had stated the chemical constituents of St. 

ramulosum and several other Stereocaulon species to be atranorin, divaricatic acid, and a 

chemically uncharacterized substance to which he gave the name "ramulosine" (not to be 

confused with the compound ramulosin produced by Pestalotia (Fungi Irnperfecti) and described 

by HESSELTINE, BENJAMIN, BRADLE and HENDERSHOT, 1963). The alleged presence of 

divaricatic acid in St. ramuIosum has not been confirmed by subsequent investigations. Dr. 

TH. ESSLINGER (Duke Univ.) investigated the chemical constitution of the pseudopodetia and 

the apothecia separately by thin-layer chromatography in material of St. ramulosum from 

Jamaica and New Zealand. Perlatolic acid, with traces of 2 unidentified substances, was 

found in both apothecia and pseudopodetia, but atranorin was present only in the latter 

(ESSLINGER, personal commun.). This investigation was suggested by the fact that in another 

64 DICKSON, Hortus Siccus Britannicus Fasc. XI (1796) no. 24, named Lichen paschalis, in BM is a 

mixture of St. ramulosum and an indeterminable species, probably St. alpinum LAUR. NO locality is 

given on the label. 

66 This is probably not a true exsiccat, as no fully printed labels were issued, most of the data being 

handwritten. 

The illustration of "Stereocaulon ramulosum" in YASUDA, 1925, PI. XIII, fig. 1 is of St. sorediiferum 

HUE.


species of this group, St. trachyphloeum LAMB, the apothecia and the pseudopodetia proved 

to contain different substances (see p. 304). 

Distrib. : very widely distributed and common in the southern hemisphere, and penetrating into the 

northern hemisphere in Central America and the West Indies as far north as Mexico; also in Hawaii. 

Southern hemisphere: Australia, Tasmania, New Zealand (including the subantarctic islands), New 

Hebrides (var. macrocarpum), New Caledonia (var. macrocarpum), Tahiti (var. macrocarpum), Fiji (?, 

somewhat doubtful), central Africa (Uganda, Zaire, Rwanda-Burundi, Kenya, Tanzania), Gough 

Island in the S. Atlantic, Argentina, Chile, Falkland Islands, South Georgia (var. pulvinare), Bolivia, 

Peru, Colombia, Ecuador, Brazil. Northern hemisphere: Venezuela, Panama, Costa Rica, Mexico, 

Jamaica, Cuba (var. perpumilum), Dominican Republic, Guadeloupe, Haiti, Curacao, Puerto Rico, 

Hawaiian Islands. Unverified records:" Himalayan region (CHOPRA, 1934; BISWAS, 1947); China 

(ZAHLBRUCKNER, 1934); Java (ZAHLBRUCKNER, 1956); Comores Islands (NYLANDER, 1860, as St. proximum); 

Amsterdam Island (S. Indian Ocean) (NYLANDER, 1886, as St. proximum); Tristan da Cunha 

(HUE, 1898-1901 ; LYNGE, 1937, as St. rnixtum); Canary Islands (KLEMENT, 1965), probably referable to 

St. atlanticum (LAMB) LAMB. Erroneous records: Japan (YASUDA, 1925,=St. sorediiferum HUE); 

Philippines (ZAHLBRUCKNER in Krypt. Exs. Vindob. no. 2067, as St. mixtum,=St. massartianum HUE); 

South Africa (DOIDGE, 1950, as St. proximum; apparently a misinterpretation of NYLANDER'S record 

from the Comores Islands mentioned above); the specimen quoted as from the Cape of Good Hope 

by TH. FRIES, 1858, p. 25, collected by FRASER, was almost certainly from Au~tralia~~. 

St. ramulosum probably represents an old Antarcto-Tertiary element, as its distribution agrees in 

the main with that of Astelia (Liliaceae), of which SKOTTSBERC (1960) writes: "Astelia is one of the best 

examples of an old Antarcto-Tertiary genus, now spread around the colder southern hemisphere, with 

scattered insular stations and extending north to the Hawaiian Islands in the Pacific and to the Mascarene 

Islands in the Indian Ocean." St. salazinum (BORY) FBE, a species very closely related to St. 

ramulosum, does occur in the Mascarene Islands (Reunion). A similar distribution pattern is shown by 

the phanerogam genera Acaena, Gunnera, Nertera and Brachycoma, included by VAN STEENIS (1953) in 

the "General Subantarctic" floral element. 

Remarks: shows considerable variation in stature and habitus. Occasionally, when 

growing among grasses, the pseudopodetia are often more slender and tend to be emorient at 

the base. The hypothecium is more or less brownish and nubilated up to the base of the 

hymenium, without any intervening hyaline stratum as in St. implexum TH. FR. This brownish 

coloration usually disappears on treatment with KHO, except in W. Indian specimens and var. 

pilophoroides (TUCK.) LAMB, in which a certain degree of pigmentation is present and some 

brown color persists in the hypothecium when thus treated. Hymenium (6%) 7%80 (-1 10) p 

high. Spores 3-5 (-7)-septate, (28-) 3240 (-90) x3.tk3.5 (4.0) p. Pycnoconidia filiform, 

arcuate, 9-1 1 x ca. 0.5 p. The ontogenetic development of St. ramulosum has been described 

and figured by JAHNS (1970). 

St. ramulosum f. genuinum TH. FR. designates the typical species; Lichen ramulosus Sw. 

is cited as synonym, and the type of the latter is also that of the nomenclaturally superfluous 

forma. Var. compactum MULL. ARG. is merely a stunted and depauperate condition. Var. 

microcarpoides MULL. ARG. is a fortuitously microcarpic state with apothecia 1 mm or less in 

diameter. F. strigosum TH. FR. is a taxonomically insignificant condition of lax and elongated 

growth habitus. St. furcatum FR. was based on a small and delicate growth state of St. 

5r TUCKERMAN, 1872, 1882-88, mentions specimens said to have been collected in North America, 

but considers such records doubtful. RIDDLE (1910) also mentions a reputed occurrence in Oregon, 

but is of the opinion that a confusion of labels was involved. We have never seen St. ramulosum from 

the continental United States, although the related St. rnicrocarpum MULL. ARC. occurs there. 

The closely related species St. meyeri B. STEIN does however occur in South Africa.


ramulosum; it was confused with St. virgatum ACH. by TH FRIES (1857), this error being pointed 

out by M~~LLER ARG. (1887). St. vimineum TH. FR. is a state of St. ramulosum with markedly 

harmonic basal phyllocladial branchlets, a not uncommon condition in this species. 

NYLANDER (1860 and elsewhere) segregated several species from St. ramulosum mainly 

or entirely on the basis of different Cyanophyceous algae in the cephalodia. He did not 

recognize them as such, but commented on their resemblance to blue-green algal genera, 

using e.g. the term "sirosiphoid" for Stigonema filaments. He considered St. ramulosum (s. 

str.) to be characterized by Nostoc-like "granula gonimia" in the cephalodia, which was an 

incorrect assumption if reference is made to the actual type specimen, which has Scytonema. 

NYLANDER'S segregates, which all belong to St. ramulosum, were St. mixtum NYL. (Scytonema 

in cephalodia), St. proximum NYL. ('Lsir~~iphoid'', i.e. Stigonema in cephalodia) and St. 

ramulosum subsp. macrocarpoides NYL. (like var. macrocarpum (RICH.) BAB., but with "sirosiphoid 

gonimia", i.e. Stigonema in cephalodia instead of Nostocoid). FORSELL (1883) however 

showed that in St. ramulosum different genera of Cyanophyceae may occur on the same 

individual and even in the same cephalodium, and the distinction of different species on this 

basis has been accordingly denied by HELLBOM (1896) and DEGELIUS (1941). Nevertheless, it 

has been observed that the typical species usually has Stigonema or Scytonema in the cephalodia 

(rarely Nostocoid algae) whereas the f. elegans has most frequently Nostocoid algae (LAMB, 

1951, p. 542-543). 

Fo. elegans TH. FR. 

De Stereoc. et Pilophor. Comment. : 11 (1857);n0 Monogr. Stereoc. et Pilophor. : 328 (sep. 24) (1858); 

HUE, NOUV. Arch. Mus. Hist. Nat. ser. 3, 10: 244 (sep. 32) (1898).80 

Typus: several collections cited from Campbell Island, Magellan Straits, Chile (Valdivia), Cape 

of Good Hope and Australia, no holotype designated. DODGE (1929, p. 137) lectotypified it on a specimen 

from Magellan Straits, leg. N. J. ANDERSON (date ?) (UPS-TH FR, lectotypus). 

Facultative synonym: Stereocaulon proximum f. traversii HUE, Nouv. Arch. Mus. Hist. Nat. ser. 

3, 10: 245 (sep. 33) (1898). 

Exsicc. : ARN. Lich. Exs. (Lich. Jurae) no. 1209 (date ?), as St. proximum (FH). - FOLLM. Lich. 

Exs. Sel. Mus. Hist. Nat. Casselensi Fasc. VII (1975) no. 136 (FH, KASSEL). 

Icon. : HOOK. F. 184547, PI. LXXX, fig. 1, as St. ramulosum. 

Remarks: a luxuriant form of the cooler southern hemisphere, reaching a height of 12 

(-18) cm and with elegantly branched phyllocladia. Common in southern Argentina and 

Chile, Australia, New Zealand, Tasmania and the subantarctic islands. Sometimes apparently 

transitional to the var. macrocarpum (RICH.) BAB. Cephalodia usually with Nostocoid algae, 

rarely Stigonema or Scytonema. 

Fo. nudatum (MULL. ARG.) LAMB (n. comb.) 

Basionym: Stereocaulon proximum var. nudatum MULL. ARG. Flora, 69: 252 (1886); DODGE, 

Ann. Cryptog. Exot. 2 (2): 136 (1929). - Stereocaulon ramulosum var. nudatum (MULL. ARG.) MULL. 

ARG. J. Linn. Soc. Bot. 32: 199 (1896); LAMB ex FREY, Bot. Jahrb. Syst. 86: 247 (1967). 

Typus: Australia; 2 collections cited, one without exact locality ("Australia Felice"). DODGE 

(1929) selected as lectotype the specimen from Brogers Creek, leg. BAUERLEN, 1884 (no. 10) (G-MULL, 

lectotypus). 

There as St. ramulosum ,5'. elegans; referred to as a forma in the accompanying text. 

Bo AS St. ramulosum var. elegans, not an intentional new combination. This taxon is accorded 

varietal rank also by several other authors, under the impression that TH. FRIES described it as such.


Remarks: characterized by the strongly denuded and ecorticate, ligneous pseudopodetia 

with smooth, almost polished surface. Appears to be transitional to the var. rnacrocarpum 

(RICH.) BAB. and the f. elegans TH. FR. Seen from Australia, Tasmania, New Zealand and 

Chile. 

Fo. subcompressum LAMB (n. f.) 

Diagn. : Ramuli phyllocladoidei leviter complanati, flabelliformiter divisi, latere inferiori 

pallidiores. Ceterum ut in typo. 

Typus: Tasmania, Mt. Wellington, on rocks, leg. R. A. BASTOW (date ?) (LD, holotypus; FH, 

isotypus). 

Remarks: phyllocladia slightly flattened, flabellately branched in one plane and with 

paler undersides. Otherwise as in the typical species. Rather uncommon; seen from Australia, 

Tasmania, subantarctic islands of New Zealand, and Chile (incl. Juan Fernandez). 

Some of the references in the literature to St. ramulosum var. compressum BAB., such as HUE 

(1898-1901) and LINDAU (1912), perhaps refer to this form of St. ramulosum and not to the 

true "var. compressum" BAB. (=St. fronduliferum LAMB). 

Fo. tomentosulum LAMB (n. f.) 

Diagn. : Pseudopodetiis tomentosulis (tomento tenuissimo, adpresso) dignoscitur; habitu 

interdum macrocarpoideo (phyllocladiis pseudopodetiorum fertilium superne rarescentibus 

vel fere nullis). 

Typus: Colombia, Dept. Santander, W. slope of PBramo Rico, altit. 3600 m s. m., on soil, leg. E. P. 

KILLIP and A. C. SMITH, 1927 (no. 17733) (s, holotypus; FH, isotypus). 

Remarks: pseudopodetia more or less continuously clothed with a very thin, adpressedfelted 

tomentum. Stature and habitus either as in the typical species or more resembling 

that of var. macrocarpum (RICH.) BAB. with upper parts of fertile pseudopodetia more or less 

destitute of phyllocladia. Rather common in Colombia and Peru; also seen from Ecuador, 

Bolivia, Venezuela, Brazil, West Indies (Dominican Republic) and New Zealand. 

Var. macrocarpum (RICH.) BAB. 

ex HOOK. F. Botany of the Antarctic Voyage of H. M. Discovery Ships Erebus and Terror in the Years 

1839-1843. . .Part 11. Flora Novae-Zelandiae. Part ii. Flowerless Plants: 294 (1855); NYL. 

Synopsis Methodica Lichenum, 1 (2): 236 (1860). - Stereocaulon macrocarpum RICH. Voyage de 

Dkouvertes de l'Astrolabe, execut6. . . pendant les annks 18261829 sous le cornmandement de M. 

Dumont d'u~ille, Botanique, I : 34 (1832); DODGE, Ann. Cryptog. Exot. 2 (2): 139 (l$29)61. - Stereocaulon 

ramulosum [subsp.]** macrocarpum (RICH.) HUE, NOUV. Arch. Mus. Hist. Nat. ser. 3, 2: 245 

(sep. 37) (1890). - Stereocaulon macrocarpum [f.] a. genuinum TH. FR. De Stereoc. et Pilophor. Comment. 

: 22 (1857). 

Typus: New Zealand, "loco dicto havre de l'Astrolabe", leg. A. RICHARD (date ?) (PC-HUE, holotypus). 

This is the specimen cited as type by DODGE, 1929, p. 139, and is labelled: "Echantillon figur6. 

Nouvelle Zelande. Stereocaulon macrocarpum No. 4. Herbarium Richard"; in red ink has been added, 

in HUE'S handwriting, the number 234. 

Facultative synonyms: Stereocaulon proximum [subsp.]** macrocarpoides NYL. Synopsis Methodica 

Lichenum, 1 (2): 238 (1860). - Stereocaulon proximum var. macrocarpoides (NYL.) NYL Ann. Sci. Nat., 

Bot. ser. 4, 15: 371 (1861). - Stereocaulon macrocarpoides (NYL.) KREMP. Lichenes, in Reise S. M. 

The description given by DODGE 

appears to be a composite one, referring partly to St. implexum 

TH. FR.


Fregatte Novara, Bot. Theil, 1 (2): 124 (1870). - Stereocaulon ramulosum var. macrocarpoides (NYL.) 

MULL. ARG. Bull. Herb. Boissier, 2, Append. I: 22 (1894). 

Exsicc.: FOLLM. Lich. Exs. Sel. Mus. Hist. Nut. Casselensi Fasc. VII (1975) no. 137 (FH, KASSEL). 

-Krypt. Exs. Vindob. Cent. XLIV (1963) no. 4339 (FH, LD, s). - V~ZDA, Lich. Sel. Exs. Fasc. XLVIII 

(1973) no. 1191 (FH). 

Icon.: RICH. 1832-33, P1. IX, fig. 4, as St. macrocarpum. - DUVIGN. 1956, fig. 10 A, as St. macrocarpum 

(apothecial structure, asci, spores). 

Remarks: phyllocladia absent or almost so in upper parts of pseudopodetia, there replaced 

by a wrinkled and interrupted cortical layer. This character is however somewhat 

inconstant, and some specimens seem to afford transitions to f. elegans and var. pulvinare. 

The apothecia are often quite large, up to 4 or 5 mm dim., but sometimes reach this size also 

in the two other taxa mentioned. Most typically developed in New Zealand and the subantarctic 

islands, Tierra del Fuego and southern Chile, but material apparently referable 

here has also been seen from New Caledonia. the New Hebrides, Tahiti, Bolivia, Venezuela 

and Mexico. 

Var. pulvinare (DODGE) LAMB (n. comb.) 

ex D. C. LINDS. Brit. Antarct. Survey Sci. Repts. no. 89: 51 (1974) (comb. inval.). 

Basionym: Sfereocaulon pulvinare DODGE, Repts. B. A. N. Z. Antarct. Research Exped. 1929- 

1931, ser. B, 7: 139 (1948). 

Typus: New Zealand, South Island, Dunedin, Mac's Valley, on rock, leg. J. S. THOMSON (date?) 

(no. 9) (DODGE, holotypus). 

Faculfative synonym (doubtful): Sfereocaulon macquariense DODGE, Nova Hedwigia, 15: 289 

(1968) (see Append. 2, p. 323). 

Remarks: pseudopodetia densely caespitose, crowded and intricated, forming hard and 

rigid clumps very firmly rooted onto stones or gravelly soil. Perhaps a modification induced 

by harsh environmental conditions, and possibly not taxonomically distinct from var. macrocarpum. 

Seen from New Zealand and the subantarctic islands, Australia, Tasmania, southern 

Chile, Tierra del Fuego, and South Georgia. 

Fo. crebratum LAMB (n. f.) 

Diagn.: Habitu maxime compcto et pulvinato, phyllocladiis coralloideis apicalibus 

congestisque, superficiem crustesam efficientibus. 

Typus: New Zealand, South Island, Port Chalmers near Dunedin, on rock, leg. J. S. THOMSON 

(date ?) (no. D. 15, T. 696) (DSIR, holotypus; CAN, MO, isotypes). 

Remarks: forming pulvinate tufts of extremely compact habitus, with the coralloid 

phyllocladia apical and very crowded, forming a more or less crustose surface. Usually 

sterile. Not uncommon in New Zealand and the subantarctic islands; also seen from Tasmania 

and Juan Fernandez. 

Var. pilophoroides (TUCK.) LAMB (n. comb.) 

Basionym: Stereocaulonpilophoroides TUCK. Proc. Amer. Acad. Arts, 6: 265 (1864); DODGE, Ann. 

Cryptog. Exot. 2 (2): 134 (1929); MAGN. in MAGN. et ZAHLBR. Ark. Bot. 31A (6): 47 (1944); LAMB ex 

FREY, Bot. Jahrb. Syst. 86: 245 (1967). 

Typus: U.S.A., Hawaii, without exact locality, leg. W. HILLEBRAND, 1864 (FH-TUCK, holotypus). 

Facultative synonyms: Stereocaulon denudatum PERS. ex GAUDICHAUD in DE FREYCINET, Voyage 

autour du Monde . . . executk sur les Corvettes de S. M. 1'Uranie et la Physicienne, pendant les annks 

1817, 1818, 1819 et 1820, Botanique: 21 1 (1826) (non St. denudatum FLORKE ex anno 1819). - Stereo-


caulon mixtum var. denudatum (PERs.) MULL. ARG. Flora, 72: 60 (1889). 

Exsicc. : ZAHLBR. Lich. Rarior. Exs. no. 137 (1910), as St. proximum (FH). 

Icon.: MAGN. in MAGN. et ZAHLBR. 1945, PI. I, fig. 1, as St. pilophoroides. 

Remarks: endemic to the Hawaiian Islands. Habitually resembles the S.E. Asiatic 

species St. staufleri LAMB. When typically developed, easily recognizable by the taller stature 

(10-12 cm, or even up to 20 cm acc. to MAGNUSSON), more or less brown-pigmented hypothecium 

and more numerous septa (up to 8) in the spores, which may be up to 90 p long. 

However, these distinctive characters are not always associated, and sometimes only one or 

two are present, making distinction from the typical species and the var. macrocarpum difficult. 

Which of the three distinctive characters should be regarded as most important 

(chief-differential) is undecided; see BLANKINSHIP (1898) for a discussion of this problem as 

met with in higher plant groups. 

Var. simplicius LAMB (n. var.) 

Diagn.: Pseudopodetia simplicia vel subsimplicia, cornuto-attenuata. Habitu ut in 

St. meyeri var. acuminatum (MULL. ARG.) LAMB, sed esorediaturn. 

Typus: Brazil, Rio de Janeiro State, Organ Mts., Pedra do Sino above Teresopolis, altit. 2000 m s. 

m., on exposed gneissic rock, leg. S. F. MARKGRAF, 1952 (M, holotypus). 

Remarks: pseudopodetia erect, stipate-caespitose, several springing from a common 

holdfast, simple or very sparingly branched, cornute-acuminate, 5-8 cm long, 1-2 rnrn thick, 

terete, clothed throughout entire length with cylindric-coralloid phyllocladia. Apices esorediate. 

Pseudopodetial surface ecorticate, glabrous. Cephalodia typical of the species, containing 

Stigonema or Scytonema. Apothecia rare, terminal, 0.61.5 mm diam., conveximmarginate. 

Rare; the only other specimen seen was from Ecuador, altit. 2200 m s.m., 

leg. G. HARLING, 1947 (no. 2040, in CAN, LIL, s). 

Var. gracilius (MULL. ARG.) LAMB (n. comb.) 

Basionym: Stereocaulon proximum [var.] ,i3. gracilius MULL. ARG. Rev. Mycol. (Toulouse), 1 : 164 

(1 879) ; DODGE, Ann. Cryptog. Exot. 2 (2) : 136 (1 929). - Stereocaulon gracilius (MULL. ARG.) DUVIGN. 

Biol. Jaarb. "Dodonaea", 9: 92 (1942)82. 

Typus: Colombia, Cauca Valley, La Horqueta, near Dolores, leg. ANDRB, 1876 (no. 2813 pr. p.) 

(G-MULL, holotypus; K, isotypus). 

Remarks: pseudopodetia of small and delicate stature, not exceeding 4 cm in length, 

usually less, but otherwise well developed and often fertile. Apothecia not over 1 mm diam. 

Does not appear to be a stunted or depauperate ecotype. Pseudopodetia often strongly corticate. 

Some specimens show whitish, mealy, pseudosorediate efflorescences at the apices of 

the pseudopodetia, such as are often present also in St. rnicrocarpum MULL. ARG. This, as 

well as the resemblance in habitus, has led some authors to misidentify St. rnicrocarpum as 

the present taxon (ZAHLBRUCKNER in Krypt. Exs. Vindob. no. 667, as St. proximum var. 

gracilius; DUVIGN. 1942, as St. gracilius). Spores (3-) 45 (-6)-septate, 35-60 x3.54.0 p. 

Var. gracilius has been seen by us from Colombia, Costa Rica and Mexico. 

Var. perpumilum LAMB (n. var.) 

Diagn.: Nanum, statura etiam humiliori quam in praecedente (var. gracilius), pseudo- 

OZ The material studied by DUV~GNEAUD 

(Krypt. Exs. Vindob. no. 667), and to which he applied this 

name, is a different species, St. microcarpum MULL. ARG.


podetiis 4-14 mm longis, apotheciis minoribus (0.6-0.8 mm), sporis brevioribus, 33-43 p 

longis, 3-septatis. 

Typus: Cuba, Oriente, Sierra Maestra, along ridge E. of Gran Piedra, altit. ca. 1250 rn s. m., on 

rock, leg. H. A. IMSHAUG, 1959 (no. 25055) (FH, holotypus; MSC, isotypus). 

Remarks: similar to the var. gracilius, but much dwarfer, with pseudopodetia only 4- 

14 mm long, smaller apothecia and shorter, less septate spores. Richly branched, mostly 

smoothly corticate, with cylindric-coralloid phyllocladia both on the main and upper branches. 

As often in var. gracilius, some of the pseudopodetial apices show small whitish mealy pseudosorediate 

efflorescences. Cephalodia normal for the species, with Scytonema in the specimens 

examined. Apothecial structure typical, with hymenium ca. 75 p high. Similar in appearance 

to dwarf states of St. microcarpum M~~LL. ARG., but chemically different (atranorin and perlatolic 

acid, PD +faint yellow). 3 different collections seen, all from Sierra Maestra in Cuba, 

leg. IMSHAUG, 1959. 

Var. exalbidum (NYL.) LAMB (n. comb.) 

Basionym: Stereocaulon exalbidum NYL. Ann. Sci. Nat., Bot. ser. 4, 11: 210 (1859); DODGE, Ann. 

Cryptog. Exot. 2 (2): 120 (1929). - Stereocaulon proximum [subsp.]* exalbidum (NYL.) NYL. Synopsis 

Methodica Lichenum, 1 (2): 237 (1860). 

Typus: Peru, Prov. Carabaya, San Juan del Oro, leg. WEDDELL (date ?) (H-NYL, no. 39773, holotypus). 

Remarks: pseudopodetia and larger phyllocladia ecorticate, spongy-tomentose, white or 

chalky-whitish, sometimes with a slight roseate tinge, in the fresh condition, but becoming 

pallid brownish or cream-colored in old herbarium specimens. Apothecial structure, spores 

etc. typical for St. ramulosum. Similar in color and tomentosity to the following variety 

(var. submollescens). Common in Peru and Bolivia; also seen from Ecuador. 

Var. submollescens (NYL.) LAMB (n. comb.) 

Basionym: Stereocaulon ramulosum [subsp.]** submollescens NYL. Compt. Rend. Hebd. Seances 

Acad. Sci., Paris, 83: 87 (1876); Lichenes Novae Zelandiae: 16 (1888); DODGE, Ann. Cryptog. Exot. 

2 (2): 146 (1929).R3 

Typus: Campbell Island (subantarctic island of New Zealand), growing among mosses and hepatics, 

leg. FILHOL, 1874 (H-NYL, no. 40089, holotypus). 

Remarks: chalky-whitish, often with a faint roseate tinge, in fresh material; fragile; 

younger parts of pseudopodetia and phyllocladia matt, with soft, spongy or tomentose surface. 

Pseudopodetia 7-1 1 cm long, loosely entangled, subsimple to moderately branched. Mature 

apothecia rare, but immature initials commonly present. The coralloid phyllocladia are of 

very small size relative to the pseudopodetia, less than 1 mm long, and this seems to be the 

principal distinguishing character from var. exalbidum. Usually growing in moss tufts or in 

carpets of Coprosma pumila, in which the lower parts are concealed. Common in the subantarctic 

islands (Campbell, Antipodes, Macquarie) and one quite typical specimen has been 

seen from New Zealand (South Island, Arthur's Pass, leg. H. A. IMSHAUG, 1971, in MSC).~~ 

- . - - - -- -- - - - -- -- - - 

63 AS Stereocaulon submollescens; a lapsus rather than an intentional new combination. ZAHLBRU- 

CKNER, 1926-27 (1927), p. 668, also lists it as St. submollescens. 

O4 The description by DODGE (1948, p. 139) of "Stereocaulon submollescens" from New Zealand 

refers to the typical St. ramulosum.


95. Stereocaulon argus HOOK. F. et TAYL. 

London J. Bot. 3: 653 (1844); in HOOK. F. The Botany of the Antarctic Voyage of H. M. Discovery 

Ships Erebus and Terror in the Years 1839-1843 . . . Part I. Flora Antarctica. Part i: 196 (1845). 

Emend. TH. FR. De Stereoc. et Pilophor. Comment.: 22 (1857)s6; Monogr. Stereoc. et Pilophor.: 333 

(sep. 29) (1858); MULL. ARG. Flora, 71: 19 (1888). - Stereocaulon macrocarpum [var.] B. argus (HOOK. 

F. et TAYL.) TH. FR. De Stereoc. et Pilophor. Comment.: 22 (1857). 

Typus: Campbell Island (subantarctic island of New Zealand), leg. J. D. HOOKER, 1840, pr. p. 

(FH-TAYL, lectotypusBB; UPS-TH FR, isolectotypus). 

Facultative synonym: Stereocaulon argodes NYL. Compt. Rend. Hebd. S6ances Acad. Sci., Paris, 

83: 87 (1876); DODGE, Ann. Cryptog. Exot. 2 (2): 108 (1929). 

Icon.: HOOK. F. 184547, PI. LXXIX, fig. 11, 1-9.07 - TH. FRIES 1858, PI. VII, fig. 4 (hymenium, 

ascus, spores). - DUYIGN. 1956, fig. 9,2, as St. argodes. 

Mat. chim. : atranorin, perlatolic acid, probably stenosporic acid, and traces of unidentified 

substances. HUNECK (1974) reported atranorin and norstictic acid, but subsequently (in litt.) 

informed us that the specimen investigated contained perlatolic acid, not norstictic acid. 

Distrib. : subantarctic islands of New Zealand (Campbell and Macquarie Islands); also Stewart 

Island, just S. of New Zealand. Unverified record: Chile (NYLANDER, 1855), probably referring to St. 

dusenii LAMB. 

Remarks: apothecia large, 3-6 mm diam., pseudolecanorine with wrinkled paler margin,B8 

later tending to become convex-irnmarginate. Hypothecium brownish-nubilated and opaque 

right up to the base of the hymenium. Hymenium 80-100 (-135) p high. Spores fusiforrn, 

stout, (3-) 5-7 (-9)-septate, 35-60 (-75)x(4-) 5-7 p. Pycnoconidia filiform, slightly to 

moderately curved, 12-15 x ca. 0.4 p. Cephalodia pale, resembling those of St. ramulosum 

(Sw.) RAUSCH., but smoother and more globose, less deeply scrobiculate-indented, containing 

Nostocoid algae. 

Two specimens collected on Campbell Island by H. A. IMSHAUG in 1969 (nos. 46382, 

46405, in MSC) resemble St. argus, but are smaller, contain atranorin and fumarprotocetraric 

acid (PDSorange-red) and have large, smoothly subglobose cephalodia. They differ from 

St. dusenii LAMB, which has similar chemical constitution, in the form of the cephalodia. 

Probably an undescribed species, but requires further investigation. 

96. Stereocaulon atlanticum (LAMB) LAMB (n. comb.) 

Basionym: Stereocaulon meyeri subsp. atlanticum LAMB ex POELT, BestimmungsschliisseI Europaischer 

Flechten: 635 (1969). 

Typus: Azores, S. Jorge, on rock, leg. SPINOLA (date ?) (no. 109) (UPS-MAGN, holotypus; FH, isoty- 

PUS). 

Mat. chim. : the full chemical spectrum consists of atranorin, stictic acid, constictic acid, 

norstictic acid, possibly connorstictic and consalazinic acids, perlatolic acid, possibly anziaic 

acid, ursolic acid, and 5 unidentified substances. There are several deficient phases, the 

commonest with atranorin, stictic acid and norstictic acid (the type specimen belongs here). 

Distrib. : Azores, central Africa (Kenya, Rwanda-Burundi, Tanzania, Uganda, Zaire), South Africa 

Bs AS St. macrocarpum [var.] B. argus. 

Consists partly of Argopsis megalospora TH. FR.; considered lectotypified on the Stereocaulon 

component by TH. FRIES'S emendation, 1857. See LAMB, 1974, p. 450452. 

B7 Illustration based partly on St. argus and partly on Argopsis megalospora TH. FR. 1 and 2, showing 

the habitus, seem to refer to St. argus, but 8 and 9, "asci", obviously represent spores of the Argopsis. 

ZAHLBRUCKNER, 

1926-27 (1927), p. 673, placed the species in subgen. Lecanocaulon.


Costa Rica, Venezuela, Ecuador, Peru, Bolivia. 

Remarks: resembling St. meyeri B. STEIN, but of smaller stature, with corticated pseudopodetia 

of "macrocarpoid" habitus and mostly harmonic phyllocladial branchlets, and 

distinct in chemical constitution. Closely related to St. microcarpum MULL. ARG., from which 

it differs in the possession of well developed true soredia and in its simpler chemical spectrum. 

Morphologically it appears to be indistinguishable from St. rugulosum LAMB, which is chemically 

different (furnarprotocetraric acid) and has a less extensive, although partly overlapping 

geographical distribution. St. atlanticum is rarely fertile. The record of "Stereocaulon 

ramulosum var. farinosum" from the Azores by DEGELIUS (1941) refers to this species. 

97. Stereocaulon colensoi BAB. 

ex HOOK. F. The Botany of the Antarctic Voyage of H. M. Discovery Ships Erebus and Terror in the 

Years 1839-1843 . . . Part 11. Flora Novae-Zelandiae. Part ii. Flowerless Plants: 295 (1855); 

NYL. Synopsis Methodica Lichenum, 1 (2): 232 (1860); HOOK. F. Handbook of the New Zealand Flora, 

Part 11: 562 (1867); DODGE, Ann. Cryptog. Exot. 2 (2): 110 (1929); MARTIN et CHILD, Lichens of New 

Zealand: 155 (1972). - Pilophoron colensoi (BAB.) KNIGHT, Trans. & Proc. New Zealand Inst. 16: 400 

(1 883) ; ZAHLBR. Catalogus Lichenum Universalis, 4 : 433 (1 926). - Corynophoron colensoi (BAB). 

NYL. Lichenes Novae Zelandiae: 15 (1888).80 

Typus: New Zealand, North Island, leg. COLENSO (date ?) (no. 2746) (K, holotypus; H-NYL, isotypus). 

Icon. : HOOK. F. 1855, PI. CXXX A. - NYL. 1860, PI. VII, fig. 8 (ceph. algae). - DUVIGN. 1956, 

figs. 4 (3, 6. - MARTIN and CHILD 1972, PI. 47, lower fig. 

Mat. chim. : atranorin, colensoinic acid, D-mannitol, D-arabitol, unidentified tetrahydroxy 

fatty acids, and traces of 5 further unidentified substances. Colensoinic acid, described from 

St. colensoi by Fox, KLEIN and HUNECK, 1970, is the depsidone analogue of the depside 

perlatolic acid, and has since been found also in some other Stereocaulon species. 

Distrib. : New Zealand and Campbell Island. Erroneous record: Fuegia (RASANEN, 1932, =St. 

glabrum (MOLL. ARG.) VAIN.). 

Remarks: a notable feature of typical St. colensoi are the peculiar cephalodia, which 

DUVIGNEAUD (1956) terms "ficoid". They are pyriform-clavate, shortly stalked, usually 

with a subaeruginose tinge, with finely granulate to cerebriform-wrinkled surface, solid internally, 

containing Stigonema or Scytonema. Their cortex is 15-50 p thick, isabelline-nubilated 

and semi-opaque in section, of indistinctly pseudoparenchymatous (not palisadic) structure 

with more or less isodiametric, thin-walled cells 3-5 p diam. The structure of the cephalodial 

cortex in St. colensoi is unique in being persistently of the primary type, not proceeding to the 

development of gelatinized-palisadic tissue as in St. implexum TH. FR. (see LAMB, 1976, p. 358). 

Young immature cephalodia may be subglobose with almost smooth surface. In var. caliginosum 

the cephalodia are of somewhat different appearance (see below). Pseudopodetia 

corticated with verrucose-wrinkled thallus-mantle, and often "macrocarpoid" in habitus. 

Apothecia terminal, commonly several clustered, 1.5-3.0mm diam., at first with prominent 

paler receptacle (pseudothalline margin) which later becomes more or less reflexed and excluded. 

Central cone densely isabelline-nubilated, opaque and compact. Hypothecium 

nubilated below, more or less hyaline in upper subhymenial part. Hymenium 9LL120 (-135) p 

high. Spores (4) 8-12-septate, finally somewhat constricted at the septa, (35-) 65-90 x 4-6 p. 

@@ The epithet Corynophoron was first published by NYLANDER, 1860, as "fere genus proprium", 

and 1888 (loc. cit.) as a proper genus, comprising St. colensoi only, mainly on the basis of the peculiar 

cephalodia.


Conidia arcuate, 9-12 ~0.5-0.7 p. KNIGHT (1883, P1. XXXIX, fig. 1) illustrated an "ascus" 

of "Pilophoron colensoi" supposedly containing 8 uniseriate simple ellipsoid spores. He 

probably mistook a single spore, constricted at the septa, for an ascus. 

Var. caliginosum LAMB (n. var.) 

Diagn. : Distinguitur cephalodiis subglobosis, simplicibus vel vulgo tuberculato-conglomeratis, 

mox obsolete verruculosis aut minute areolato-verruculosis (haud cerebriformicorrugatis), 

breviter pedicellatis aut sessilibus, intrinsecus solidis et dense griseo- vel nigricantinubilatis 

opacisque. 

Typus: New Zealand, North Island, Kaimanawa Range, Thunderbolt Peak, altit. 1500 m s. m., 

On rock, leg. D. J. GALLOWAY, 1972 (FH, holotyp~s; GALWWAY, ~SO~YPUS). 

Remarks: cephalodia both basal and lateral on pseudopodetia, concolorous with the 

phyllocladia or somewhat darker (grayish or brownish-gray), irregularly subglobose, 0.7- 

2.0 rnrn diam., sessile or shortly stalked, single or usually agglomerated into compound masses 

3-5 (-8) rnrn diarn., at first more or less smoothly corticate, soon becoming obsoletely verruculose 

to minutely areolate-verruculose, not cerebriform-wrinkled. Their cortical structure 

as in the typical species, but their internal tissue densely dark gray or blackish and opaque in 

section (not relieved by KHO). Chemistry, apothecial structure and spores as in var. colensoi. 

Known from 2 different localities in the Kaimanawa Range of North Island, New Zealand. 

98. Stereocaulon corticatulum NYL. 

Flora, 41: 117, footnote (1858); Synopsis Methodica Lichenum, 1 (2): 240 (1860); Lichenes Novae 

Zelandiae: 15 (1888); LINDS. Trans. Linn. Soc. London, 25: 529 (1866); DODGE, Ann. Cryptog. Exot. 

2 (2) : 1 14 (1929) ; MART^ et CHILD, Lichens of New Zealand : 155 (1972), pr. p. 

Typus: New Zealand, exact locality not stated, "supra saxa calcarea", leg. COLENSO (date ?) (no. 

5144) (H-NYL, no. 40026, holotypus). 

Facultative synonyms: Stereocaulon Ieptaleum NYL. Synopsis Methodica Lichenum, 1 (2): 251 

(1860). - Stereocaulon corticatulum [subsp.]* detergens NYL. Lichenes Novae Zelandiae: 16 (1888). 

- Stereocaulon corticatulum var. detergens (NYL.) MULL. ARG. Bull. Herb. Boissier, 2, Append. I: 

22 (1894).70 

Icon. : MARTIN and CHILD 1972, PI. 32, upper right hand figure (field photograph in color).71 - 

LAMB 1976, fig. 3 (cephalodia). 

Mat. chim. : atranorin, rangiformic acid, norrangiformic acid, an unidentified fatty acid 

and an unidentified pigment; the chemical constitution is the same as that of St. delisei BORY. 

Distrib.: New Zealand and its subantarctic islands, Australia, Tasmania, Argentina (NeuquBn, 

Rio Negro, Chubut, Tierra del Fuego), Chile (Magallanes, Aysen, Ruble, Juan Fernandez), South 

Africa (var. capense), Tristan da Cunha, Gough Island. Erroneous record: Lapland and North 

America (OZENDA and CLAUZADE, 1970; based on a confusion with St. capitellatum MAGN. and other 

species). 

Remarks: NYLANDER correctly compared St. corticatulum with St. delisei BORY; the two 

species are in fact very closely related. Other closely related species are St. wadei LAMB and 

St. melanopotamicum LAMB. St. corticatulum is a small species, often not over 1 cm high, 

ZAHLBRUCKNER, 1926-27 (1927), p. 646, lists it as "Stereocaulon detergens NYL.", a lapsus, not 

an intentional new combination. 

Another photograph, PI. 46, lower figure, said to be of St. corticatulum, obviously represents a 

different species, possibly St. ramulosum var. submollescens (NYL.) LAMB. The description of St. 

corticatulum given by MARTIN and CHILD is a composite one.


but may reach 3 cm or slightly over. Pseudopodetia partly or largely corticate, glabrous, 

rarely becoming more or less ligneous. Phyllocladia often very indistinctly developed; most 

of the assimilation seems to be carried on by the corticate thallus-mantle. Apical soredia 

often present but not constant. Cephalodia very variable in appearance, often not outwardly 

recognizable as of the sacculate type; pale (concolorous with the thallus-mantle) to brownish, 

subglobose and divided into botryose verruculae, indented-corrugated, or more rarely distinctly 

scrobiculate-foveolate. The depressions on their surface are often darker (aeruginoseglaucescent). 

They are of loose-cored sacculate structure, and their cortical layer is dimorphic, 

showing alternation of two different types of tissue (LAMB, 1976). Apothecia up to ca. 1 mrn 

in diameter, becoming convex. Hymenium 60-80 p high. Spores 8, straight, acicularfusiform, 

3(-5)-septate, (28-) 35-50 (-65) x 34 (4.5) p. Pycnoconidia filiform, curved to 

almost straight, 8-13 x ca. 0.5 p. 

Var. humile (MULL. ARG.) LAMB 

ex FREY, Bot. Jahrb. Syst. 86: 244 (1967). - Stereocaulon humile MULL. ARC. Bull. Herb. Boissier, 4: 

88 (1896); DODGE, Ann. Cryptog. Exot. 2 (2): 123 (1929). 

Typus: Australia, Victoria, "verisimiliter in cacuminibus montium", leg. KNIGHT, 1887 (no. 60) 

(G-MULL, holotypus). 

Remarks: pseudopodetia dwarf, 4-12 (-15) mm long, upright or subdecumbent, forming 

a crust, more or less sorediate at their apices; phyllocladia indistinct. Seen from Australia, 

Tasmania, New Zealand and its subantarctic islands, and Chile (Magallanes). 

Var. complanatum LAMB (n. var.) 

Diagn. : Habitu applanato, dorsiventrali, apicibus pseudopodetiorum expansis, spathulatis, 

in latere ventrali sorediosis; soredia vulgo ramulis perminutis, congestis, leprosis suffulta (ut 

in St. delisei BORY). 

Typus: New Zealand, South Island, Otago, Taieri Mouth, on coastal rock face, leg. J. MURRAY, 

1958 (no. 1421) (FH, holotypus). 

Remarks: characterized by the flattened, dorsiventral, spathulate habitus, with soredia 

on the ventral side of the expanded apical portions, the general habitus resembling that of a 

very short St. spathuliferum VAIN. or a coarse St. nanodes TUCK.; the soredia commonly borne 

on minute, crowded, leprose branchlets, as in St.delisei BORY. The nature of the soredia in 

this variety affords an additional indication of the close affinity, already shown by the identical 

chemistry, between St. corticatulum and St. delisei. 2 specimens seen, from New Zealand and 

Australia. LAUDER LINDSAY (1866, p. 529) mentions a condition of St. corticatulum from 

New Zealand which must refer to this variety: "a condition of minute branching and subfarinosity 

of the terminal divisions of the podetia." 

Var. capense (LAMB) LAMB (n. comb.) 

Basionym: Stereocaulon capense LAMB, Lilloa, 26: 416 (1953). 

Typus: South Africa, Western Cape Province, Tulbagh Division, Great Winterhoek, altit. 1500 m 

s. m., on rock face above a stream in steep gully, leg. E. ESTERHUYSEN, 1951 (no. 19862 pr. p.) (CAN, 

holotypus; BOL, isotypus). 

Icon. : LAMB 1953, PI. 11, fig. 5. 

Remarks: distinguished by its more dorsiventral habitus with the terminal branches 

tending to be more or less strongly flattened and flabellate-squamulose, somewhat resembling


the phyllocladia of St. evolutum GRAEWE or certain states of St. sterile (SAV.) LAMB. Soredia 

present or absent. Several collections seen from South Africa (Table Mountain, Western 

Cape Province, Natal). 

Var. subcorticatum (RAs.) LAMB 

Canad. J. Bot. 29: 581 (1951). - Stereocaulon subcorticatum RAS. Ann. Bot. Soc. Zool. Bot. Fenn. 

Vanamo, 2 (1) : 24 (1932). 

Typus: Chile, Magallanes, Fjordo de Agostini, on rocks at seashore, leg. H. ROIVAINEN, 1929 

(H-R~S, holotypus; S, isotypus). 

Remarks: pseudopodetia almost entirely clothed with corticate thallus-mantle. Often 

fertile, but seldom sorediate. Grows preferentially in more shaded locations. Rather 

common; seen from Chile (Magallanes, Aysen, Juan Fernandez), Argentina (Tierra del 

Fuego, Rio Negro), Australia, Tasmania, and Gough Island in the south Atlantic. 

99. Stereocaulon crambidiocephalum LAMB (n. sp.) 

ex VARGAS, Revista Univ. (Cuzco), no. 94: 3 (1948) (nomen nudum). 

Diagn. : Pseudopodetia erecta, substrato arcte affixa, 4.5-8.0 cm alta, robusta, ramis 

majoribus crassit. 2.Ck3.5 mm, apicalibus 0.8-1.2 mm; parce subdichotomice ramosa, albida, 

fere in toto decorticata et subfarinosa; apicibus dense corymbiformiter sorediosis, sorediis 

ramulis tenuibus congestis suffultis et capitula 2-6 mm lata formantibus. Phyllocladia parce 

evoluta (praesertim basin versus, superne fere deficientia), cylindrico-coralloidea, vulgo 

ramosa, longit. 0.8-2.0 (-3.0) mm, divaricata, corticata, partim parce subfarinosa. Cephalodia 

jam basalia, jam lateraliter insidentia, cinereo-albida, subglobosa, sacculata, 1.0-2.3 mm lata, 

scrobiculato-foveolata vel cerebriformia. (Apothecia et pycnidia ignota.) Atranorinum, 

acidum perlatolicum et acidum anziaicum continet (KSbene flavescens, PDSpallide flavescens). 

-A St. meyeri B. STEIN, cui peraffine, habitu magis robusto, pseudopodetiis decorticato-farinosis, 

phyllocladiis superne rarescentibus aut nullis, et praecipue soraliis apicalibus 

multoties majoribus et ramulis minutis congestis suffultis differt. 

Typus: Peru, Dept. Cuzco, Prov. Paucartambo, Llulluchayocc, altit. 3200 m s. m., on rock, leg. C. 

VARGAS, 1944 (no. 4301) (VARGAS, holotypus; CAN, MICH, isotypes). 

Mat. chim. : atranorin, perlatolic acid, anziaic acid. 

Distrib. : Peru and Colombia. 

Remarks: closely related and similar to St. meyeri B. STEIN, differing chiefly in larger 

and more robust stature and large apical heads of soredia borne on fine congested branchlets. 

The cephalodia, containing Stigonema, are similar to those of St. ramulosum (Sw.) RAUSCH. 

Rare; known from 4 collections. 

100. Stereocaulon delisei BORY 

ex DUBY, Aug. Pyrami de Candolle Botanicon Gallicum, ed. 2, Pars secunda: 619 (1830); SCHAER. 

Enumeratio Critica Lichenum Europaeorum: 178 (1850); OLIV. Flore Analytique et Dichotomique 

des Lichens de lYOrne, Part 1 : 59 (1882); CROMB. A Monograph of Lichens found in Britain: 117 (1894); 

HARM. Lichens de France, Fasc. 111: 363 (1907); A. L. SM. A Monograph of the British Lichens, ed. 

2, Part I: 410 (1918); MAGN. Goteborgs K. Vetensk.- o. Vitterh.-Samh. Handl. ser. 4,30 (7): 44 (1926); 

Flora over Skandinaviens Busk- och Bladlavar: 67 (1929); DODGE, Ann. Cryptog. Exot. 2 (2): 116 

(1929); FREY, Rabenhorsts Krypt.-F1.9, Abt. IV, 1. Halfte: 82, 106 (1932); LAMB, J. Bot. 77: 76 (1939); 

MOT. Porosty (Lichenes), in Flora Polska, 3 (2): 378 (1964); DAHL et KROG, Macrolichens of Denmark,


Finland, Norway and Sweden : 133 (I 973). - Stereocaulon coralloides [subsp.]' delisei (BORY) NYL. 

Synopsis Methodica Lichenum, 1 (2): 242 (I 860). - Leprocaulon delisei (BORY) OLIV. Expod Systkmatique 

et Description des Lichens de l'Ouest et du Nord-Ouest de la France, 2 (4): 378 (1903). 

Typus: France, Dept. Calvados, Vire, leg. DELISE and DESPREAUX (PC, fide DODGE, 1929, p. 116; 

not seen by us. We have however studied the authentic, possibly isotype material distributed in DEL. 

Lich. de France no. 18, from the same locality). 

Exsicc.: DEL. Lich. de France Fasc. I (1828) no. 18 (authentic, possibly isotype material) (FH). - 

HAV. Lich. EXS. Norveg. no. 608 (1939) (BG, WS). 

Icon.: FREY 1932, fig. 14. - KERSH. 1960, fig. 6. - Oz. et CLAUZ. 1970, fig. 387 C. - DAHL et 

KROG 1973, fig. 40. - NYL. 1860, PI. VII, fig. 17 (ceph. algae), as St. coralloides subsp. delisei. 

Mat. chim. : atranorin, rangiformic acid, norrangiformic acid, and an unidentified fatty 

acid. 

Distrib.: France, England, Scotland, Norway, Sweden, Madeira, South Africa. Unverified record: 

Italy (JATTA, 1909-11). 

Remarks: a rare suboceanic species, closely related to St. corticatulum NYL.; the only 

member of the subgenus Holostelidium found in Europe. Cephalodia (not always present) 

similar in appearance and structure to those of St. corticatulum. Apothecia were previously 

unknown in this species, but a specimen in BM from Scotland, Argyll, Strontian, on boulders 

from lead mine spoil heaps, leg. P. W. JAMES, 1962 (no. 295) is fertile. Apothecia scarce, 

terminal, up to ca. 1 mrn diam., convex, irnrnarginate. Central cone compact, sordid isabelline; 

hypothecium colorless, hyaline; hymenium (somewhat immature) ca. 60 p high. (No 

mature asci or spores found.) 

10 1. Stereocaulon didymicum LAMB (n . sp .) 

Diagn.: Pseudopodetia caespitosa, erecta aut partim subdecumbentia, substrato arcte 

affixa, 2.0-3.5 cm longa, 0.8-1.0 mm crassa, teretia, subsimplicia vel irregulariter aut subdichotomice 

ramosa, vel in toto corticata vel partim decorticata, omnino glabra, sordide 

cinerascentia. Phyllocladia (passim parcissime evoluta) cylindrico-coralloidea, simplicia vel 

parce ramosa, longit. 0.5-1.5 mm, crassit. ca. 0.2 mm, laevia et corticata, in partibus superioribus 

pseudopodetiorum breviora et verruculosa aut passim omnino deficientia (habitu macrocarpoideo). 

Cephalodia lateralia, sessilia subsessiliave, latit. 0.7-1.5 mm, isabellino-albescentia 

aut cinerascentia, haud nitida, primum laeviuscule subglobosa, dein mox scrobiculato-corrugata 

(ut in St. ramuloso), sacculata, intus laxe contexta, cortice 30-60 p crasso, 

nec palisadico nec gelatinoso, sordide nubilato et lamina tenui viso fere opaco, hyphis valde 

pachydermaticis 5-9 p latis contexto. Apothecia terminalia, 0.8-2.0 (-2.3) mm lata, e plano 

mox pileato-convexa et immarginata, fusconigricantia. Conus centralis sat compactus et 

dense cinerascenti- aut isabellino-nubilatus. Hypothecium sordide nubilatum aut fuscescens 

sed haud vere pigmentiferum (KHO fere decoloratum). Hymenium 75-90 {L altum. Sporae 

rectae aut subrectae, 34-septatae, aciculari-fusiformes, longit. 32-50 p, crassit. 3.0-3.5 p. 

Atranorinum, acidum didymicum, acidum sticticum et substantiam ignotam acido didymico 

affinem geret (KSsubflavescens, PDi-aurantiaco-rubescens). - Habitu statum minorem St. 

ramulosi var. macrocarpi (RICH.) BAB. in memoriam revocans, St. rubiginoso PERS. vere arctius 

affine (ut e structura corticali cephalodiorum videtur), sed materiis chimicis et distributione 

geographica diversum. 

Typus: Costa Rica, Prov. Cartago, between Palmital and La Sierra, altit. 1800 m s. m., on steep 

stony roadside soil bank, leg. I. M. LAMB and U. V. METZGER, 1969 (FH, holotypus; DUKE, isotypus). 

Mat. chim.: atranorin, didymic acid, stictic acid, and an unidentified substance related to


didymic acid. 

Distrib. : Costa Rica and Ecuador. 

Remarks: this is the only species of Stereocaulon known to contain the dibenzofuran 

didymic acid, and appears to be most closely related to the Hawaiian species St. rubiginosum 

PERS. (incl. St. roccelloides (TH. FR.) LAMB), having the same peculiar cephalodial structure as 

the latter. The cephalodial cortex in both these species has in effect the structure of pseudopodetial 

cortex, being sordid brownish- or isabelline-nubilated, mostly semi-opaque, not 

gelatinized-conglutinated or palisadic but composed of randomly interwoven or vaguely vertically 

subparallel, very thick-walled hyphae compacted together but distinct and separate, 

those on the surface emerging separate and discrete. Algae Scytonema or Stigonema. In 

the type specimen the apices of some of the sterile pseudopodetia bear terminal, whitish, 

granulose, pseudosorediate efflorescences, similar to those seen in St. microcarpum MULL. 

ARG. Morphologically the new species resembles a small state of St. ramulosum (Sw.) 

RAusc~. or its variety macrocarpurn (RICH.) BAB. Known from 2 collections only; that from 

Ecuador (FH, MSC) was collected by G. W. PRESCOTT in Prov. Carchi, 1953 (no. An L-5). 

102. Stereocaulon dusenii LAMB (n. sp.) 

Diagn.: Pseudopodetia robusta, rigida, erecta, caespitosa, substrato arcte adnata, 4-7 

(-8) cm longa, 1.0-2.5 (4) mm crassa, teretia aut partim leviter complanata, parce modiceve 

irregulariter ramosa, albida aut cremea, in partibus superioribus corticata, pallio thallino continuo, 

verrucoso aut rugoso vestita, tantum partibus infimis denudata et ecorticata. Soredia 

nulla. Phyllocladia cylindrico-coralloidea, valida, crassit. 0.2-0.4 mm, teretia, vulgo nonnihil 

toruloso-verrucosa, majora ramosa et longit. 4(-5) mm attingentia; pseudopodetia aequaliter 

vestientia aut passim superne parum evoluta. Cephalodia lateralia, vulgo breviter pedicellata, 

subglobosa, plumbeo-cinerascentia, olivaceo-cinerea aut subalbida, 1-2 (-2.5) mm lata, scrobiculato-corrugata 

(ut in St. ramuloso), sacculata, intus laxe contexta, cortice hyalino aut 

isabellino-nubilato, conglutinato-palisadico. Apothecia terminalia, 1.5-3.0 (-5) mm lata, 

mox pulvinato-convexa et immarginata, disco fusconigricanti, aetate nonnumquam divisa et 

discos secundarios proferentia, receptaculo pseudothallino pallido rugosoque reflex0 inferne 

bene conspicuo. Conus centralis cinerascens et subopacus, laxe aut compacte contextus. 

Hypothecium parte basali sordide isabellino-nubilatum, parte superiori (50-100 p) incolor et 

hyalinum. Hymenium 90-100 p altum. Sporae Clnae, rectae, cylindrico-fusiformes, 3-6 

(-8)-septatae, longit. 35-60 p, crassit. 4.0-5.5 (-6) p. Pycnidia infra apothecia sita, maculis 

nigricantibus indicata; conidia filiformia, arcuata, longit. 9-15 p, crassit. 0.3-0.5 p. Atranorinum, 

acidum fumarprotocetraricum, acidum protocetraricum, acidum perlatolicum et 

acidum anziaicum continet (K+subfuscescens, PDSminiato-rubescens, saltem in strato 

medullari). -Simile St. argo HOOK. F. et TAYL. et St. stenospermo (LAMB) LAMB, ab iis 

materiis chimicis aliisque notis discrepans. 

Typus: Chile, Magallanes, Isla Desolacibn, Puerto Angosto, altit. ca. 150 m s. m., on rocks, leg. 

P. DUSBN, 1896 (no. 186) (s, holotypus; CAN, UPS, isotypes). 

Mat. chim.: atranorin, fumarprotocetraric acid, protocetraric acid, perlatolic acid, 

anziaic acid. 

Distrib. : Chile (Magallanes, AysBn, Juan Fernandez) and Argentina (Tierra del Fuego). 

Remarks: closely related and similar morphologically to St. argus HOOK. F. et TAYL. em. 

TH. FR., and especially to St. stenospermum (LAMB) LAMB, both of which differ in chemistry 

and geographical distribution. NYLANDER'S record of St. argus from Chile (1855) probably


refers to this species. There is a specimen of St. dusenii in s said to be from North America, 

labelled "Menzies misit. L. rarnulosus Sw. N. America Herb. Swartzii. Stereocaulon ramulosum 

Achar.", and annotated by TH. FRIES "St. macrocarpum A. Rich.". St. dusenii is sometimes 

"macrocarpoid" in habitus, with the upper parts of the pseudopodetia more or less 

destitute of phyllocladia. The cephalodia contain Stigonema or Scytonema. 

103. Stereocaulon fronduliferum LAMB (n. sp.) 

ex MARTIN, Trans. Roy. Soc. New Zealand, Bot. 3 (8): 152 (1966) et MARTIN et CHILD, Lichens of New 

Zealand : 158 (1972) (nomen nudum). 

Diagn. : Pseudopodetia arcte affixa, plus minusve erecta, ad 3.5 cm alta (sed saepe minora), 

ramosa, teretia aut leviter complanata, habitu dorsiventrali (uno latere corticata, glaucescentia, 

saepe transversim rugosa aut fissa, altero latere decorticata, glabra, ochraceo-albida), ramificatione 

bilaterali, distincte pinnatifida. Rarnuli phyllocladoidei complanati, simplices aut 

flabellatim divisi, superficie dorsali glauco-corticati, ventrali pallidiores; longit. 1-2 mm, sed 

basin pseudopodetiorum versus longiores. Soredia nulla. Cephalodia lateralia, sessilia aut 

breviter pedicellata, subglobosa, 0.7-1.3 mm lata, primo laevia, dein leviter aut sat distincte 

foveolata; fuscescentia, subnitida, intus solida (protosacculata), algas Nostocaceas continentia, 

cortice 40-50 p crasso, incolori aut pallide isabellino, palisadico. Apothecia terminalia, fere 

ab initio hemisphaerico-convexa et immarginata, tandem fere globosa, 1.0-1.5 (-2.0) mm lata, 

fusconigricantia, subnitida opacave. Conus centralis laxe contextus, hyphis hyalinis saepe 

materia viridiflava et granulosa (KHO dissolventi et solutionem aurantiaco-rubescentem 

effundenti) inspersis. Hypotheciurn pigrnentiferum (fuscum, KHO fere immutatum), ad 

150 p crassum. Hymenium 90-110 (-120) p altum. Sporae (6) 8nae, rectae vel subrectae, 

cylindrico-fusiformes, (4) 5-9 (-10)-septatae, ad septa passim leviter constrictae, longit. 40- 

70 (-80) p, crassit. 3.5-4.5 (-5.5) p. Atranorinum, acidum fumarprotocetraricum et fortasse 

parce acidum protocetraricum continens (K - vel leviter fuscoflavescens, PD intense miniatorubescens). 

- Species valde peculiaris, phyllocladiis complanatis et flabellato-divisis fere St. 

foliolosum NYL. simulans. A cel. BABINGTON jam anno 1855 recognitum et nomine St. ramulosum 

var. compressum appellatum fuit. 

Typus: New Zealand, exact locality, collector and date not indicated, commun. LOCKE TRAVERS 

to the Exposition Universelle, Paris, 1889 (PC-HUE, holotypus, named by HUE St. ramulosum var. 

compressum). 

Facultative synonyms: Stereocaulon ramulosum var. compressum BAB. in HOOK. F. The Botany of 

the Antarctic Voyage of H. M. Discovery Ships Erebus and Terror in the Years 1839-1843 . . . Part 11. 

Flora Novae-Zelandiae. Part ii. Flowerless Plants: 294 (1855). - Stereocaulon ramulosum [subsp. 

macrocarpum] f. compressum (BAB.) HUE, NOUV. Arch. Mus. Hist. Nat. ser. 3, 2: 245 (1890), nom. 

illegit., non St. ramulosum f. compressum NYL. ex anno 1860.72 

Mat. chim. : atranorin, fumarprotocetraric acid, and probably a trace of protocetraric acid. 

Distrib.: New Zealand. Unverified records: Colombia (HUE, 1898-1901; LINDAU, 1912), perhaps 

referring to St. ramulosum f. subcompressum LAMB. 

Remarks: ZAHLBRUCKNER, Catalogus Lichenum Universalis, vol. I (1922) p. 692, published 

the combination Stereocaulon compressum, referring to a species of Sphaerophorus, by 

a lapsus, wrongly ascribing it to LINDSAY. For this reason it seems advisable to use a different 

name for the present species. Short descriptions (as St. ramulosum var. compressum) 

were given by TH. FRIES (1857, p. 12, footnote; 1858, p. 26, footnote), NYLANDER (1860, p. 

72 A facultative synonym of St. strictum var. compressum (NYL.) LAMB.


236) and DODGE (1929, p. 138). The chemical composition of the greenish-yellow granular 

pigment, colored orange-red by KHO, commonly present in the central cone tissue, is unknown; 

a similar or possibly the same substance is frequently found in the hypothecium of 

St. exutum NYL. 

104. Stereocaulon halei LAMB (n. sp.) 

Diagn.: Pseudopodetia erecta, caespites laxos formantia, simplicia vel parce ramosa, 

apicibus cornuto-attenuata, (3-) 46 cm longa, satis robusta, glaberrima, pro maj. parte 

decorticata; phyllocladia cylindrico-coralloidea, divaricata, majora ad 2.5 (-3.0) mm longa et 

ramificantia. Soredia nulla. Cephalodia subglobosa aut pulvinuliformia, pallida, tandem 

scrobiculato-contortuplicata (nullo mod0 complanata nec cristata), protosacculata, intus 

solida, Scytonema continentia. Apothecia parva, ad 1 mm lata, ramulos laterales simplicesque 

singulatim terminantia, mox convexa, haud hypophysata, margine (receptaculo) haud 

distincte evoluto. Conus centralis compactus, isabellino-nubilatus et serniopacus. Hypothecium 

angustum, incolor, con0 centrali multo crassiori impositum. Hymenium 6C80 

(-85) p altum. Sporae 6-8nae, cylindrico-fusiformes, rectae, (I-) 3-septatae, longit. 25-35 p, 

crassit. ca. 3 p. Mat. chim. : atranorinum et acidum lobaricum (phyllocladiis K flavescentibus, 

PD leviter flavescentibus). -St. pseudomassartiano LAMB habitu persimile et certe arcte 

cognatum, forma cephalodiorum et materiis chimicis praecipue differens. 

Typus: Malaya, Pahang, below Ganong Brinchang, altit. ca. 1800 m s. m., on exposed rocks and 

soil, leg. M. E. HALE, 1965 (no. 29793) (FH, holotypus; us, isotypus; isotype material distributed in 

VEZDA, Lich. Sel. Exs. no. 683). 

Exsicc.: KUROK. Lich. Rarior. et Crit. Exs. Fasc. I11 (1971) no. 141, as St. massartianum (FH, LD).- 

VBZDA, Lich. Sel. Exs. Fasc. XXVIII (1968) no. 683, as St. massartianum (isotype material) (FH). 

Mat. chim. : atranorin and lobaric acid. Traces of some unidentified substances also often 

present. 

Distrib.: Indonesia (Java, Borneo) and Malaya. 

Remarks: closely related and habitually similar to St. pseudomassartianum LAMB, but 

differing in its chemical constitution (lobaric instead of perlatolic and anziaic acids) and in 

the form of its cephalodia, which are not flattened and plate-like or cristate as in St. pseudomassartianum, 

but subglobose and finally scrobiculate-foveolate to cerebriform. The cephalodial 

structure is however similar in both species (solid-cored except immediately around 

the algae, with dimorphic cortex showing alternation of clear-palisadic and nubilated-pseudoparenchymatous 

tissue). St. halei is the unnamed species referred to in LAMB, 1976, p. 358. 

In spite of the fact that it is apparently a rather rare species, it may in some of its localities 

occur in considerable profusion. 

105. Stereocaulon implexum TH. FR. 

De Stereoc. et Pilophor. Comment.: 23 (1857); Monogr. Stereoc. et Pilophor.: 335 (sep. 31) (1858). - 

Stereocaulon ramulosum var. implexum (TH. FR.) NYL. Synopsis Methodica Lichenum, 1 (2): 236 (1860). 

Typus: 2 collections cited by TH. FRIES: Magellan Straits, leg. N. J. ANDER~SON (date ?) and Chile, 

Child, near Ancud ("Aneud"), leg. LECHLER (Pl. Magell. no. 981). As lectotype we select the former 

specimen (UPS-TH FR, lectotypus, lectotypif: nov.). 

Facultative synonym: Stereocaulon proximum var. compactius ZAHLBR. in SKOTTSBERG, The Natural 

History of Juan Fernandez and Easter Island, 2, Botany, Part 111, no. 11: 373 (1924). 

Icon.: TH. FR. 1857a, unnumbered fig., b (spore); 1858, PI. VIII, fig. 1. - NYL. 1860, P1. VII, fig. 

11 (ceph. algae), as St. ramulosum var. implexum.


Mat. chim. : 3 chemical strains are here distinguished: 

Strain I (the typical and common strain), with atranorin and lobaric acid, or sometimes 

in a deficient phase with atranorin only demonstrable. Cfr. HUNECK (1972). 

Strain 11, with atranorin and miriquidic acid. 

Strain III, with atranorin, norstictic acid and salazinic acid. Strains I1 and 111, which 

probably merit the status of independent species, are found only in the islands of the S. Atlantic 

Ocean (Tristan da Cunha, Gough Island). 

Distrib. : Strain I: Chile (from Magallanes northwards to Bio-Bio, also Juan Fernandez) and Argentina 

(from Tierra del Fuego northwards to NeuquCn). Strain 11: Tristan da Cunha. Strain 111: 

Tristan da Cunha and Gough Island. Unverified records: New Zealand (KREMPELHUBER, 1870; 

MULLER ARG., 1894; HELLBOM, 1896), all probably referring to St. colensoi BAB. Erroneous record: 

Brazil (VAINIO, 1890,=St. microcarpurn M~~LL. ARG.). The occurrence, outside of South America, of 

the morphologically indistinguishable chemical strains I1 and 111 in the S. Atlantic islands is parallelled 

by a similar distribution pattern in some other genera, e. g. certain species of Placopsis (LAMB, 1947) 

and some Pteridophytes (TRYON, 1966). 

Remarks: St. implexum attains its optimal development in the more northerly provinces 

of Chilean and Argentine Patagonia, there being of very different appearance from the stunted 

populations of Magallanes and Tierra del Fuego (such as the lectotype material). The 

important features of the species in its best development may be briefly summarized as follows: 

Pseudopodetia erect, 4.5-5.5 (-7.0) cm tall, 1-2 mm thick, glabrous, for the most part 

decorticate and subligneous, but in upper parts clothed with an interrupted or continuous, 

areolate-verrucose, corticate thallus-mantle. Phyllocladia coralloid, terete, usually slightly 

verruculose-torulose; lower ones longer and harmonically developed, upper ones short and 

divaricate; uppermost parts of pseudopodetia destitute of phyllocladia (habitus notably 

"macrocarpoid"). Cephalodia becoming very deeply and closely scrobiculate-foveolate, 

much more so than in St. ramulosum, finally verruculose-scrobiculate. Apothecia usually 

ca. 1.5 mm diam., rarely up to 3.5 mm. Central cone densely gray-nubilated and opaque. 

Hypothecium colorless and transparent, contrasting sharply with the underlying central cone 

tissue. Spores 3-5 (-6)-septate, (25-) 4&55 (-65) x(3.c) 3.54.0 (4.5) p. The deeply and 

sharply wrinkled cephalodia and the transparent hypothecium are reliable distinguishing 

characters against St. ramulosum. 

The southern (Magellanic) population of the species is generally more stunted, fails to 

show the "macrocarpoid" habitus, has less deeply and closely scrobiculate cephalodia, and 

its spores are shorter and broader, ca. 25-28 (-32) x (4) 5 (-6) p, not however so short and 

broad as represented by TH. FRIES (1857a). All these features however intergrade, as specimens 

are taken from south to north, with those described above. The character of the 

transparent hypothecium remains constant throughout the entire range. 

The cephalodial cortex of St. implexum shows a structural transition from primary 

(nubilated, indistinctly pseudoparenchymatous) tissue to secondary (hyaline, gelatinizedpalisadic) 

tissue. The secondary gelatinized tissue is developed from the outer side of the 

primary tissue. The youngest cephalodia have a cortex consisting entirely of primary tissue. 

A similar superimposition of cortical tissues, but in reverse order, is seen in the cephalodia 

of St. staufferi LAMB. See discussion in LAMB, 1951, p. 537, and 1976, p. 358. 

The type material of St. proximum var. compactius ZAHLBR. from Juan Fernandez is shorter 

and more compacted, otherwise typical St. implexum. 

St. implexum is the commonest Stereocaulon species in the Great Lakes region of Argentine 

Patagonia (Rio Negro, Chubut), forming extensive pure associations on exposed rocks.

Journ. Hattori Bot. Lab. No. 43 1977 

106. Stereocaulon melanopotamicum LAMB (n. nom.) 

Syn. Stereocaulon corticatulum var. procerum LAMB, Farlowia, 4: 462 (1955). 

Typus: Argentina, Prov. Rio Negro, between Lago Frias and the Frias Glacier, on rock, leg. 

I. M. LAMB, 1950 (no. 6086) (CAN, holotypus; FH, isotypus). 

Icon. : LAMB 1955, fig. 27, as St. corticatulum var. procerum. 

Mat. chim.: atranorin, lobaric acid, and traces of unidentified substances. Usnic acid 

also demonstrated in one specimen, but perhaps due to a contaminant. The report by 

HUNECK and FOLLMANN (1965) of atranorin, lecanoric acid and parietin in a Chilean specimen 

was based on a misidentification, the plant in question being Leprocaulon tenellum (TUCK.) 

NYL.; see LAMB and WARD, 1974, p. 540. 

Distrib.: Argentina (Neuquen, Rio Negro, Chubut) and Chile (Malleco). 

Remarks: morphologically similar to St. corticatulum NYL., but sometimes considerably 

larger (up to 5 cm tall, as in the type specimen), although usually only ca. 2 cm, so that for 

certain identification chromatography is necessary to demonstrate the presence of lobaric 

acid instead of rangiformic and norrangiformic acids, the characteristic constituents of St. 

corticatulum. The apical soredia, conspicuous in the type material, are not always distinctly 

developed. May occur closely associated with St. corticatulum in Argentine Patagonia. 

107. Stereocaulon meyeri B. STEIN 

Jahresber. Schles. Ges. Vaterl. Cult. 66: 134 (1888); LAMB ex FREY, Bot. Jahrb. Syst. 86: 220 (1967). 

Typus: Africa, Tanzania, Mt. Kilimanjaro, altit. 4000 m s. m., leg. H. MEYER, 1887. Holotypus 

(BRSL ?) not seen by us, but an isotype sent by B. STEIN studied in UPS-TH FR. 

Facultative synonymsT3: Stereocaulon meyeri var. bornmulleri B. STEIN, Jahresber. Schles. Ges. 

Vaterl. Cult. 66: 134 (1888). - Stereocaulon ramulosum var. bornmulleri (B. STEIN) M ~L. ARG. Engler's 

Bot. Jahrb. Syst. 15: 513 (1893). - Stereocaulon haumanianum DUVIGN. Lejeunia, Mbm. no. 14: 102 

(1956). - Stereocaulon karisimbiense DUVIGN. IOC. cit. p. 94 (1956). - StereocaulonpenicilliurnD~~~~ 

loc. cit. p. 98 (1956). 

Exsicc.: ABB. Lich. Madagasc. et Borbon. Sel. Exs. Fasc. I1 (1966) no. 39 (FH). 

Icon.: DUVIGN. 1956, figs. 2, 27, as St. penicillium; fig. 26, as St. karisimbiense; figs. 28, 29, 30, as 

St. haumanianum. 

Mat. chim.: atranorin, perlatolic acid, and 2 unidentified substances. The chemistry 

therefore agrees with that of the closely allied species St. ramulosum (Sw.) RAUSCH. 

Distrib. : central Africa (Kenya, Rwanda-Burundi, Uganda, Tanzania), South Africa, Madagascar, 

Rkunion, Tristan da Cunha, Mexico, Venezuela, Colombia, Peru, Bolivia, Argentina (prov. C6rdoba). 

Remarks: whether St. meyeri should be maintained as a distinct species separate from 

St. ramulosum (Sw.) RXusc~. is perhaps open to question. The only reliable distinguishing 

character is the presence of soralia on the apices of the pseudopodetia, and these are variable 

in their degree of development. Generally speaking, the abundance of soralia in St. meyeri 

is inversely proportional to the degree of fertility (presence of apothecia). In well fertile 

African specimens the soredia may be reduced to vanishing point, and the distinction from 

St. ramulosum becomes subjective. POELT (1970, 1972a) has drawn attention to the existence 

of paired ascocarpic and sorediate taxa in various lichen genera; St. ramulosum and St. meyeri 

afford a good example of such an "Artenpaar". See also discussion in BRODO and HAWKS- 

WORTH, 1977, pp. 3C32. Morphologically subsimilar but chemically different species related 

to St. meyeri are St. atlanticum (LAMB) LAMB and St. rugulosum LAMB. 

73 Stereocaulon proximum f. sorediata NYL. (syn. Stereocaulon sorediatum (NYL.) VAIN.) may also 

possibly be a synonym of Sr. meyeri, but is an untypified doubtful taxon (see Append. 2, p. 324).


Fo. acuminatum (MULL. ARG.) LAMB (n. comb.) 

Basionym: Stereocaulon ramulosum var. acuminatum MULL. ARC. Flora, 73: 335 (1890); DODGE, 

Ann. Cryptog. Exot. 2 (2): 138 (1929). 

Typus: Africa, Tanzania, Mt. Kilimanjaro, altit. 3000 m s. m., leg. VON HOHNEL, 1890 (no. 208) 

(G-MULL, holotypus). 

Remarks: pseudopodetia simple or only sparingly branched, acuminate-attenuate at the 

apices, which are farinose-sorediate. Habitus resembling that of St. cornutum MULL. ARG. 

Rare; 2 specimens seen, from Tanzania and Kenya. 

Fo. tucumanum LAMB (n. f.) 

Syn. Stereocaulon meyeri var. tucumanum LAMB ex GRASSI, Lilloa, 24: 363 (1950) (nomen nudum). 

Diagn. : A specie typica differt phyllocladiis distincte complanatis et flabellato-divisis. 

Typus: Argentina, Prov. Tucumhn, Dept. Chicligasta, between La Cascada and Las Cuevas, altit. 

2800 m s. m., on rock in subalpine pasture at the tree limit, leg. T. MEYER, 1949 (LL, holotypus; CAN, 

isotypus). 

Remarks: phyllocladia flabellately branched, distinctly flattened at the ends, often paler 

(whitish) on the underside. Apices of pseudopodetia sorediate. (Sterile.) Pycnoconidia 

filiform, arcuate, 8-10 xca. 0.5 p. Known only from the type specimen. 

Var. farinosum (TH. FR.) LAMB 

Canad. J. Bot. 29: 582 (1951). - Stereocaulon ramulosum [var.] 6. farinosum TH. FR. De Stereoc. et 

Pilophor. Comment.: 12 (1857); DODGE, Ann. Cryptog. Exot. 2 (2): 138 (1929). - Stereocaulon ramulosum 

f. farinosum (TH. FR.) HUE, Nouv. Arch. Mus. Hist. Nat. ser. 3,lO: 244 (sep. 32) (1898). - Stereocaulon 

farinosum (TH. FR.) VAIN. Dansk Bot. Ark. 4 (11): 6 (1926). 

Typus: Mexico, Pico de Orizaba, altit. 10-12000 feet, leg. LIEBMANN, 1841 (no. 74; PI. Mex. no. 

7663) (WS-TH FR, holotypus; c, FH, isotypes). 

Icon. : TH. FR. 1858, PI. VII, fig. 1, as St. ramulosum 6. farinosum. - LAMB 1951, PI. 111, fig. B. 

Remarks: distinguished from the typical species by its compact, prostrate-caespitose, 

dorsiventral habitus. Specimens seen from Mexico, Peru, Bolivia, Chile and South Africa. 

The description given by HUE (I 898-1901), based on a Colombian specimen (not seen by us) 

may refer to St. strictum TH. FR. The specimens from the Azores cited as St. ramulosum 

var. farinosum by DEGELIUS (1941) belong to St. atlanticum (LAMB) LAMB. 

108. Stereocaulon microcarpum MULL. ARG. 

Flora, 62: 162 (1879); DODGE, Ann. Cryptog. Exot. 2 (2): 126 (1929). 

Typus: Brazil, Slo Paulo State, near Apiahy, on rock, leg. J. I. PUIGGARI, 1877 (no. 151) (G- 

MULL, holotypus). 

Facultative synonyms: Stereocalrlon mixtum var. tenellum MULL. ARG. Flora, 63: 260 (1880). - 

Stereocalrlon proximum status ferruginascens MCL. ARG. Hedwigia, 30: 220 (1891). - Stereocaulon 

implexum f. sorediosa VAIN. Etude sur la Classification Naturelle et la Morphologie des Lichens du 

Brhil, Pars prima : 68 (1 890). 

Exsicc.: Krypt. Exs. Vindob. Cent. VII (1901) no. 667, as St. proximum var. gracilius (c, FH, LD, 

UPS, US, WELC). -VAIN. Lich. Bras. Exs. (1891-92) no. 1188, as St. implexum f. sorediosa (FH); no. 

1361, as St. implexum (FH). 

Mat. chim.: the chemistry of this species is complex. The presumed full chemical spectrum 

(seen in 1 specimen from Peru) comprises atranorin, stictic acid, constictic acid, norstictic 

acid, perlatolic acid, anziaic acid, glomelliferic acid, bourgeanic acid, and fumarproto-


cetraric acid. The following deficient phases have been found: 

Deficient phase I, with atranorin, stictic acid, constictic acid, norstictic acid, perlatolic 

acid, anziaic acid, glomelliferic acid. Seen from Brazil, Peru, Costa Rica and Mexico. 

Deficient phase 11, with atranorin, stictic acid, constictic acid, norstictic acid, perlatolic 

acid, anziaic acid. The type specimen of St. mixtum var. tenellum MULL. ARG. belongs here. 

Deficient phase 111, with atranorin, stictic acid, constictic acid, norstictic acid, perlatolic 

acid. Seen from Brazil, Peru, Oceania (Tahiti) and U.S.A. (Tennessee). 

Deficient phase IV, with atranorin, stictic acid, constictic acid, norstictic acid. The 

type specimen of St. microcarpurn belongs here. One specimen seen also from Galipagos 

Islands. 

Deficient phase V, with atranorin, stictic acid, perlatolic acid, anziaic acid. One specimen 

seen, from Peru. 

Deficient phase VI, with atranorin and stictic acid. One specimen seen, from Peru. 

Stictic acid being the dominant primary constituent, specimens of St. microcarpurn 

normally react PDtorange-red. One specimen from Brazil was found to give the reaction 

K + yellow then red, PD +persistent strong yellow. Chromatography however showed it to 

belong to Deficient phase IV, with an unusually high concentration of norstictic acid. 

Distrib.: Brazil, Peru, Costa Rica, Guatemala, Mexico, Oceania (Tahiti), GalApagos Islands, 

U.S. A. (Tennessee). 

Remarks: St. microcarpum varies considerably in stature, with pseudopodetia often 

only 1-2, but sometimes up to 6 cm in length. Tall states are morphologically hardly to be 

distinguished from St. ramulosurn (Sw.) RAUSCH. In many specimens the apices of the pseudopodetia 

are whitened, mealy and decorticated, giving a pseudosorediose appearance, probably 

not due to damage by animal agencies, and not to be confused with the true apical soredia in 

the related species St. atlanticum (LAMB) LAMB and St. rugulosum LAMB. The cephalodia are 

of the normal ramulosum-type, loose-cored with gelatinized-palisadic cortex. Pycnoconidia 

filiform, curved, 8-1 1 x ca. 0.5 p. St. proximum status ferruginascens MULL. ARG. was 

based on a rusty-colored specimen growing on iron-bearing rocks. St. microcarpum is the 

only species of subgen. Hofostelidium known to occur in the continental United States. It is 

found at the top of Roan Mountain in Tennessee. 

109. Stereocaulon novogranatense LAMB (n. sp.) 

Diagn.: Pseudopodetia disco basali substrato affixa, erecta sed nonnihil dorsiventralia 

(phyllocladiis uno latere crebrioribus), ca. 6 cm alta, modice ramosa, rigida sed vix lignosa, 

sordide cinereo-albida, decorticata, superficie laevi aut submollescenti, haud tomentosa. 

Rami principales tereti, I .0-1.7 (-2.5) mm crassi. Phyllocladia cylindrico-coralloidea, divaricata, 

corticata, laevigata, subulata, primo simplicia, mox congeste ramosa, glaucoalbida, 

ca. 0.7-1.3 mm vel basin versus ad 3.5 mm longa et magis ramosa, 0.1-0.2 mm crassa. Cephalodia 

pseudopodetiis lateraliter insidentia, pulvinata, 1-2 rnrn lata, valde contortuplicatoscrobiculata 

et corrugata, rugis complanatis et cristatis (fere ut in St. pseudomassartiano 

LAMB), isabellino-glaucescentia, marginibus sinuatis et pallidioribus; sacculata, intus laxe 

contexts, cortice hyalino et gelatinoso, palisadico. Apothecia terminalia, latit. 1.5-2.0 

(-3.0) rnrn, haud hypophysata, primo subplana et margine pallidiori haud prominenti cincta, 

dein bene convexa et irnmarginata, disco obscure fusco. Conus centralis cum hypothecio 

dense fuscidulo-nubilatus (KHO decoloratus). Hymenium 100-120 p altum. Sporae 6 

8nae, in asco leviter spiraliter contortae, nonnihil sinuosae, 7-1 1 (-16)-septatae, longit. 70-95 p,


crassit. 3.54.0 p. Atranorinum, acidum fumarprotocetraricum, acidum protocetraricum, 

acidum perlatolicum et acidum anziaicum continet (K+flavofuscescens, PD+aurantiaco- vel 

miniato-rubescens). - Species peculiaris, affinitate nonnihil dubia, fere ad subsect. Holostelidium 

accedens. 

Typus: Colombia, Volcan Purac6 between Popayan and La Plata, altit. 3300 m s. m., in paramo 

formation, apparently on rock, leg. G. GUZMAN, 1964 (no. 5543 pr. max. p.) (FH, holotypus; MEXPOL, 

isotypus). 

Mat. chim. : atranorin, fumarprotocetraric acid, protocetraric acid (trace), perlatolic acid, 

anziaic acid, and traces of unidentified substances (possibly including glomelliferic acid). 

Distrib. : Colombia. 

Remarks: the cephalodia are distinctive, outwardly somewhat resembling those of St. 

pseudomassartianum LAMB, but sacculate (loose-cored). The chemistry is similar to that of 

St. pachycephalum VAIN., which differs in the form of the cephalodia, lower hymenium and 

less septate spores. Seems to occupy a position transitional to the subsect. Holostelidium. 

Known only from the type material. 

1 10. Stereocaulon pachycephalum VAIN. 

Dansk Bot. Ark. 4 (11): 7 (1926). 

Typus: Mexico, Oaxaca State, Sierra de Oajaca (Oaxaca), El Pelado, altit. 9-10000 feet, on rock, 

leg. LIEBMANN, 1842 (no. 71b, Plant. Mex. no. 7654) (TUR-VAIN, no. 3868, holotypus; c, UPS-TH FR, 

isotypes). 

Mat. chim.: the full chemical spectrum (not yet found) appears to comprise atranorin, 

fumarprotocetraric acid, lecanoric acid, perlatolic acid, sublimbatic acid and bourgeanic acid. 

Material of the following deficient phases has been found: 

Deficient phase I, with atranorin, fumarprotocetraric acid, lecanoric acid, sublimbatic 

acid (trace), and bourgeanic acid. One specimen from Peru. 

Deficient phase 11, with atranorin, fumarprotocetraric acid, perlatolic acid, and probably 

a trace of sublimbatic acid. The type material belongs to this phase, also one specimen 

from Peru. 

Distrib. : Mexico, Peru. 

Remarks: related to St. microcarpum MULL. ARG. Habitually similar to St. ramulosum 

(Sw.) RAUSCH., but mng in its chemistry, almost complete cortication of the pseudopodetia, 

predomiaantly harmonic phyllocladia, slightly higher hymenium (1W110 p) and 

longer spores (not described by VAINIO) which are 4-6-septate, 55-85 x3.54.0 (-4.5) p. 

Cephalodia (with Scytonema) sacculate, anatomically like those of St. ramulosum. The 

presence of lecanoric acid is unique in the genus. Rare; known only from 4 collections. 

1 1 1 . Stereocaulon philippinense Rjis. 

Arch. Soc. Zool. Bot. Fenn. "Vanamo", 3: 79 (1949). 

Typus: Philippines, Negros, Canlaon Volcano, "ad terram argillaceam et arenosam", leg. E. D. 

MERRILL, 1910 (no. 6874) (H-RAS, holotypus). 


Distrib. : Philippine Islands. 

Remarks: this species, known only from the type specimen, remains poorly understood 

even after restudy of the holotype. RASANEN compared it to Cladonia cariosa (AcH.) SPRENG., 

but habitually it more calls to mind a very robust state of Stereocaulon condensatum HOFFM. 

The primary thallus is well developed and persistent, consisting of crowded, upright, closely


branched stipes (undeveloped pseudopodetia) densely covered at their apices with a more or 

less continuous crust of dactylaeform-coralloid phyllocladia of minute size (ca. 0.5 mm long, 

not over 0.1 mm diam.). Pseudopodetia simple or very sparingly branched, 1.5-2.0 cm high, 

0.61.0 mm thick, erect, rigid, decorticate, glabrous, pallid ochraceous. Phyllocladia densely 

clothing pseudopodetia from base to apex, cinerascent-whitish, terete-coralloid, simple or 

rarely furcate, minute, similar to those on the primary thallus. Cephalodia mostly on primary 

thallus, a few also lateral on pseudopodetia; inconspicuous, concolorous with the phyllocladia 

or with a slight aeruginose tinge, irregularly subglobose to pulvinate-verrucose, 0.3-1.0 mm 

diam., the larger ones becoming divided into more or less discrete verruculae; often with a 

few dark spots on the surface due to the Cyanophyceous algae (Stigonema) showing through; 

sacculate, loose internally, with cortex 25-60 p thick, colorless and hyaline, highly gelatinized, 

with fistulose, branched and anastomosing lumina intricated in various directions (not pali- 

sadic). Apothecia terminal, convex-pileate, large, up to 4.5mm diam., dark brown to 

blackish-brown, immarginate. Central cone colorless, compact. Hypotheciurn faintly 

sordid yellowish in section (not truly pigmented), 45-70 p deep. Hymenium 65-78 p high, 

with asci 50-60 p long. Spores seen 4-6 in ascus, cylindric-fusiform, straight, 3-septate, 

30-40 x 3.54.0 p. The cephalodia resemble those of St. corticatulum NYL., to which the 

species may be related, but it requires further investigation. 

1 1 2. Stereocaulon pseudomassartianum LAMB 

ex FREY, Bot. Jahrb. Syst. 86: 246 (1967); J. Jap. Bot. 51 (12): 357 (1976). 

Typus: Philippines, Luzon, Mountain Province, between Banaue and Mt. Polis, altit. 1060-1212 m 

s. m., on soil, leg. A. W. HERRE, 1923 (FH, holotypus; UPS-MAGN, isotypus). 

Exsicc.: Lich. Exs. Univ. Colorado Mus. Fasc. VIII (1971) nos. 300, 301, 303 (COLO, FH). 

Icon.: FREY 1967, fig. 3: PI. 17, fig. 3. - LAMB 1976, fig. 4 (cephalodia). 

Mat. chim. : atranorin, perlatolic acid, sometimes also trace amounts of anziaic acid. 

Wrongly stated to contain lobaric acid by LAMB ex FREY (1967). 

Distrib. : Philippines, Celebes and New Guinea. Erroneous records: Java and Borneo (LAMB 

ex FREY, 1967), =St. halei LAMB. 

Remarks: habitually resembles St. massartianum HUE on account of the apothecia borne 

on lateral stalks, but immediately distinguishable by the peculiar plate-like or cristate cephalodia, 

which are solid-cored and have a dimorphic cortical layer (LAMB, 1976). 

1 13. Stereocaulon rubiginosum PERS. 

ex GAUDICHAUD in DE FREYCINET, Voyage autour du Monde . . . executB sur les Corvettes de S. M. 

1'Uranie et la Physicienne, pendant les annBes 1817, 1818,181 9 et 1820, Botanique: 212 (1826); MAGN. 

in MAGN. et ZAHLBR. Ark. Bot. 31A (6): 48 (1944). - Stereocaulon sanguineum DEL. ex GAUDICHAUD, 

loc. cit. (illegitimate obligate synonym). 

Typus: U.S.A., Hawaii ("Iles Sandwich"), exact locality not stated, on soil, leg. C. GAUDICHAUD 

(date ?) (no. 72) (L, holotypus, not seen by us; isotypes studied in PC and H-NYL, also a probable isotype 

in UPS-TH FR). 

Facultative synonyms: Stereocaulon ramulosum [subsp.]* roccelloides TH. FR. De Stereoc. et 

Pilophor. Comment.: 13 (1857); Monogr. Stereoc. et Pilophor.: 330 (sep. 26) (1858); MAGN. in MAGN. 

et ZAHLBR. Ark. Bot. 31A (6): 43 (1944). - Stereocaulon roccelloides (TH. FR.) LAMB, J. Jap. Bot. 51 

(12): 354, footnote (1976)r4 (Strain 11). 

-- 

r4 ZAHLBRUCKNER (192627) and DODGE (1929) cite the taxon as Stereocaulon roccelloides, but this 

was a lapsus, not an intentional new combination.


Exsicc. : VEZDA, Lich. Sel. Exs. Fasc. XVII (1965) no. 415, as St. flavireagens (LD) (typical strain); 

Fasc. XIX (1966) no. 466 (FH, KLEM, LD) (Strain 11). 

Mat. chim. : 2 chemical strains are here recognized, which may merit the status of separate 

species : 

Strain I (the typical or rubiginosum strain), with atranorin, norstictic acid, connorstictic 

acid (trace) and perlatolic acid (trace). No stictic acid present. 

Strain I1 (the roccelloides strain), with presumed full chemical spectrum (found in 1 

specimen) consisting of atranorin, stictic acid, constictic acid, norstictic acid, perlatolic acid, 

anziaic acid and bourgeanic acid. The following deficient phases have been found: 

Deficient phase I, with atranorin, stictic acid, norstictic acid, perlatolic acid, anziaic acid, 

bourgeanic acid. 

Deficient phase 11, with atranorin, stictic acid, perlatolic acid, anziaic acid, bourgeanic 

acid (V~ZDA, Lich. Sel. Exs. no. 466). 

Deficient phase 111, with atranorin, stictic acid, norstictic acid, bourgeanic acid. 

Deficient phase IV, with atranorin, stictic acid, norstictic acid. 

Distrib. : both strains are sympatric (endemic to the Hawaiian Islands). 

Remarks: St. rubiginosum, together with St. caespitosum RED. and St. didymicum LAMB, 

is characterized by the peculiar structure of the cephalodial cortex, which anatomically resembles 

that of the pseudopodetial cortex, consisting of more or less distinct and separate 

thick-walled hyphae in irregularly interwoven arrangement, sordid isabelline and cloudy in 

section (not colorless, hyaline and conglutinated-palisadic as in St. ramulosum (Sw.) RAUSCH.); 

see LAMB, 1976, p. 354. The structure is the same in both the chemical strains. The cepha: 

lodia are sacculate, cerebriform-convoluted, sometimes flattened and tongue-like in the 

juvenile condition, with Scytonema or Stigonema. The species is of low, compact, rigid, 

divaricately branched habitus, very firmly attached to rock or soil, and the pseudopodetia are 

for the most part corticate. The spores seem to be constantly 3-septate, (22-) 2545 p long. 

The pycnoconidia are filiform, curved, 9-12xca. 0.5 p. The crimson-reddish color of 

PERSOON'S type material is obviously due to the secondary decomposition of norstictic acid. 

1 14. Stereocaulon rugulosum LAMB (n. sp.) 

Diagn.: Pseudopodetia caespitosa, irregulariter ramosa, substrato disco basali arcte 

affixa, longit. 24 (-7) cm, teretia, sat robusta (diametro 1-2 mm), rigida, habitu macro- 

carpoideo (phyllocladiis in partibus superioribus haud vel vix evolutis), pro max. parte strato 

corticali (pallio thallino) fere continuo, scrobiculato-ruguloso vestita, apicibus sorediata, 

soraliis terminalibus, capitatis, 0.8-1.5 mm latis, albidis, granuloso-pulverulentis. Phyllo- 

cladia cylindrico-coralloidea, simplicia ramosave, vulgo 1.0-1.5 mm longa, 0.2-0.4mm 

crassa, cremeo-albida, basin pseudopodetiorum versus longiora (ad 8 mm) et ibi bene ramosa; 

superne vix ulla. Cephalodia lateraliter sessilia aut breviter pedicellata, subgloboso-pulvinata, 

1-3 (4) mm lata, glaucescentia aut subaeruginoso-cinerea, tandem confertirn contortuplicata 

vel scrobiculato-rugosa, sacculata, intus laxe contexta, cortice 25-70 ,u crasso, fuscopallescenti, 

haud vel parum nubilato, manifeste pseudo- aut prosoparenchymatico, cellulis aut rotundatis 

et ca. 6 p latis aut palisadico-elongatis, ca. 10 x 3 p. pot he cia terminalia, primo scutelli- 

formia, mox pileato-convexa et irnmarginata, 1.5-2.5 (-3.5)mm lata, fusconigrescentia. 

Conus centralis sublaxe contextus, dense isabellino-inspersus et opacus. Hypothecium tota 

crassitudine (60-170 p) fuscidulo-nubilatum (KHO decoloratum). Hymenium 75-1QO 

(-120) p alturn. Sporae 8nae vel pauciores, rectae, elongato-fusiformes, (3-) 5-7-septatae,


40-70 (-75) p longae, 3.54.5 (-5.0) p latae. Atranorinum et acidum fumarprotocetraricum 

geret, accessorie etiam acidum perlatolicum et acidum anziaicum (K fuscoflavescens, PD 

miniato-rubescens). - St. meyeri B. STEIN subsimile est sed minus, habitu macrocarpoideo, 

pseudopodetiis ruguloso-corticatis, et materiis chimicis differens; St. atlantico (LAMB) LAMB 

vere arctius cognatum. 

Typus: Africa, Uganda, Ruwenzori Mts., Scott Elliott Pass, altit. 4000 m s. m., on rock in scree 

slope, leg. E. ESTERHUYSEN, 1956 (no. 25233) (FH, holotypus; BOL, isotypus). 

Mat. chim. : the full chemical spectrum (seen in several specimens) comprises atranorin, 

fumarprotocetraric acid, perlatolic acid, anziaic acid, and an unidentified substance. Protocetraric 

acid may also possibly be present in traces. 

Deficient phase I, with atranorin, fumarprotocetraric acid, perlatolic acid, and an unidentified 

substance. The type material belongs here. 

Deficient phase 11, with atranorin, fumarprotocetraric acid, and an unidentified substance. 

Distrib. : Central Africa (Uganda and Kenya). 

Remarks: a sorediate species morphologically resembling St. meyeri B. STEIN, but usually 

smaller, of more or less distinctly "macrocarpoid" habitus, for the greater part corticate with 

a rugulose thallus-mantle, and chemically different. Very similar in appearance to St. atlanticum 

(LAMB) LAMB, which has a wider geographical range (overlapping in Central Africa) 

and also differs in the chemical constituents. 

1 1 5. Stereocaulon salazinum (BORY) FI~E 

Essai sur les Cryptogames des Ecorces Exotiques Officinales: xcvii (1824) ("salazianum"). -Lichen 

salazinus BORY, Voyage dans les Quatre Principales Iles des Mers d'Afrique, 3: 106, footnote (1804). 

- Chlorocaulum salazinum (BORY) CLEMENTS, The Genera of Fungi: 175 (1909). 

Typus: Rbunion (Bourbon) Island, "les rochers de la plaine des chicots. Plus beau aux Salazes", 

leg. BORY DE SAINT-VINCENT (date ?) (PC, lectotypus; lectotypified by RIDDLE ex DODGE, 1929, p. 140). 

Probable isotypes (not all with complete data) seen in L, PC and PC-HUE. 

Facultative synonyms: Stereocaulon assimile NYL. Ann. Sci. Nat., Bot. ser. 4, 11 : 251 (1859). - 

Stereocaulon mixtum [subsp.]* assimile (NYL.) NYL. Synopsis Methodica Lichenum, 1 (2): 239 (1860). 

-Stereocaulon ramulosum var. assimile (NYL.) LAMB, Canad. J. Bot. 29: 582 (1951).75 

Exsicc.: ABB. Lich. Madagasc. et Borbon. Sel. Exs. Fasc. I1 (1966) no. 35, as St. ramulosum var. 

macrocarpum. 

Icon.'? BORY 1804, PI. XVI, fig. 3, as Lichen salazinus. - LAMB 1951, PI. 111, fig. 7 A, as St. ramu- 

Iosum var. assimile. 

Mat. chim.": the presumed full chemical spectrum (not yet seen in the same specimen) 

consists of atranorin, perlatolic acid, anziaic acid, stictic acid, norstictic acid, usnic and 

isousnic acids, and probably stenosporic acid. The material seen represents the following 

deficient phases : 

Deficient phase I, with atranorin, perlatolic acid, anziaic acid, stictic acid, usnic and 

isousnic acids. (Reunion.) 

'5 The epithet was emended by LAMB loc. cit. because NYLANDER'S isotype material in H-NYL has a 

slight intermixture of St. scutelligerum TH. FR.; but since NYLANDER'S description is perfectly unambiguous, 

this emendation seems unnecessary. 

Ffi~, 1824. PI. 111, fig. 7, as St. salazianum [sic] seems to represent a different species. 

'' The depsidone salazinic acid of ZOPF was isolated from "Stereocaulon salazinum" (actually St. 

scutelligerum TH. FR.). It is now known that this substance was in fact norstictic acid (DWIGNEAUD, 

1942; MORS, 1952), but in spite of this the name salazinic acid is still used for a different depsidone (see 

C. CULBERSON, 1969, p. 162).


Deficient phase 11, with atranorin, perlatolic acid, anziaic acid, usnic and isousnic acids. 

(Rkunion.) 

Deficient phase 111, with atranorin, perlatolic acid, stictic acid, norstictic acid. (Madagascar.) 

Deficient phase IV, with atranorin, perlatolic acid, anziaic acid and probably stenosporic 

acid. (Rkunion, including isotype material.) 

Usnic acid is a very unusual constituent in the genus Stereocaulon; it has been found also 

in one specimen of St. melanopotamicum LAMB, in which however it may be an accidental 

contaminant. But its occurrence in St. salazinum (2 specimens) seems to be indisputable. 

Distrib.: islands of the Indian Ocean (Rkunion, Madagascar). 

Remarks: the nomenclatural history of this species is very complicated. BORY'S original 

material consists of a mixture of 2 species, which were later designated as Stereocaulon assimile 

NYL. (1859) and Stereocaulon scutelligerum TH. FR. (1861). BORY'S original description combines 

the characters of both these species, but his figure, PI. XVI, fig. 3, shows a plant of the 

ramulosum-type. NYLANDER (1859, p. 250, and 1860, p. 233) applied the name St. salazinum 

to the St. scutelligerum component, and in this was followed by several other authors, e.g. 

HUE (1898-1901) and VAIN10 (1915). Other authors, such as ACHARIUS (1810), TH. FRIES 

(1861) and DODGE (1929) applied the epithet to the other component, which resembles St. 

ramulosum var. macrocarpum (RICH.) BAB." NYLANDER, considering the epithet salazinum to 

refer to the scutelligerum component, was automatically obliged to give a name to the ramulosum-like 

component, which he did as Stereocaulon assimile NYL. (1859, p. 251). In agreement 

with the views expressed by ACHARIUS and TH. FRIES (see above), RIDDLE (ex DODGE, 1929, 

p. 140) finally definitely lectotypified Lichen salazinus BORY on the ramulosum-like component 

(=St. assimile NYL.), thereby clearing up the longstanding confusion. The emendation of 

Lichen salazinus in the present sense can probably be credited to ACHARIUS (1810, p. 580). 

St. salazinum, in the presently accepted sense, is closely related to St. ramulosum (Sw.) 

RAUSCH., having a strong resemblance to the var. macrocarpum (RICH.) BAB. It is smaller 

and more slender than the latter, has a tendency in some specimens at least to produce harmonic 

branches (cfr. the illustration in LAMB, 1951, P1. 111, fig. 7 A), and has a distinctive chemical 

constitution as well as a different distribution pattern, confined to the Mascarene Islands and 

Madagascar. 

1 16. Stereocaulon staufleri LAMB 

ex FREY, Bot. Jahrb. Syst. 86: 247 (1967). 

Typus: New Guinea, North-east Region, Eastern Highlands Distr., Mt. Wilhelm, Goroka, Keglsugl, 

Pengage Creek, on route to Lake Aunde, altit. 3250-3500 m s. m., on granitic rock, leg. H. U. 

STAUFFER, 1964 (no. 5668-B) (FH, holotypus; FREY, isotypus). 

Exsicc.: Lich. EXS. Univ. Colorado Mus. Fasc. VIII (1971) no. 305 (COLO, FH). 

Icon. : FREY 1967, fig. 4: PI. 17, fig. 4. 

Mat. chim.: the typical species contains atranorin, perlatolic acid, anziaic acid (in trace 

" FFBE (1824, p. xcvii) made the combination Stereocaulon salazinum (BORY) FFBE (as "salazianum") 

but gave no description, and his illustration (PI. 111, fig. 7) is quite unrecognizable. Chlorocaulum 

salazinum (BORY) CLEMENTS was applied to the scutelligerum component, but this does not affect the nomenclature. 

TH. FRIES (1857, p. 25, and 1858, p. 43) mistakenly applied the name Stereocaulon vulcani 

(BORY) ACH. to the St. scutelligerum component, but after this mistake was pointed out by N~ANDER 

(1859) rectified it by introducing the nomen novum scutelligerum, reserving the name salazinum for the 

ramulosum-like component of BORY'S original material (TH. FRIES, 1861).


amounts) and an unidentified substance. The var. borneense has the same constituents with 

the addition of glomelliferic acid. The statement in the original description that St. staufleri 

contains lobaric acid was an error. 

Distrib.: New Guinea (including Irian and Papua); N. Borneo (var. borneznse). It seems to be 

common in the Bismarck Ranges of northeastern New Guinea. Var. borneense has been found to date 

only in the Kinabalu National Park of North Borneo (Sabah), where it also appears to be quite common. 

Remarks: a large and conspicuous species, outwardly resembling large states of St. ramulo- 

sum var. pilophoroides (TUCK.) LAMB. The anatomical structure of the cephalodial cortex is 

like that found in St. implexum TH. FR. in consisting of two superimposed tissues, but in St. 

staufleri the nubilated-pseudoparenchymatous tissue is developed outside of the hyaline- 

gelatinized layer. 

Var. bornehe LAMB (n. var.) 

Diagn.: Minus et tenuius quam in typo, ramulis phyllocladoideis basin versus magis 

evolutis, cephalodiis partim laevigatis; atranorinurn, acidum perlatolicum, acidum anziaicum 

et insuper acidum glomellifericum continens. 

Typus: North Borneo (Sabah), Kinabalu National Park, altit. ca. 3500 m s. m., at base of rock 

slabs in open forest on tourist trail, leg. M. E. HALE, 1964 (no. 28310) (US, holotypus; FH, isotypus). 

Remarks: smaller and more slender than the typical species, with more development of 

phyllocladioid branchlets towards the base. Cephalodia partly with smooth surface. Glomelliferic 

acid present in addition to atranorin, perlatolic acid and anziaic acid. Spores seen 

3-7-septate, 70-90x&5 p, sometimes slightly spirally wound inside ascus. 8 collections 

seen, all from the Kinabalu National Park. 

117. Stereocaulon stenospermum (LAMB) LAMB (n. comb.) 

Basionym: Stereocaulon argus var. stenospermum LAMB, Farlowia, 4: 460 (1955). 

Typus: Argentina, Prov. Rio Negro, near foot of Frias Glacier S. of Lago Frias, on exposed rock, 

leg. I. M. LAMB, 1950 (no. 6088) (CAN, holotypus). 

Icon.: LAMB 1955, fig. 26, as St. argus var. stenospermum. 

Mat. chim.: atranorin, perlatolic acid, anziaic acid, glomelliferic acid, and traces of 

unidentified substances. 

Distrib. : Argentine Patagonia (Rio Negro, NeuquCn, Chubut). 

Remarks: similar to St. argus HOOK. F. et TAYL. em. TH. FR., but differs in the narrower 

spores, partly or entirely transparent hypothecium, and presence of glomelliferic acid. 

1 18. Stereocaulon trachyphloeum LAMB 

in GALWWAY, LAMB et BRATT, Lichenologist, 8: 64 (1976). 

Typus: Tasmania, Hansen's Peak, altit. 1136 m s. m., on quartzite-schist rock, leg. G. C. BRAIT, 

1967 (no. 67-527) (FH, h010typuS; BRATT, ~SO~YPUS). 

Icon.: GALLOWAY, LAMB et BRATT 1976, P1. 2, figs. A, B. 

Mat. chim.: atranorin, fumarprotocetraric acid and protocetraric acid (in the thallus); 

perlatolic acid and anziaic acid (in the apothecia). 

Distrib. : Tasmania and New Zealand. 

Remarks: habitually resembles a coarse state of St. colensoi BAB. with roughened, areolateverruculose 

thallus-mantle. Of interest is the occurrence of different chemical constituents in 

the thallus and in the apothecia.


119. Stereocaulon wadei LAMB (n. sp.) 

Diagn.: Pseudopodetia caespitosa, erecta aut intricata, substrato arcte affixa, parva, 

altit. 5-14 mm, crassit. 0.3-0.9 rnm, teretia, irregulariter ramosa, passim corticata (pallio 

thallino laevigato aut rimoso), passim decorticata et glabra, pro maj. parte soredioso-pulverulenta. 

Thallus primarius effusus, granuloso-sorediosus. Phyllocladia haud distincte evoluta. 

Soralia apicalia, effusa, 1-3 (-5) rnrn lata, albida, granuloso-pulverulenta, ramulis pertenuibus 

corymbosis terminalibus laxe suffulta (ut in St. delisei BORY). Cephalodia subglobosa, sessilia 

aut brevissime pedicellata, 0.61.8 mm lata, plumbeo-cinerascentia, distincte confertimque 

scrobiculata aut botryoso-scrobiculata, sacculata, intus laxe contexta. (Apothecia haud visa.) 

Atranorinum, acidum perlatolicum, acidum glomellifericum et acidum glomellicum continet. 

- Ad stirpem St. corticatuli pertinet et cum St. delisei BORY similitudinem praebet, sed materiis 

chimicis ornnino diversum. 

Typus: New Zealand, North Island, Tongariro National Park, near Whakapapa, leg. A. E. WADE, 

1966 (FH, holotypus; BM, isotypus). 

Mat. chim. : atranorin, perlatolic acid, glomelliferic acid and glomellic acid. The latter 

substance occurs in brown Parmeliae (Melaenoparmelia). The same series of compounds 

(perlatolic acid, glomelliferic acid, glomellic acid) occurs very commonly in brown Parmeliae; 

see LEUCKERT, POELT, SCHULTZ and SCHWARZ, 1975. 

Distrib. : New Zealand. 

Remarks: known only from the type material. The morphological appearance falls 

within the range of St. corticatulum NYL., but the apical soralia are larger and more copiously 

developed than is usual in that species, and the cephalodia more closely and distinctly scrobiculate, 

resembling those of St. implexum TH. FR. The unique chemistry sets it apart from both 

St. corticatulum and St. delisei. 

Subsect. C. Dactyloideum (LAMB) LAMB 

120. Stereocaulon strictum TH. FR. 

De Stereoc. et Pilophor. Comment.: 24 (1857); Monogr. Stereoc. et Pilophor.: 346 (sep. 42) (1858). 

- Stereocaulon substrictum HUE, NOUV. Arch. Mus. Hist. Nat. ser. 3, 10: 253 (1898) (ut nom. nov.). 

- Stereocaulonpeladense VAIN. Dansk Bot. Ark. 4 (1 1): 7 (1926) (ut nom. nov.). 

Typus: TH. FRIES cited 3 Mexican collections made by LIEBMANN. AS lectotype we select the specimen 

from Oaxaca, Sierra de Oajaca (Oaxaca), altit. 9-10000 feet, leg. LIEBMANN, 1842 (no. 76, Plant. 

Mex. no. 7661) (UPS-TH FR, lectotypus, lectotypif. nov.; c, isolectotypus). 

Facultative synonyms: Stereocaulon lecanoreum NYL. Flora, 41: 117, footnote (1858); Synopsis 

Methodica Lichenum, 1 (2) : 233 (1 860) ; DODGE, Ann. Cryptog. Exot. 2 (2) : 124 (1929). - Stereocaulon 

strictum var. lecanoreum (NYL.) LAMB, Canad. J. Bot. 29: 582 (1951). - Stereocaulon mixtum sorediatum 

NYL. Synopsis Methodica Lichenum, 1 (2): 239 (1860). -Stereocaulon mixtum var. sorediatum 

(NYL.) HUE, NOUV. Arch. Mus. Hist. Nat. ser. 3, 2: 246 (1890). 

Exsicc.: V~ZDA, Lich. Sel. Exs. Fasc. XLVIII (1973) no. 1190 (BRNU). 

Icon.: TH. FRIES 1858, PI. VIII, fig. 4. - LAMB 1951, fig. 3 (cephalodia); 1976, fig. 2 (cephalodia). 

Mat. chim.: atranorin and porphyrilic acid. Cfr. HUNECK, 1974. 

Distrib. : Mexico, Costa Rica, Venezuela, Colombia, Ecuador, Peru, Bolivia. 

Remarks: St. strictum is characterized by the frequent presence of spathulate unifacial 

soredia at the apices of the pseudopodetia or at the tips of the phyllocladia, dactylaeform 

cephalodia, dark pigmented hypothecium, vermiform multiseptate spores, and the presence of 

porphyrilic acid. In fertile specimens soredia may be almost or entirely absent; in the var. 

compressurn they are very well developed. The cephalodia are solid internally and bounded


by a uniform, transparent, gelatinized cortical layer (LAMB, 1976). The apothecia are often 

pseudolecanorine with pale pseudothalline margin, as in the type material of St. lecanoreurn 

NYL. ZAHLBRUCKNER (1926-27), following NYLANDER (1860), included St. Iecanoreurn in 

subgen. Lecanocaulon. Although occasional symbiotic algae (Trebouxia) may be present 

entangled among the loosened hyphae on the outer side of the excipulum, the structure of the 

apothecium in St. stricturn, as in all other Stereocaulon species, is not truly lecanorine. St. 

substricturn HUE (1898) and St. peladense VAIN. (1926) are illegitimate nomina nova for St. 

stricturn, unnecessarily given on account of Stereocaulon rarnulosurn var. stricturn BAB. (1852, 

syn. St. foliolosurn var. stricturn (BAB.) LAMB). 

Var. compressum (NYL.) LAMB 

ex VBZDA, Lich. Sel. Exs. no. 1435 (1976). - Stereocaulon proximum var. compressum NYL. Synopsis 

Methodica Lichenum, 1 (2): 237 (1860). - Stereocaulon explanatum LAMB, Canad. J. Bot. 29: 582 

(1951) (ut nom. nov.). 

Typus: Colombia (Nova Granata), exact locality not stated, leg. GOUDOT, 1844 (PC, holotypus). 

Facultative synonym: Stereocaulon ramulosum f. compressum NYL. Synopsis Methodica Lichenum, 

1 (2): 236 (1860). 

Exsicc.: Lich. Exs. Sect. Bot. Mus. Hist.-Nut. Hungar. (1969) no. 86, as St. proximum (LD). - 

V~ZDA, Lich. Sel. Exs. Fasc. LVIII (1976) no. 1435 (BRNU). 

Icon.: LAMB 1951, fig. 5, as St. explanatum. 

Remarks: apices of pseudopodetia flattened, spathulate, corticated on one side, sorediate 

on the other. Apothecia rarely developed. Transitional states between the variety and the 

typical species are sometimes found. The nomen novum explanaturn LAMB was unnecessarily 

created on account of the mention of "Stereocaulon cornpressurn LINDS." in ZAHLBRUCKNER'S 

Catalogus, but that proved to be merely a lapsus, for LINDSAY'S reference is to Sphaerophoron 

cornpressurn. 

Sect. 2. Redingeria LAMB 

121. Stereocaulon caespitosum RED. 

Hedwigia, 76: 132,138 (1936); MARTIN et CHILD, Lichens of New Zealand: 155 (1972). - Gymnocaulon 

caespitosum (RED.) DWIGN. Lejeunia, Mtm. no. 14: 14 (1956) (comb. inval.). 

Typus: New Zealand, South Island, Otago, Mt. Maungatua near Dunedin, altit. 900m s. m., 

on schistose rock, leg. J. S. THOMSON (date ?); 2 collecting numbers cited, 185 and 485, of which we 

designate the former as lectotype (DSIR, lectotypus, lectotypif. nov.). 

Icon.: RED. 1936, figs. 1 A, 2 E, F. -LAMB 1951, figs. 3, 4. 

Mat. chirn.: the internal tissues give a strong red reaction with PD, and most specimens 

contain atranorin, protocetraric acid, furnarprotocetraric acid, and an unidentified pigment. 

Two specimens from Tasmania, however, although giving a similar reaction, were found to 

be of different chemical constitution: one contained atranorin and bourgeanic acidrs, the other 

apparently atranorin only. The chemistry of this species requires further investigation. 

Distrib.: New Zealand (South Island), Australia, Tasmania, Campbell Island. 

Remarks: a specimen from the type locality in New Zealand formed a patch about 1 m 

in diameter, estimated as being more than 400 years old (J. MURRAY, personal communication). 

The ontogenetic development of this species was studied by JAHNS (1970). 

70 Bourgeanic acid, an aliphatic compound, was first named, isolated and characterized (from species 

of Ramalina) by BODO, HEBRARD, MOLHO and MOLHO, 1973. It occurs also in several other species of 

Stereocaulon.


122. Stereocaulon gregarium RED. 

Hedwigia, 76: 139 (1936); MARTIN et CHILD, Lichens of New Zealand: 155 (1972). - Gymnocaulon 

gregarium (RED.) DUVIGN. Lejeunia, MBm. no. 14: 14 (1956) (comb. inval.). 

Typus: New Zealand, South Island, Otago, Flagstaff Hill near Dunedin, on schistose rock, leg. J. 

S. THOMSON (date ?) (no. 149) (DSIR, holotypus). 

Icon.: RED. 1936, fig. 3. - MARTIN and CHILD 1972, PI. 47, upper left. 

Mat. chim.: the full chemical spectrum consists of atranorin, protocetraric acid, fumarprotocetraric 

acid, perlatolic acid, colensoinic acid, anziaic acid, and possibly sublimbatic 

acid.80 

Deficient phase I, with atranorin, protocetraric acid and furnarprotocetraric acid. 

Deficient phase 11, with atranorin, protocetraric acid and perlatolic acid. 

Distrib. : New Zealand (North Island, South Island, Stewart Island). In North Island it appears 

to be rare. 

Remarks: very similar to St. caespitosum RED., but differing in the pale, wrinkled-scrobiculate 

cephalodia, the almost unpigmented hypothecium, the longer and narrower spores, and 

the filiform conidia. Ontogenetic development investigated by JAHNS (1970). 

123. Stereocaulon loricatum LAMB 

in GALLOWAY, LAMB et BRATT, Lichenologist, 8: 61 (1976). 

Typus: New Zealand, South Island, West Matukituki Valley, Mt. French, altit. 1600 m s. m., on 

rock, leg. D. J. GALLOWAY, 1967 (FH, h010typuS; BM, CHR, isotypes). 

Icon.: GALLOWAY, LAMB and BRATT 1976, PI. I, A, B. 

Mat. chim.: atranorin, colensoinic acid, fumarprotocetraric acid, salazinic acid (often in 

only trace amounts). 

Distrib. : New Zealand (South Island). 

Remarks: known from 7 collections. 

APPENDIX 1 

St. paschale (L.) HOFFM. 

St. paschale is the type species of the genus Stereocaulon, and the type specimen must be 

one of those seen by LINNB in drawing up his description of Lichen paschalis (1753) and 

preserved in his herbarium at the Linnean Society of London (LINN). VAINIO, 1886, cited 4 

sheets seen by him in the Linnean herbarium, which he referred, without comments, to Stereocaulon 

pascale [sic]. The situation is however somewhat more complicated. There are 

actually 6 sheets bearing material of "Lichen paschalis", of which 4 were labelled by other 

authors (LINN. fil., EHRHART) and were probably inserted after the date of publication and 

hence not to be considered as type material. The 2 sheets labelled by LINNB himself are nos. 

1273-254 and 1273-259. The former is labelled "Lichen pascalis globuliferus" by LINNB and 

~aschalis" by LINNAEUS fil., and bears one specimen of Stereocaulon ramulosum (Sw.) 

RAUSCH., as correctly determined by RIDDLE; the latter bears in LINNB'S handwriting "paschalis" 

and the numbers 982 and 68, and has 8 specimens, of which RIDDLE has stated, in an 

Sublimbatic acid is a substance of uncertain chemical affinity described by HALE (1967) from 

Parrnelia hypomelaena HALE and several other species of subgen. Xanthoparmelia. It has also been 

found in Stereocaulon pachycephalum VAIN.


attached annotation, "This is the authentic Type of Lichen paschalis Linn. The central 

specimen is certainly that species. The others are less typical." The number 982 refers to 

the number of the species on p. 359 of the first edition of LINNJYS Flora Suecica (1745), where 

the polynomial nomenclature ("Lichen fruticulosus solidus tectus, foliolis crustaceis") is used; 

68 refers to the number of the species in LINNE'S Species Plantarum, vol. 2 (1753), which is 

the original publication of Lichenpaschalis. We inspected the specimens on this sheet in 1954 

and again in 1958. The large central specimen which RIDDLE seems to suggest as lectotype 

is not St. paschale as now understood, but is either St. tomentosum FR. or St. myriocarpum 

TH. FR., with the numerous small lateral apothecia characteristic of both those species. The 

lectotypification of RIDDLE (if such was intended) should therefore be rejected. The remaining 

7 specimens, grouped around the large central specimen, are as follows (starting at the uppermost 

one and going clockwise): (2) St. paschale (fert.); (3) St. paschale (ster.); (4) St. 

paschale (fert.); (5) St. paschale (sparingly fert.); (6) apparently referable to St. grande (MAGN.) 

MAGN. (fert.); (7) St. paschale (sparingly fert.); (8) St. paschale (ster.). We would suggest 

as lectotype of St. paschales' specimen no. (5), at the bottom of the sheet, not only because 

it represents the species in the generally accepted sense, but also because LINN~ has written 

the Flora Suecica number 982 close against it, which suggests that this may be the specimen 

which he regarded as most characteristic of Lichenpaschalis. This specimen shows the typical 

dark cephalodia. The origin of the specimens on the sheet is not known, but the placing 

of the Flora Suecica number 982 against the lectotype would seem to indicate that it was from 

Sweden (probably Lapland). 

It must be remembered that prior to the end of the first quarter of the 19th. century, when 

St. dactylophyllum FLK. (1819), St. tomentosum FR. (1824) and St. alpinum LAUR. (1827) were 

first discriminated, the names Lichen paschalis and Stereocaulon paschale were used in a very 

wide sense, including these species, and probably some others also. 

St. paschale f. typicum DU RIETZ 

No type designated. A superfluous nomenclatural synonym for the typical species. 

St. paschale [var.] a. vulgare TH. FR. 

Although intended to designate the typical species, this variety is probably not to be considered 

as typified by the Linnean material, which according to TH. FRIES'S remarks (1857, 

p. 34) he had not seen, but should be typified on one of the specimens cited by him in his original 

description of that taxon, i. e. either EHRHART, PI. Cryptog. Linn. no. 118 or FLOTOW, 

Deutsche Lich. Ens. no. 12 A-C (both in herb. TH. FRIES). Through the courtesy of Dr. 

R. MOBERG (UPS) we received both these specimens for examination, and verified that they are 

both referable to St. paschale. The EHRHART specimen is somewhat atypical in having pale 

Nostoc-cephalodia of the alpinum-type, but FLOTOW 12A and 12B both have the typical dark 

Stigonema-cephalodia. No. 12A is fertile, and we designate it as the lectotype of var. vulgare 

(lectotypif. nov.), a synonym of the typical species. These FLOTOW specimens bear no indication 

of locality. 

St. paschale [var.] 6. serpens TH. FR. 

Typus: Sweden, "circa Upsaliam sat frequens" (UPS-TH FR). It is from Vitulfsberg, leg. TH. 

FRIES, 1852. 

Lectotypif. nov.


St. paschale f. colligatum LAMB 

Typus: Canada, Nova Scotia, Cape Breton, Victoria Co., Ingonish, Middle Head, leg. I. M. 

LAMB, 1952 (no. 6946) (CAN). 

Lichen crustaceus GILIB. 

No holotype designated; specimens mentioned from France and Lithuania (not seen). 

Lichen paschalis L. given as a synonym, and a description supplied which conforms to some 

species of Stereocaulon. Possibly intended as a new species based on one of the specimens 

mentioned, but more likely merely an illegitimate change of name for Lichen paschalis. 

St. tomentosum [var.] y. thyrsoideum SCHAER. 

Switzerland, various specimens mentioned, no holotype designated. The specimens 

in question (G) examined by FREY, who stated them (1932, p. 153) to be typical St. alpinum 

LAUR.; not seen by us. FREY, 1. c., p. 172, also mentions a specimen in herb. SCHAER., called 

St. tomentosum var. thyrsoideum, from Switzerland, Gauligletscher, which is St. glareosum 

(SAV.) MAGN., but as this locality is not listed in SCHAERER'S protologue, it is not part of the 

type material of var. thyrsoideum, and was probably added later by SCHAERER. 

St. tomentosum [subsp.]* alpinum [f.] a. verrucosum TH. FR. 

No type designated; specimens mentioned from Spitsbergen, Norway, Scotland, Ger- 

many, Switzerland, France and Italy. St. alpinum LAUR. given as synonym. A superfluous 

nomenclatural synonym for the typical form of that species. 

St. alpinum f. adpressum MAGN. 

No holotype designated; specimens mentioned from Italy and Switzerland. Dr. R. 

MOBERG (UPS) informed us that there are no specimens in ups (incl. UPS-MAGN) identified as 

St. alpinum f. adpressurn. Therefore a lectotype would have to be selected. It seems obvious 

from the description that the name refers to a mere growth-state or modification of St. alpinum. 

St. botryosum var. subincrustatum FREY 

3 specimens mentioned, 2 from Switzerland and 1 from Austria, no holotype designated. 

As lectotype we have chosen that from Switzerland, Berner Alpen, Faulhornkette, Schoniwang 

near Breitenboden, leg. E. FREY, 1928 (no. 294) (FREY) (lectotypif. nov.). It is a juvenile 

saxicolous state of St. alpinum LAUR. with short pseudopodetia still firmly attached to the rock. 

FREY, in litt. 1957, concurred with the opinion that this taxon is a synonym of St. alpinurn. 

The other Swiss specimen mentioned does indeed belong to St. botryosum ACH. em. FREY, 

but as its characters do not agree with those described in the protologue, we excluded it from 

consideration in selecting a lectotype. 

St. fastigiatum ANZI 

No holotype designated; "in molte alpi del distretto di Bormio, e della Valtellina". The 

authentic material distributed in ANZI, Lich. Rar. Langob. Exs. no. 16 should probably be 

designated as isotype or isolectotype subject to study of the material in ANZI'S own herbarium, 

for which we did not have the opportunity. 

St. botryosum subsp. islandicum DUVIGN. 

Typus: Iceland, Arnessjhla, Thingvellir, "ad lavam inter muscos", leg. P. VAN OYE,


1938 (no. 5) (formerly BRLU, but destroyed during World War 11, according to Prof. Du- 

VIGNEAUD (oral commun., 1958)). It was supposed to differ from the typical St. botryosum 

in its chemistry, but the identity of the substances present was not elucidated. To be dis- 

carded as a nomen dubium. 

St. fastigiaturn f. congesturn MAGN. 

3 specimens cited (2 from Sweden and 1 from Norway), no holotype designated. We 

select the one from Sweden, Tome Lappmark, Koppargsen. It is fertile, although MAGNUSSON 

stated it to be sterile. 

St. farinaceurn MAGN. 

Typus: Sweden, Tome Lappmark, par. Jukkasjami, Vassitjlkko, altit. ca. 600 m s. m., "on soil in 

alpine situations", leg. A. H. MAGNUSSON, 1919 (no. 3371) (UPS-MAGN, holotypus). 

Patellaria pileata [var.] a. crustacea WALLR. and [var.] ,B. thamnodes WALLR. 

Both described from Germany (Thiiringen), without exact localities, "ad terram arenosam 

sterilem". Not seen by us. 

St. acaulon NYL. 

Typus: France, Limoges (prov. Lemovicensi), Barsac, "supra terram jugorum aridorum", leg. E. 

LAMY, 1873 (H-NYL, no. 40022, holotypus). 

St. condensaturn [var.] a. minor EGELING 

No type material now present in KASSEL; probably destroyed during World War I1 (G. 

FOLLMANN, personal commun.). 

St. condyloideum ACH. 

Typus: Sweden, no exact locality stated, "ad terram glareosam" (H-ACH, holotypus). 

St. condensatum f. athallinum SANDST. ex ERICHS. 

3 localities mentioned in Germany, Schleswig-Holstein, leg. ERICHSEN, 1926 and 1936, 

no holotype designated. Not seen by us, and not lectotypified. 

St. condensatum f. septentrionale MAGN. 

3 localities in Sweden (Harjedalen, Lycksele Lappmark, Lule Lappmark) mentioned, 

no holotype designated. We select as lectotype (lectotypif. nov.) the specimen from Harjedalen, 

Linsall, leg. STENHOLM, 1917 (UPS-MAGN). Color, after 38 years in the herbarium, not 

noticeably different from that of other herbarium material of St. condensatum. 

St. coralloides FR. 

Type from Sweden, but no locality or date designated. As the original description 

accompanied the exsiccat FR. Lich. Suec. Exs. Fasc. IV (1825) no. 118, the latter may be 

considered isotype material. 

St. dactylophyllum [var.] ,B. major SOMM. 

Type from Swedeish Lapland, but no locality designated. Not seen by us. Referred to 

St. coralloides a. dactylophyllum by TH. FRIES, 1861. 

St. coralloides var. orientalis RAS. 

Typus: U.S.S.R., Karelia ladogensis, Lake Ladoga, Kurkijoki, Lapinlahti, "auf Uferfelsen am

offenen See" (H-RAs, holotypus). 


St. spissum NYL. ex HUE 

Based on Zw. Lich. Exs. no. 997, from Germany, Oldenburg, Edewecht near Zwischenahn, 

leg. SANDSTEDE (date?). Two examples of this number are present in H-NYL, both placed 

in the cover of St. spissum; one (herb. NYL. no. 39877) is sterile, PD-, and probably St. 

saxatile MAGN.; the other (herb. NYL. no. 39878) is fertile, PD+ orange-red, and is referable 

to St. dactylophyllum var. occidentale; we selected it as the lectotype of St. spissum (LAMB, 

1954, p. 272). 

St. spissum f. dactylinum FREY 

Described from Germany, Schleswig-Holstein, "Gebiet der grossen Endmoranen", coll. 

C. F. E. ERICHSEN. Several localities cited, no holotype designated. Two specimens of this 

type material seen by us in HBG; one of them is St. saxatile MAGN., the other, with FREY'S 

annotation "f. dactylinum", is St. dactylophyllum var. occidentale; we select the latter as lecto- 

type of St. spissum f. dactylinum (lectotypif. nov.). 

St. fastigiatum f. confluens MAGN. 

Typus: Sweden, Torne Lappmark, par. Jukkasjarvi, KopparAsen, altit. 600 m s. m., leg. A. H. 

MAGNUSSON, 1921 (no. 6073) (UPS-MAGN, holotypus). 

St. evolutum var typicum TH. FR. 

TH. FRIES quotes this as synonymous with St. evolutum GRAEWE, and so it can be considered 

as based on the latter and therefore an obligate synonym. 

St. evolutum f. planum MAGN. 

3 localities in Sweden mentioned, all in Bohuslan, no holotype designated. We select 

the specimen from par. Naverstad, Klagerod, leg. A. H. MAGNUSSON, 1917, as lectotype 

(UPS-MAGN, lectotypif. nov.). It was marked as type specimen by MAGNUSSON himself. 

St. lusitanicum MAGN. 

Typus: Portugal, Minho, Serra do Gerez, near Cabril, altit. 1000 m s. m., leg. C. TAVARES, 1944 

(no. 196) (UPS-MAGN, holotypus). 

St. patagonicum LAMB 

Typus: Argentina, Chubut, Lago Menendez, leg. I. M. LAMB, 1950 (no. 5790) (CAN, holotypus; 

FH, isotypus). 

St. colensoi var. reagens RAS. 

Typus: Chile, Magallanes, Finlandia Fjord, leg. ROIVAINEN, 1929 (H-RAS, holotypus). 

St. patagonicum f. subirregulare LAMB 

Typus: Argentina, Rio Negro, Lago Frias, Cerro Rigi, altit. ca. 1750 m s. m., leg. I. M. LAMB, 

1950 (no. 5789) (CAN, holotypu~; FH, isotypus). 

St. paschale f. ramulgerum NYL. 

Typus: U.S.S.R., Kola Peninsula, Murmansk region, Ponoi, leg. N. I. FELLMAN, 1863 (no. 47) 

(H-NYL, holotypus).


St. glareosum f. congestum MAGN. 

Typus: Sweden, Lycksele Lappmark, par. Tarna, Syterbacken, leg. A. H. MAGNUSSON, 1924 

(no. 8377) (UPS-MAGN, holotypus; FH, isotypus). 

Patellaria tomentosa ,B. erecta WALLR. 

WALLROTH, descr. orig., gives as synonym St. incrustatum FLORKE, quoting the latter's 

Deutsch. Lich. no. 77, which is probably isotype material of St. incrustaturn, and which may be 

considered as type material of WALLROTH'S epithet. 

St. abduanum ANZI 

Several localities in Italy mentioned: River Adda (Abdua) and its tributaries in the Valle 

Tellina plain (Fusine, Delebio, Chiavenna); no holotype designated. Lectotypified by DODGE 

(1929, p. 106) on the Adda specimen, which we have not seen. Paratype material from Valle 

Tellina and Valle Chiavenna (Clavenna) is distributed in ANZI'S exsiccat Lich. Rar. Langob. 

Exs. no. 14, as St. incrustaturn. 

St. incrustatum var. elatum OKSN. 

Typus: U.S.S.R., E. Siberia, Irkutsk Oblast, near Sljudianka, Chamar-Daban, leg. A. N. OKSNER, 

5. viii. 1927 (herb. OKSNER, holotype, not seen by us). We did however see a specimen in UPS-MAGN, 

labelled "Stereocaulon conf. incrustaturn", which appears to be isotype material, as the locality, collector 

and date on the label are the same. It is quite typical St. incrustaturn. OKSNER corresponded 

with MAGNUSSON. 

St. myriocarpum [var.] ,B. orizabae TH. FR 

Typus: Mexico, Veracruz State, Mt. Orizaba, altit. 12000 feet (3700 m s. m.,) leg. LIEBMANN, 1841 

(no. 72b) (UPS-TH FR, holotypus; c, isotypus). 

St. tomentosum [subsp.]*** St. myriocarpoides NYL. 

Typus: N. India, Himalayas, exact locality not stated, altit. 12000 feet (3700 m s. m.), leg. J. D. 

HOOKER and THOMSON (date ?) (Herb. Ind. Or. no. 2170) (PC, holotypus; L, LD, UPS-TH FR, isotypes). 

St. condensatum var. sorediatum HARM. 

Lectotypified by DODGE (1929, p. 11 1) on the material distributed in CLAUD. et HARM. Lich. Gall. 

praecip. Exs. no. 474, from France, Herault, La Salvetat, leg. MARC (seen in FH, isolectotypus). 

St. pileatum var. sorediiferum NYL. 

Typus: France, Vosges, Dept. Moselle, Bitche, leg. ?, date ?. There is no specimen in H-NYL. 

St. pileatum f. macrum MAGN. 

Typus: Sweden, Bohuslan, par. Ljung, Kolbengtsered, leg. A. H. MAGNUSSON, 1925 (no. 9513) 

(UPS-MAGN, hblotyp~s; FH, ~SO~YPUS). 

St. saxonicum BACHM. 

several 'collections from Germany and Czechoslovakia mentioned in BACHMANN'S 

2 

papers (1926, 1927) in which for the most part the taxon is designated as Stereocaulon tiro- 

liense or Stereocladiurn tiroliense; no holotype designated. One of the collections cited, 

from E. Germany, Sachsen, Giesing mountain near Altenberg, was distributed in Krypt. Exs. 

Vindob. no. 2275, as St. alpinum var. tyroliense on the original label, St. saxonicurn on the se- 

cond emended label. This could possibly be selected as isotype material, but inspection


of the copy used by BACHMANN would be necessary, because it is heterogeneous; in co~o, 

FH and us a depauperate meager state of St. pileaturn AcH., but in PR a mixture of St. pileatum 

and St. nanodes TUCK., and in s entirely St. nanodes. The present location of BACH- 

MANN'S herbarium is unknown to us. 

St. pileatum f. ramificans MAGN. 

Typus: Sweden, Vastergotland, Alingsls, leg. STENHOLM, 1896 (UPS-MAGN, holotypus). 

St. paschale var. evolutoides MAGN. 

Over 11 specimens from Sweden (Bohuslan, Vastergotland, Varmland and Medelpad) 

cited, no holotype designated. We have selected as lectotype a specimen from Varmland, 

Sillerud, Noretjarn, leg. A. H. MAGNUSSON, 1912 (UPS-MAGN, lectotypus, lectotypif. nov.). 

St. tomentosum var. graphiticola RAS. 

Typus: Finland, Savonia borealis, Kuopio, Laivo Island, "supra saxa schistosa graphitica juxta 

fodinam", leg. V. RASANEN and A. S. LEPPALA, 1946 (H-RAS, holotypus; isotypes distributed in Licheno- 

theca Fenn. no. 247; that in LD examined by us). 

St. paschale subsp. evolutoides f. laxatum LAMB 

Typus: Sweden, Varmland, Sillerud, Jarnsjon, leg. A. H. MAGNUSSON, 1912 (UPS-MAGN, holotypus) 

St. paschale f. taeniarum MAGN. 

2 specimens from Sweden, Bohuslan, cited, no holotype designated. As lectotype we 

selected that from par. Ockero, Bjorko, leg. A. H. MAGNUSSON, 1916 (UPS-MAGN, lectotypus, 

lectotypif. nov.). 

St. leporinum TH. FR. 

Several specimens cited from Madeira, leg. DIEDRICHSEN et al., and Canary Islands, 

leg. BOURGEAU et al., no holotype designated. As lectotype we have chosen the specimen 

from Madeira, Rio Frio, leg. F. DIEDRICHSEN (date?) (UPS-TH FR, lectotypus, lectotypif. nov.). 

It is quite typical St. azoreum (SCHAER.) NYL. (= "St. sphaerophoroides" mult. auctt., non 

TUCK.). 

St. granulosum LAUR. 

Specimens cited in HARTUNG (1857) from the Canary Islands (Lanzarote and Fuertaventura) 

and from Madeira; no holotype designated. HEPP, Flechten Europas no. 305 is 

quoted; this is from Madeira, Cural, leg. HEER and HARTUNG, and may be proposed as isolectotype 

material (lectotypif. nov.). Examples in FH-TUCK, PC and s studied by us. 

St. maderense TUCK. 

Typus: Madeira, Pico Ruivo, leg. PICKERING, U. S. Exploring Exped. 1838-42 (FH-TUCK, holotypus). 

It was subsequently annotated by TUCKERMAN himself "S. sphaerophoroides (S. Maderense 

perperam mihi olim)". 

St. leporinum [var.] a. elatum TH. FR. and [var.] p. pumilum TH. FR. 

Included in the several specimens cited under St. leporit~um (see above), holotypes not 

designated. BOURGEAU, Plant. ,Canar., iter secund. no. 1594, collected in the Canary Islands, 

Teneriffe, Selva de Las Mercedes, by E. BOURGEAU, 1855, as St. ramulosum, might eventually 

be considered appropriate as isolectotype material of var. elatum, and is typical St. azoreum


(SCHAER.) NYL. However, we failed to examine the material of these two varieties studied by 

TH. FRIES himself (UPS-TH FR). 

St. tomentosum [var.] ,!I. inciso-crenatum SCHAER. 

A general distribution in boreal and montane Europe indicated, localities being mentioned 

in Switzerland and France. FR. Lich. Suec. Exs. no. 90 cited, which is isotype material of 

St. tomentosum. Obviously intended by SCHAERER to indicate the typical form of the species, 

so that the above-mentioned exsiccat may be considered as the type material also of P. incisocrenatum. 

St. tomentosum [var.] a. campestre KORB. 

K~RBER gives no description or localities for this variety, but cites FR. Lich. Suec. Exs. 

no. 90, as well as several other exsiccata. The typification is therefore the same as that of var. 

inciso-crenatum above. 

Patellaria tomentosa [var.] y. decumbens WALLR. 

Mentioned as from Thiiringen and the Harz Mountains; Stereocaulon tomentosum FR. 

Sched. Critic. no. 90 given as a synonym, and DODGE (1929) lectotypified it on FR. Lich. Suec. 

Exs. no. 90. It is therefore an illegitimate obligate synonym of St. tomentosum. 

St. tomentosum [subsp.]*. magellanicum TH. FR. 

Typus: Chile, Magallanes, Magellan Strait; specimens collected by ANDERSON and from near 

"Sandy Point" (Punta Arenas) by LECHLER (Plant. Magell. no. 997); no holotype designated. As 

lectotype we select a specimen from "Fretum Magellanicum", leg. N. J. ANDERSSON, 1852, in UPS-TH 

FR (lectotypif: nov.); it was annotated by G. E. Du RIETZ as St. tomentosum. 

St. cupringorme NYL. 

Typus: Finland, Asikkala, leg. NORRLIN (date ?) (H-NYL, no. 39921, holotypus). A poorly developed 

condition of St. tomentosum, as stated by TH. FRIES (1871, p. 49). 

St. tomentosum f. truncatum FREY 

Typus: Germany, Mecklenburg, Rostock, Bernstorffer Tannen, leg. ? (B, holotypus, destroyed 

in World War I1 and not seen by us). From the description of it, and the illustration of a Tyrolean 

specimen, by FREY (1932, p. 185, PI. 111, fig. 8) it would seem to be merely a robust condition of the 

species. 

St. tomentosum f. tectorum TOM. 

Typus: U.S.S.R., Smolensk Oblast, Smolensk, "sur le toit pourri et moussu d'un hangar", leg. 

M. P. TOMIN, 1915 (no. 82) (VORAG, holotypus; UPS-MAGN, isotypus). Only the isotype seen by us. 

Stereocladium tyroliense NYL. 

Typus: Austria, Tirol, Brenner, leg. ARNOLD, 1874 (H-NYL, no. 40148, holotypus). 

St. tyroliense var. lapponicum MAGN. 

3 localities mentioned in Sweden, Lycksele Lappmark, par. Tba, Bjorkfors, by Syterbacken 

and Kvarnbacken, leg. A. H. MAGNUSSON, 1924. As lectotype we have selected 

the specimen from Syterbacken, altit. 475 m s. m., by a brook (MAGN. no. 8238) (UPS-MAGN, 

lectotypus, lectotypif. nov.).


St. hypopetraeum VAIN. 

Typus: U.S.S.R., Karelia ladogensis, Lake Ladoga (Laatokka), Kotiluoto Island, leg. E. A. 

VAINIO, 1923 (TUR-VAIN, no. 3764, holotypus; annotated "ex orig." by VAINIO). 

St. ostrobottniae MAGN. 

Typus: Finland, Ostrobottnia media, Karleby, Vittick, on granitic rock of a wall, leg. S. AHLNER, 

1938 (UPS-MAGN, holotypus). 

St. denudatum FLORKE 

Typus: cited from Germany, Harz Mts.: Brocken and Konigsberg; also from Norway (unlocalked); 

no holotype designated. As isolectotype material (lectotypif. nov.) we select FLORKE, Deutsch. 

Lich. no. 79 (1819), issued in conjunction with the original description. No locality is cited on the label. 

Seen in PR and UPS. 

Patellaria paschalis [var.] /I. bracteata WALLR. 

Listed from several localities in Germany, of which the only one mentioned by name is 

"Bructero Hercyn.", referring to the BroAen in the Harz Mts. of central Germany. The type 

material seen by us in STR is unlocalized, and VAINIO (1886a) also does not mention any locality 

for the type material seen by him in B. Probably a neotype from the same region should be 

designated. 

St. denudatum [status] a. tenue LAUR. ex FR., [status] b. 

validum LAUR. ex FR. and [status] c. digitatum LAUR. ex FR. 

No localized type material of these taxa cited. Pre-Linnean illustrations of b. validum and c. 

digitatum (DILLENIUS, MORISON) are mentioned, and these might possibly be considered as the types 

(STAFLEU et al., Int. Code, 1972, Art. 9). 

St. denudatum [f.] b. capitatum FLOT. 

Typus: Polish-Czechoslovakian border, Riesengebirge, no exact locality cited. B, holotypus, 

destroyed in World War I1 and not seen by us, but studied by FREY (1932, p. 133). 

St. denudatum [f.] P. commune TH. FR. 

No type locality cited; obviously intended to designate the typical condition of St. denudatum, 

and may therefore be considered as based on the same type material. 

St. denudatum [var.] a. genuinum TH. FR. 

No type locality cited; ff. validum LAUR. and commune TH. FR, cited as synonyms; therefore obviously 

intended to designate the typical condition of St. denudatum. 

St. nabewariense ZAHLBR. 

Typus: Japan, Honshu, prov. Kotsuke (=Gumma Pref.); 2 localities mentioned, Mt. Akagi and 

Mt. Nabewari, leg. A. YASUDA. We lectotypify it on the specimen from Mt. Nabewari, leg. YASUDA, 

1911 (no. 16) (w, lectotypus, lectotypif. nov.). 

St. denudatum [f.] d. compactum FLOT. 

Typus: Polish-Czechoslovakian border, Riesengebirge, Schneekoppe (Koppenkegel), leg. NEES 

v. ESENBECK and J. FLOTOW, 1836. B, holotypus, destroyed in World War I1 and not seen by us. 

FLOTOW'S description is vague, but it probably refers to f. umbonatum (WALLR.), the taxon with which 

it is placed by ZAHLBRUCKNER, 192627 (1927), p. 644.


St. confluens MULL. ARG. 

Typus: Java, Gedk Volcano, altit. ca. 7500 feet, leg. SOLMS, 1883 (G-MULL, holotypus). 

St. confluens var. fuscescens MULL. ARG. 

Typus: Africa, Tanzania, Mt. Kilimanjaro, altit. ca. 3000-4000 rn s. m., leg. L. v. HOHNEL, 1890 

(no. 204) (G-~LL, holotypus). 

St. violascens MULL. ARC. 

Typus: Ecuador, Mt. Chimborazo, Tambo de Totorillas, leg. E. ANDRB, 1876, altit. 3910 m s. m., 

(no. 3961 bis) and 4200 m s. m. (no. 3929). As lectotype we select the former specimen (G-M~~LL, 

lectotypus, lectotypif. nov.; isolectotypes in K, UPS, us). 

St. ruwenzoriense DUVIGN. 

Typus: Africa, Uganda, Ruwenzori, W. side, altit. 4200 m s. m., leg. HUMBERT, 1929 (BRLU, 

holotypus). 

St. microthuja DUVIGN. 

Typus: Africa, Uganda, Ruwenzori, Mt. Stanley, altit. 4250 m s. m., "roches nues horizontales", 

leg. HAUMAN, 1932 (no. 928) (BRLU, holotypus). The fac. abietina DUVIGN. (nom. inval.) is based on the 

same specimen. 

St. vulcani subsp. richardianum MONT. 

Typus: RCunion, "com. cl. Richard" (PC-MONT, holotypus; L, UPS-TH FR, isotypes). TH. FRIES 

(1858) erroneously reported the type as from Mauritius. 

St. speciosum LAMB 

Typus: Argentina, Chubut, Lago Menendez, below the Torrecillas Glacier, altit. ca. 650 m s. m., 

leg. I. M. LAMB, 1950 (no. 6069) (CAN, holotypus). 

St. virgatum f. achariana VAIN. 

Typus: Windward Islands, Dominica, Roseau Valley, leg. W. R. ELLIOT^, 1919 (no. 110) (TUR- 

VAIN, holotypus; could not be found by us, but a paratype specimen from St. Vincent, Souffrikre, leg. 

ELLIOTT thus inscribed was seen, herb. VAIN. no. 3840). VAINIO states that this taxon is the same as 

the type in herb. ACH. 

St. virgatum f. applanata VAIN. 

Typus: Windward Islands, St. Vincent, Richmond Valley, leg. W. R. ELLIOTT, 1919 (no. 193) 

(TUR-VAIN, holotypus, not seen by us). 

St. virgatum f. primaria VAIN. 

Typus: 2 localities cited in Windward Islands, leg. W. R. ELLIOTT, 1919: Dominica, Laudat (no. 

889) and St. Vincent, SouBritre crater (no. 143). As lectotype we select the specimen from Dominica 

(TURVAIN, no. 3847, lectotypus, lectotypif. nov.). 

St. giltayi DUVIGN. 

Typus: "Scottia", leg. L. GILTAY, 1937 (BRLU, holotypus). It is a perfectly typical specimen of 

St. virgatum, and must su~ely have come the West Indies. According to Dr. DWIGNEAUD (oral communication) 

the specimen was presented by L. GILTAY together with other specimens collected during 

a visit to Scotland in 1937, and he admitted the possibility of a confusion of labels.

I. MACKENZIE LAMB: A Conspectus of S~ereocaulon 

St. sphaerophoroides TUCK. 

TUCKERMAN'S protologue (1845) mentions no place of origin, but states that it was collected 

by L. GOULDING (ex herb. LAMBERT). He sent this type specimen to TH. FRIES, who wrongly 

identified it with his St. leporinum. This specimen, which appears to be the holotype of 

St. sphaerophoroidess2, is preserved in UPS-TH FR, labelled "In monte ignivomo Ind. Occid. ? 

Rev. L. GOULDING", and is typical St. virgatum ACH. See further discussion on pages 206,207. 

St. Jlavireagens GYELN. 

Typus: U.S.A., Hawaii, Glenwood ("Clenwood"), altit. 4000 feet, on lava, leg. E. SCHNELL (date 

?) (holotypus presumably in BP, not seen by us, but an isotype specimen seen in UPS-MAGN). 

St. Jlavireagens f. cinerascens MAGN. 

Typus: U.S.A., Hawaii, Maui, 2 localities mentioned, no holotype designated. As lectotype we 

select the specimen from Haleakala, along Halemau trail, leg. 0. SELLING, 1938 (no. 5886) (GB, lectotypus, 

leclolypif. nov. ; UPS-MAGN, ~SO~~C~O~YPUS). 

St. Jlavireagens f. densum MAGN. 

Typus: U.S.A., Hawaii, Hawaii, top of Hualalai, altit. 2600 m s. m., leg. C. SKOTTSBERG, 1922 (no. 

1569) (UPS-MAGN, holotypus). 

St. foliiforme HUE 

Typus: Japan, Hokkaido, Shiribeshi-yama, leg. FAURIE, 1905 (no. 6999) (PC, holotypus; FH, 

isotypus). 

St. subramulosum MULL. ARG. 

2 Japanese localities mentioned (Mt. Fusijama and Mt. Ontake), both collected by MI- 

YOSHI, 1890. As lectotype we have selected that from Mt. Fujiyama (no. 7) (G-MULL, lectotypus, 


St. subramulosum f. approximans HUE 

Typus: Japan, Honshu, Iidesan, leg. FAURIE (date ?) (no. 862) (PC-HUE, holotypus, but could not 

be found by us in 1966). 

St. subramulosum f. complanatum HUE 

Typus: Japan, Honshu, Yokosuka, leg. ONO and SAVATIER (date ?) (no. 546) (PC-HUE, holotypus, 

but could not be found by us in 1966). 

St. subramulosum var. humile MULL. ARG. 

Typus: Japan, Shikoku, Prov. Tosa (=Kochi Pref.), leg. MIYOSHI, 1890 (no. 54) (G-MULL, holotypus). 

St. uvuliferum MULL. ARG. 

Typus: Japan, Honshu, Prov. Tamba ("Tomba") (now part of Kyoto and Hyogo Prefectures), 

leg. MIYOSHI, 1890 (G-MULL, holotypus). Illustrated by ASAHINA, 1961b, fig. 1. 

St. armatulum ZAHLBR. 

Typus: Taiwan (Formosa), Soozan, Taihoku, on volcanic rock, leg. Y. ASAHINA, 1925 (no. 215) 

(w, holotypus). 

82 NO isotype duplicate could be found by us in FH-TUCK.


St. octomerelloides ASAH. 

Typus: Japan, Honshu, Prov. Etigo, Sasagamine Pasture, leg. Y. ASAHINA, 1949 (ASAH, no. 1521, 

holotypus; CAN, isotypus). 

St. sandwicense MAGN. 

3 specimens from the Hawaiian Islands (Maui and Hawaii) mentioned in the protologue, 

no holotype designated. As lectotype we have selected that from Maui, Haelauu, leg. 0. 

SELLING, 1938 (no. 5820) (s, lectotypus, lectotypif. nov.); it was marked "Typus" by MAGNUS- 

SON on the packet. 

St. octomerum f. robustior ASAH. 

Typus: Japan, Honshu, Prov. Ettyu, Mt. Tateyama, leg. Y. ASAHINA, 1928 (no. 612) (ASAH, 

holotypus). 

St. verruculigerum var. formosanum ASAH. 

Typus: "saxicola in the higher mountain range of central Formosa", no exact locality or collector 

designated. As lectotype we have selected a specimen from Taiwan (Formosa), Rengechi, leg. Y. 

ASAHINA, 1925 (no. F. 222) (ASAH, lectotypus, lectotypif. nov.; CAN, isolectotypus). 

St. exutum var. sorediata RAS. 

Typus: Japan, Kiushiu, prov. Hizen, leg. YASUDA, 1919 (no. 604) (H-RAS, holotypus). 

St. nesaeum var. lecideoides VAIN. 

4 specimens from the Philippines (Luzon and Mindanao) cited in the protologue, no holotype 

designated. As lectotype we selected the one from Mindanao, prov. Misamis, Mt. Malindang, 

leg. MEARNS and HUTCHINSON, 1906 (no. 4806) (TUR-VAIN, no. 3851, lectotypus, 


St. nesaeum var. zeorina VAIN. 

4 specimens from the Philippines (Luzon) cited in the protologue, no holotype designated. 

As lectotype we selected the one from prov. Benguet, Mt. Tonglon (Santo Tomas), leg. A. D. 

E. ELMER, 1904 (no. 6544) (TUR-VAIN, no. 3859, lectotypus, lectotypif. nov.). 

St. stalagmit$erum DWIGN. 

Typus: Java, Mt. Pangerango, altit. 2600 m s. m., leg. J. MASSART (date ?) (formerly BRLU, holotypus, 

but could not be located there in 1958. Dr. DWIGNEAUD informed us that most probably it was 

destroyed or lost during World War 11). 

St. nesaeum NYL., 1859, non auctt. plur. 

The only locality mentioned in the original description is "supra lapides et saxa in insulis 

Sandwich". No specimens thus identified from Hawaii could be found in H-NYL or PC, and 

the identity of the species is uncertain. It is certainly not the same as St. massartianum HUE, 

to which the name nesaeum was subsequently applied; see discussion on p. 273. 

St. sinense HUE 

Typus: China, Yiinnan, Lopin-chan above Lan-kong, leg. R. P. DELAVAY, 1888 (PC, holotypus). 

St. mamillosum DUVIGN. 

Typus: Africa, Uganda-Zaire border, Ruwenzori, Vallde de la Kiterere ("Ruterere" acc. to label),


altit. 4000 m s. m., on rock, leg. LEBRUN, 1931 (no. 4554) (BRLU, holotypus). 

St. claviceps var. yunnanense HUE 

Typus: China, Yiinnan, summit of Mt. Tsangchan, above Ta-li, altit. 4000 m s. m., on rock, leg. 

R. P. DELAVAY, 1884 (no. 664) (PC-HUE, holotypus). 

St. furcatum FR. 

Typus: West Indies, exact locality not stated, collector's name and date not given, "ded. Casstrom" 

(UPS-TH FR, ex herb. FR., holotypus). 

St. mixtum NYL. 

Typus: Bolivia, Cinti, on soil, leg. WEDDELL (date ?) (H-NYL, no. 40085, holotypus). 

St. proximup NYL. 

Typus: Colombia; 3 specimens cited, collected by LINDIG (date ?), nos. 385, 866, 1005. As 

lectotype we select no. 1005, from Piramo de Tolima, altit. 13000 feet (H-NYL, no. 40043, lectotypus, 


St. ramulosum var. compactum MULL. ARG. 

Typus: Australia, Queensland, on rock, leg. SHIRLEY, 1893 (no. 1738) (G-M~~LL, holotypus). 

St. ramuIosum [f.] y. strigosum TH. FR. 

2 collections cited: New Zealand, leg. A. RICHARD (date?) and southern Pacific Ocean 

region ("Soderhafs-landerna"), leg. SPARRMAN (date?). We have selected the latter as lectotype, 

since the RICHARD specimen could not be found in UPS; "Soderhafslandema" is the only 

localization on the label (UPS-TH FR, lectotypus, lectotypif. nov.). SPARRMAN was on Capt. 

COOK'S second voyage, which places the date of collection between 1772 and 1775. 

St. ramuIosum var. microcarpoides MULL. ARG. 

Place of collection unknown; collected by WALKER, 1875, on a journey through Fiji, 

New Zealand and Australia (G-M~~LL, holotypus). 

St. vimineum TH. FR. 

Typus: Mexico, Tiuzutlan, altit. 7000 feet, leg. F. LIEBMANN, 1841 (no. 73) (UPS-m FR, holotypus). 

St. proximum f. traversii HUE 

Typus: New Zealand, exact locality not stated, ''envoy6 par Sir Locke Travers a I'Exposition Universelle 

de Paris en 1889" (PC-HUE, holotypus). 

St. proximum [subsp.]** macrocarpoides NYL. 

Typus: 2 collections cited: Tasmania and Chile, without collectors' names or exact localizations. 

As lectotype we select the specimen from Tasmania, ex herb. HOOKER (H-NYL, no. 40029, lectotypus, 


St. denudatum PERS. (non FLORKE) 

Typus: Hawaiian Islands ("11. Sandwich"), leg. GAUDICHAUD, 1817-1820 (L-PERS, holotypus; 

PC, isotypus). It is typically developed St. rarnulosurn var. pilophoroides (TUCK.) LAMB. 

St. argodes NYL. 

Typus: Campbell Island, leg. FILHOL, 1874 (H-NYL, 

no. 40073, holotypus; MEL, UPS-TH FR, US, 

isotypes).


St. leptaleum NYL. 

Typus: Tasmania, no exact locality stated, on granitic rocks, leg. ?, ex herb. HOOK. F. (H-NYL, 

no. 39978, holotypus). 

St. corticatulum [subsp.]* detergens NYL. 

Typus: New Zealand, South Island, Otago, on rock, leg. ?, ex herb. JONES (H-NYL, no. 40025, 

holotypus). 

St. ramulosum var. compressum BAB. 

Typus: New Zealand, North Island, leg. COLENSO (date ?) (BM-BAB, holotypus; PC-MONT, isotypus), 

=St. fronduliferum LAMB. 

St. proximum var, compactius ZAHLBR. 

2 collections cited, both from Chile, Juan Fernandez, Masafuera, leg. C. and I. SKOT- 

TSBERG, 1917. As lectotype we select that from Las Torres, no. 350 (ups, lectotypus, lectotypif: 

nov.; SGO, isolectotypus). The other specimen, from near Correspondencia, no. 343, 

is identical (ups). 

St. meyeri var. bornmiilleri B. STEIN 

Typus: Africa, Tanzania, Mt. Kilimanjaro, altit. 4000 m s. m., leg. H. MEYER, 1887. Holotypus 

(BRSL ?) not seen by us, but an isotype sent by B. STEIN studied in UPS-TH FR. 

St. haumanianum DUVIGN. 

Typus: Africa, Uganda, Ruwenzori, Mt. Stanley, altit. 4200 m s. m., leg. HAUMAN, 1932 (no. 

931) (BRLU, holotypus). 

St. karisimbiense DUVIGN. 

Typus: Africa, Rwanda-Burundi, Karisimbi Volcano, altit. ca. 3600 m s. m., leg. HUMBERT, 

192983 (BRLU, holotypus). 

St. penicillium DUVIGN. 

Typus: Africa, Uganda, Ruwenzori, Mt. Stanley, Campi Achupa, altit. 4000 m s. m., leg. HAUMAN, 

1932 (no. 930) (BRLU, holotypus). 

St. mixtum var. tenellum MULL. ARG. 

Typus: Brazil, Slo Paulo State, near Apiahy, on rock, leg. J. I. PUIGGARI, 1879 (no. 151) (G-MOLL, 

holotypus). This specimen and the type of St. microcarpum MULL. ARC. were both collected near 

Apiahy by PUIGGARI, and both have the same collector's number (151). But according to the labels, 

that of St. microcarpum was collected in 1877, and that of St. mixtum var. tenellum in 1879. In spite of 

this apparent discrepancy in the dates, it seems possible that both may be portions of the same collection. 

St. proximum status ferruginascens MULL. ARG. 

Typus: Brazil, Minas GeraEs State, Ouro Preto, on weathered iron-mica schist, leg. H. SCHENCK, 

1891 (no. 4523) (G-MULL, holotypus). 

St. implexum f. sorediosa VAIN. 

Typus: Brazil, Minas Gerais State, Serra do Frio, on rock, leg. E. A. VAINIO, 1885 (TUR-VAIN, 

holotypus, not seen by us, but the isotype material distributed in VAIN. Lich. Bras. Exs. no. 1188 studied 

83 1931 acc. to the published description, but 1929 on the label.


St. ramulosum [subsp.]* roccelloides TH. FR. 

Typus: Hawaiian Islands, Oahu, near Honolulu, leg. N. J. ANDERSON (date ?) (UPS-TH FR, lectotypus; 

lectotypified by DODGE, 1929, p. 140). 

St. assimile NYL. 

Typus: RBunion (Bourbon) Island, Piton des Neiges, "a 1600 toises sur les rochers", leg. BORY 

DE SAINT-VINCENT (date ?) (PC, holotypus; H-NYL, no. 40077, isotypuss4). 

St. lecanoreum NYL. 

Typus: Colombia, near Antioquia, leg. D. JERVIS (date ?) (H-NYL, no. 40142, ex herb. HOOKER, 

holotypus). 

St. mixtum sorediatum NYL. 

Typus: Mexico, exact locality not stated, leg. GALEOTTI (date ?) (no. 6921) (H-NYL or PC ? Not 

seen by us). VAINIO, who apparently saw the type material, stated it to be synonymous with St. peladense 

VAIN. (=St. stricturn TH. FR.) (VAINIO, 1926). 

St. ramuIosum f. compressum NYL. 

Typus: "In America aequinoctiali", locality not stated, leg. BONPLAND (PC, not seen by us). 

NYLANDER'S description leaves no doubt as to its identity with St. stricturn var. cornpressurn (NYL.) 

LAMB. Not to be confused with St. ramulosum var. cornpressurn BAB. (=St. fionduliferurn LAMB). 

APPENDIX 2 

A. Doubtful taxa 

St. botryosum subsp. islandicum DUVIGN. 

Biol. Jaarb. "Dodonaea", 8: 160 (1941). 

Typus: Iceland, Thingvellir, on lava among mosses, leg. VAN OYE, 1938 (no. 5) (formerly in BRLU, 

destroyed in World War 11, acc. to P. DUVIGNEAUD, personal comrnun.). 

According to DUVIGNEAUD'S description, morphologically similar to St. botryosum ACH. 

em. FREY but differing in its chemical constituents as shown by the form of the crystals produced 

in GE solution after extraction with acetone. Since the substance present was not 

characterized chemically, the identity of the taxon remains dubious. 

St. confluens ff. typica, gracilius, ramosa, fastigiata DUVIGN. 

Lejeunia, MCm. no. 14: 84 (1956) (all nomina nuda). 

No material seen by us; from DUVIGNEAUD'S illustrations most of them would appear to 

refer to St. vesuvianum PERS. (var. vesuvianum). In any case they are to be ignored as nomina 

nuda. 

St. coralligerum MEYER 

Nebenstunden meiner Beschaefftigungen im Gebiete der Manzenkunde, Erster Theil: 156 (1825). 

Material mentioned from Chile (leg. CHAMISSO) and Brazil (leg. BEYRICH). Very incompletely 

described. There is a specimen in UPS-TH FR labelled "Stereocaulon coralligerum 

Called lectotype specimen in LAMB, 1951, p. 528.


Mey. Specimen origin. Loc. nat. non indic. Ex hb. Berol. accepi. Berol. Dec. 1861. Th. Fr."; 

it is St. montagneanum LAMB, and must have been collected in Indonesia. It cannot be regarded 

as type material, as only Chilean and Brazilian specimens are cited in the protologue. 

St. coralligerum must therefore be regarded as a doubtful species; see LAMB, 1965, p. 274. 

St. curtulum NYL. 

Flora, 59: 232 (1876). - Cereolus condensatus subvar. curtulus (NYL.) BOIST. Nouvelle Flore des Lichens 

2e. Partie: 33 (1903). 

Typus: France, Puy-de-DBme, Mont Dore, La Tache, leg. LAMY, 1875 (H-NYL, no. 40002, holo- 

typus). 

Very scanty and fragmentary material, not recognizable with certainty, as already stated 

by HARMAND, 1905-13 (1907), p. 362. 

St. denudatulum NYL. 

Flora, 57: 6 (1874). 

Typus: Finland, near Helsingfors, among mosses (Andraeaea), leg. NORRLIN, 1872 (H-NYL, no. 

39879, holotypus). 

Resembles St. vesuvianum var. nodulosum (WALLR.) LAMB, but smaller in all its parts. 

Phyllocladia PD+ red, many with an ill-defined darker central spot. It may possibly be 

synonymous with St. subdenudatum HAV., although the locality from which it was collected 

falls well outside of the known area of that species. The type material is very fragmentary, 

and the taxon must be regarded as a doubtful species. 

St. denudatum var. pulvinatum f. saxicolum SAV. 

Bot. Materialy, Notul. System. ex Inst. Cryptog. Hort. Bot. Petropol. 2 (11): 171 (1923). 

Described from U.S.S.R., Kamtchatka, "ad saxa in regionibus subalpinis alpinisque", 

no exact locality designated. 2 specimens are present in LE under this name, collected by 

V. P. SAVICZ: Truba Volcano, 1908, no. 2263 pr. p., and unlocalized, 1909, no. 6401. The 

latter is St. vesuvianum var. nodulosum f. depressum (MAGN.) LAMB; the former appears to be 

an undescribed species of the vesuvianum-arenarium group, PD- and with rather large terminal 

apothecia. Similar to this is another specimen from Krascheninnikov Crater, leg. 

SAVICZ, 1909 (no. 6582), labelled by him Stereocaulon denudatum and listed op. cit. p. 172. 

Unfortunately this interesting and probably new species could only be examined very briefly, 

and no data on its microscopic characters or chemical constitution were obtained. For the 

time being therefore it must be left as a doubtful and incompletely known taxon. 

St. gracillimum MULL. ARG. 

Flora, 64: 505 (1881). 

Typus: Japan, Honshu, Mt. Hakone, leg. BRAUNS, 1881 (no. 11) (G-MULL, holotypus). 

The somewhat scanty and depauperate type material is a species of the vesuvianum group 

rather resembling St. vulcani (BORY) ACH. in appearance, PD+ red, fertile, with apothecia 

typical morphologically and anatomically of the sect. Denudata. SAT^ (1941) wrongly considered 

it to be a species of Leprocaulon; see LAMB and WARD, 1974, p. 541. It cannot be 

identified with certainty, and must be considered as a doubtful species. 

St. kotiluotoense VAIN. 

ex RAs. Ann. Bot. Soc. Zool. Bot. Fenn. Vanamo, 12 (1): 55 (1939); Ann. Univ. Turkuensis, ser. A, 

7 (1) : 10 (1940) ("Kotiluotense"). 

Typus: U.S.S.R., Karelia ladogensis, specimens cited from 3 localities. We have selected


as lectotype one of those from Kotiluoto Island in Lake Ladoga, Serdobol' (Sortavala), on 

littoral rocks, leg. E. A. VAINIO, 1926 (TUR-VAIN, no. 34680, lectotypus, lectotypif. nov.). The 

syntypes seen (same locality, TUR-VAIN no. 34680A, and Lapinlahti, leg. V. RASANEN, 1934, 

in H) are the same. The lectotype consists of a number of small pieces. Thallus crustose, 

growing on detritus, mosses and other lichens (mainly Ephebe), forming small clumps up to 

2.5 cm diam., not effigurate, consisting of densely crowded, whitish nodules or verrucose 

squamules 0.34.8 (-1.0) rnrn diam. Most of these are sessile, but some are borne on congested, 

brownish, branching stipes penetrating into the substratum; not true pseudopodetia, 

but rather radicate basal extensions of the thalline nodules. No soredia or apothecia. Thallus 

K+ yellow, PD+ pale yellow; chromatography demonstrated atranorin, lobaric acid, 

and an unidentified pigment. The general habitus somewhat calls to mind an Ochrolechia 

species; VAINIO and RASANEN compared it in appearance with Lepraria neglecta (NYL.) 

ERICHS. and Lecidea ramulosa TH. FR. Whether it really belongs to Stereocaulon is doubtful. 

St. macquariense DODGE 

Nova Hedwigia, 15: 289 (1968). 

Typus: subantarctic islands of Australia, Macquarie Island, Wind Desert, "top of hills", leg. H. 

HAMILTON, B. A. N. Z. Antarct. Research Exped. 1929-31 (no. 108) (DODGE, holotypus, not seen by us). 

The description given is inconclusive, but obviously refers to some entity of the St. ramulosum 

complex. Material sent on loan at our request did not include the holotype; the specimens 

seen thus named from Macquarie Island were referable to St. ramulosum var. pulvinare 

(DODGE) LAMB. Probably the holotype is the same, but this cannot be definitely stated 

at present. 

St. meissnerianum FLORKE 

Deutsche Lichenen, Lief. IV: 14 (1819). 

No type designated, only the cephalodia described. In FH-TUCK, sheet 2393, there is 

an authentic specimen, unlocalized, labelled: "Stereocaulon Meisnerianum Floerke ex ejus 

herbario Herb. Reg. Berol.". It is the sterile thallus of St. condensatum HOFFM., also revised 

and annotated as such by TUCKERMAN. Therefore the epithet probably refers to St. condensatum, 

but is doubtful. 

St. nanum f. mundum FR. 

ex TH. FR. De Stereoc. et Pilophor. Comment.: 37 (1857). - St. quisquiliare f. mundum (FR.) ZAHLBR. 

Catalogus Lichenum Universalis, 4: 663 (1927). 

No type material could be found in ups; probably not referable to the genus Leprocaulon 

NYL. TO be discarded as a nomen dubium. 

St. paschale f. squamescens NYL. 

Lichenes Scandinaviae: 64 (1861). - St. paschale var. squamescens (NYL.) RAs. Lich. Fenn. Exs. no. 57 

(1935). - Stereocaulon squamescens (NYL.) RAS. Ann. Bot. Soc. Zool. Bot. Fenn. Vanamo, 12 (1): 53 

(1939). 

No type designated, merely a reference to this name given by NYLANDER in Mus. Fenn. 

Neither M~c~usso~.(1926, p. 51) nor the present author could find any specimens thus named 

by NYLANDER in H. TH. FRIES, who saw authentic material in Mus. Fenn., states that it may 

possibty be St. evolutum GRAEWE, but not determinable with certainty; one of the specimens, 

he states, was St. coralloides (=St. dactylophyllum FLORKE) (TH. FRIES, 1871, p. 46). It must 

therefore be regarded as a doubtful taxon. RASANEN used the name Stereocaulon squamescens


to designate St. evolutoides (MAGN.) FREY (=St. saxatile MAGN.). 

St. paschale [f.] c. subcrustosum FR. 

Lichenographia Europaea Reformata: 202, 203 (1831). 

No type designated. The description runs: "c. subcrustosum, rarnuloso-inaequabile, 

apotheciis maximis. . . primo obtutu P. subfuscam terrestrem refert". No material with 

this name could be found in ups. MAGNUSON (1926, p. 49) treated it as an "accidental form" 

of St. paschale, but the description of "apotheciis maximis" does not accord well with that 

species; St. rivulorum MAGN. would be more likely. A doubtful taxon. 

St. paschale f. velutinum FREY 


Typus: Poland or Czechoslovakia (former W. Schlesien), Heinersdorf, leg. FLOTOW 

(date?) (B, destroyed in World War I1 and not seen by us). From FREY'S description and 

illustration (PI. 111, fig. 3) it seems to agree with either St. alpinurn var. erectum FREY or St. 

grande (MAGN.) MAGN., but cannot be identified with certainty. 

St. proximum f. sorediata NYL. 

Synopsis Methodica Lichenum, 1 (2): 237 (1860). - Stereocaulon sorediatum (NYL.) VAIN. Dansk Bot. 

Ark. 4 (1 1): 7 (1926). 

No type specimen is cited in NYLANDER'S original description, and VAINIO mentions no 

original material. Being impossible to typify, the taxon is a doubtful one, but possibly refers 

to St. meyeri B. STEIN. 

St. rivulorum f. nana CENGIA SAMBO 

Nuovo Giorn. Bot. Ital. n. s. 40: 284 (1933). 

Typus: Italy, Piedmont, Passo Piccolo San Bernardo, leg. ?, date ?. 

No specimen bearing this name could be found in CENGIA SAMBO'S herbarium (FI), which 

contains only one specimen labelled "Stereocaulon rivulorum", from Passo San Bernardo, 

Lake Verney, and published as such by CENGIA SAMBO up. cit. p. 287; it is St. alpinum LAUR. 

This misidentification makes it seem very likely that the f. nana also does not belong to St. 

rivulorum MAGN., a species which we have not seen from Italy. 

St. solutum LAMB 

ex LYNGE, Skr. om Svalbard og Ishavet, no. 76: 50 (1939) (nomen nudum). 

This taxon, never validly published, was based on a specimen from Jan Mayen, Fishburn 

Valley, leg. R. S. RUSSELL, 1938 (no. 510 pr. p.) in BM. It is a sorediate species with some 

resemblance to St. capitellatum MAGN. and St. coniophyllum LAMB. Chromatography revealed 

only atranorin and a weak trace of some unidentified substance. Its identity is doubtful. 

St. spissum f. granuliferum FREY 

Rabenhorsts Krypt.-F1. 9, Abt. IV, 1. Halfte: 141 (1932). - St. dactylophyllum f. granuliferum (FREY) 

GRUMM. Catalogus Lichenum Germaniae: 24 (1963). 

FREY'S statement on typification is unclear, but it would seem that he refers to a specimen 

from Germany, Harz, Lauterburg, in herb. ZSCHACKE. This specimen (in B) was destroyed 

in World War 11, and was not seen by us; its identity is uncertain. 

Flora, 58: 358 (1875). 

St. subintricans NYL.


Typus: Finland, Hollola, Hayhto, on rock, leg. E. LANG (VAINIO), 1874 (H-NYL, no. 39967, holotypus). 

Obviously belongs to the paschale-saxatile group, but is peculiar in its copious fertility, 

with numerous minute apothecia, and its quite glabrous pseudopodetia. Phyllocladia PD-. 

We could not see any affinity with St. subcoralloides (NYL.) NYL., to which NYLANDER 

relates it. FREY also saw the type specimen, and gives some anatomical details (1932, p. 116). 

Although several specimens from Europe and N. America have been seen which correspond 

more or less to it, we cannot be sure of the identity, and prefer to consider St. subintricans as a 

doubtful species. 

St. tenuissimum DEL. 

ex NYL. Synopsis Methodica Lichenum, 1 (2): 242 (1860). 

Published by NYLANDER under St. coralloides FR., as "forma videtur sterilis transiens 

fere in Ster. Delisei. Cephalodia illius tenuissimi interdum cinereo-olivacea", from France, 

Dept. Calvados, St. Sever near Vire. The specimen not seen by us, and its identity doubtful. 

St. tomentosum f. jlabelliforme OHL. 

Schr. Konigl. ~hys.-bkon. Ges. Konigsberg, 11: 8 (1870). 

3 localities in former E. Prussia (now Poland) mentioned (Oletzo, Angerburg, Neustadt), 

no holotype designated. We have not seen any of the type material,86 nor apparently did 

FREY, who from the description equates it with typical St. tomentosum (1932, p. 184). 

OHLERT describes it as flabellately expanded with the phyllocladia forming a continuous crust 

on the upper side, and the pseudopodetia glabrous on the underside. The latter feature does 

not conform to St. tomentosum, and the f. flabelliforme, until the type material is found, must 

be regarded as a doubtful taxon. 

St. tomentosum var. hercynicum FREY 

Rabenhorsts Krypt.-Fl. 9, Abt. IV, 1. Halfte: 187 (1932). 

Typus: Germany, Harz, Lautenberg in Lutter Valley, leg. ZSCHACKE (date?), destroyed 

with ZSCHACKE'S herbarium (B) in World War 11, and not seen by us. From FREY'S description 

it seems to be probably synonymous with St. tomentosum var. compactum FREY. 

St. tomentosum f. leptothamnium FREY 

Rabenhorsts Krypt.-FI. 9, Abt. IV, 1. Halfte: 185 (1932). 

Typus: Poland (formerly Schlesien), Hartau, probably near Katzengebirge, leg. FLOTOW 

(date?), destroyed with FLOTOW'S herbarium (B) in World War 11, and not seen by us. From 

FREY'S description it appears to be a non-taxonomic modification of St. tomentosum, but 

its status remains doubtful. 

St. tomentosum f. walamoense NYL. 

Synopsis Methodica Lichenum, 1 (2): 244 (1860). -St. tomentosum var. walamoCnse (Nn.) Nn. 

Lichenes Scandinaviae: 65 (1861). - Stereocaulon valamoCnse [sic] (NYL.) RAS. Meddeland. Soc. Fauna 

Fl. Fenn. 50: 39 (1925). -St. alpinum f. walamoCnse (NYL.) FREY, Rabenhorsts Krypt.-FI. 9, Abt. IV, 

1. Halfte: 164 (1932). 

Typus: U.S.S.R., Karelia ladogensis, Valarno Island in Lake Ladoga, "supra terram nudam aridissimam", 

leg. W. NYLANDER, 1851 (H-NYL, no. 39952, holotypus; PC, isotypus). 

A very minute and poor specimen, resembling a much depauperated condition of St. 

s6 OHLERT'S herbarium is reputed to be at the University of Konigsberg (now Kaliningrad).


alpinum LAUR. Pseudopodetia distinctly tomentose. Phyllocladia minute, grain-like to 

lobate, PD-. Sterile. It gives the impression of a juvenile state of St. incrustaturn FLORKE, 

as remarked also by FREY loc. cit. MAGNUSSON (1926) could not decide on its real affinity. 

Must be considered as a doubtful taxon. 

St. tortuosum NYL. 

ex HULTING, Bih. Kongl. Svenska Vetensk.-Akad. Handl. 26, Afd. I11 (3): 17 (1900) (non DEL. in herb.). 

Typus: Sweden, Dalsland, Trollholmarne near Amal, leg. J. HULTING, 1870 (GB, holotypus). 

The above material somewhat resembles St. alpinum LAUR. and St. vesuvianum var. 

nodulosum (WALLR.) LAMB in habitus; phyllocladia PD+ pale yellow; sterile. A description 

of it is given by MAGNUSSON (1926, p. 83). Neither MAGNUSSON nor the present author, 

after study of it, could decide on its identity. NYLANDER, loc. cit., stated it to be synonymous 

with St. subintricans NYL.; we have studied the type of the latter, a doubtful species (see 

p. 324), and do not consider it to be the same. In the protologue the epithet tortuosum is 

ascribed to DELISE; the latter author never published any Stereocaulon of that name, but applied 

it to herbarium specimens of St. depreaultii DEL. from Newfoundland; numerous specimens 

thus named exist in various herbaria. The identification of St. depreaultii with 

HULTING'S Swedish material is incorrect. The epithet in the sense of DELISE (=St. depreaultii) 

has no taxonomic validity, since DELISE never published it. The first and only publication 

was by NYLANDER ex HULTING as above, and HULTING'S material must be considered the 

type of the epithet. As its identity cannot at present be satisfactorily established, St. tortuosum 

NYL. must be considered a doubtful species. 

St. turfosum BORY 

in DUMONT-D'URVILLE, Mkm. Soc. Linn. Paris, 4: 596 (1826). 

Typus: Falkland Islands, leg. DUMONT-D'URVILLE (date ?). Could not be found in PC. 

The very brief and inadequate description is insufficient for recognition. Cono~ (1915) 

suggested that it might be synonymous with St. tornentosum subsp. magellanicum TH. FR. 

(syn. St. tomentosum FR.), but he had seen no authentic material. To be regarded as a doubtful 

taxon, unless the type material is eventually found. 

B. Excluded taxa 

Baeomyces paschalis P. nanum (AcH.) WAHLENB. Flora Lapponica : 450 (18 12) = Stereocaulon 

nanum AcH., = Leprocaulon microscopicurn (VILL.) GAMS. ZAHLBRUCKNER, 192627 

(1927), p. 667, erroneously gives this taxon as a synonym of St. subcoralloides (NYL.) Nn. 

Stereocaulon aciculare (AcH.) TUCK. = Pilophorus acicularis (AcH.) NYL. 

St. aggregatum (SW.) RAUSCH. =Cladia aggregata (Sw.) ACH. 

St. albicans TH. FR.=Leprocaulon albicans (TH. FR. emend.) NYL. See LAMB and WARD, 

1974, p. 513. 

St. andinum B. DE LESD. Bull. Soc. Bot. France, 83: 7 (1936). Typus: Argentina, San Antonio 

de Los Cobres, leg. DUBUS, 1935 (B. DE LESD, destroyed in World War I1 and not seen 

by us). From the description, probably a stunted condition of Leprocaulon gracilescens 

(NYL.) LAMB et WARD, 1974, p. 523. 

St. arbuscula (NYL.) N~~.=Leprocaulon arbuscula (NYL.) NYL.; see LAMB and WARD, 1974, 

p. 516. 

St. arbuscula var. aberrans AsAH.=Leprocaulon arbuscula (NYL.) NYL., Ch. strain 11; see


LAMB and WARD, 1974, p. 516. 

St. buchanani KNIGHT =Bacidia buchanani (KNIGHT) HELLB. 

St. cereolinum ACH. =mainly Pilophorus cereolus (AcH.) TH. FR. This taxon was interpreted 

by some authors, e. g. TH. FRIES (1857, 1858) and NYLANDER (1860, 1861) as referring to 

St.pileatum AcH., but TH. FRIES subsequently showed (1871, p. 56) that the type material 

in herb. ACH. consists partly of Pilophorus cereolus (AcH.) TH. FR. and partly of Stereocaulon 

vulcani (BORY) ACH. 

St. cereolinum var. cereolus (AcH.) TH. FR. =Pilophorus cereolus (AcH.) TH. FR. 

St. cereolus ACH. = Pilophorus cereolus (AcH.) TH. FR. 

St. chlorellum TuC~.=Ramalina sp., perhaps R. pollinaria (AcH.) AcH., acc. to NYLANDER 

(1863, p. 11, footnote). 

St. cinchonae (WILLD.) RXUSCH. = Usnea cinchonae (WILLD.) ZAHLBR. 

St. confine (0. M~~LL.) HOFFM. =Lichina confinis (0. MULL.) AG. 

St. congestum NYL. =Leprocaulon congestum (NYL.) LAMB et WARD, 1974, p. 519. 

St. corallinoides HOFFM. = Parmeliella corallinoides (HOFFM.) ZAHLBR. 

St. corallinum (L.) SCHRAD. = Pertusaria corallina (L.) ARN. 

St. denudatum var. platycladum LAMB, Nytt Mag. Naturvidensk. 80: 278 (1940), cum icon. 

(PI. 11, figs. 1, 2). - Stereocaulon platycladum (LAMB) LAMB Canad. J. Bot. 29: 582 

(1951). Typus: E. Greenland, Kangerdlugssuak Glacier, altit. ca. 1300 m s. m., leg. 

L. R. and H. G. WAGER, 1935-36 (BM, holotypus). This is a compact, blackened, high 

alpine modification of Toninia conglomerata (AcH.) BOIST. resembling Stereocaulon 

vesuvianum PERS. to a remarkable degree and with the same chemical constitution and 

reactions. Only the longer, filiform conidia (16-24 x 0.5-0.7 p) supplied a clue to its 

correct identity. Quite similar specimens of Toninia conglomerata were subsequently 

seen from high altitudes in Switzerland (leg. FREY). See also LAMB, 1954a, p. 137. 

St. fibula TUCK. =Pilophorus cereolus (AcH.) TH. FR. 

St. fragile (L.) HOFFM. =Sphaerophorus fragilis (L.) PERS. 

St. fungiforme SCHAER. =Baeomyces rufus (HuDs.) DC. 

St. globiferum HOFFM. =Sphaerophorus globosus (Hum.) VAIN. 

St. gracilescens NYL. =Leprocaulon gracilescens (NYL.) LAMB et WARD, 1974, p. 523. 

St. intricatum MORIS = Lethariella intricata (MORIS) KROG, Norw. J. Bot. 23 : 94 (1976). 

St. laccatum FR. =Sphaerophorus globosus (Hms.) VAIN. 

St. laseroni DODGE, Repts. B. A. N. Z. Antarct. Research Exped. 1929-31, ser. B, 7: 141 

(1948) = Bacidia fibrosa LAMB, Rhodora, 56 : 128 (1954) (ut nom. nov.). 

St. madreporiforme HOFFM. =Pertusaria corallina (L.) ARN. 

St. melanocarpum (Sw.) RAUSCH. =Sphaerophorus melanocarpus (Sw.) DC. 

St. microscopicum (VILL.) FREY =Leprocaulon microscopicum (VILL.) GAMS ex D. HAWKSW.; 

see LAMB and WARD, 1974, p. 526. 

St. nanum (AcH.) Ac~.=Leprocaulon microscopicum (VILL.) GAMS ex D. HAWKSW. ; see 

LAMB and WARD, 1974, p. 526. 

St, nanum f. pulverulentum TH. FR. = Leprocaulon microscopicum (VILL.) GAMS ex D. HAWKSW. ;



St. novo-arbuscula ASAH. =Leprocaulon arbuscula (NYL.) NYL. (deficient phase); see LAMB 

and WARD, 1974, p. 533. 

St. obtusatum (VAHL) ACH. =Pertusaria oculata (DICKS.) TH. FR. 

St. paschale var. cereolinum (AcH.) SCHAER. =Pilophorus cereolus (AcH.) TH. FR. 

St. pseudoarbuscula ASAH. = Leprocaulon pseudoarbuscula (ASAH.) LAMB et WARD, 1974, p. 533. 

St. pulvinatum Ac~.=Siphula torulosa (THUNB.) NYL.; see MATHEY in Nova Hedwigia, 22 

(314) : 820 (1971). 

St. pygmaeum VAIN. = Lecania (Thamnolecania) brialmontii (VAIN.) ZAHLBR. ; see LAMB, 1948, 

p. 236. 

St. quisquiliare (LEERS) HOEEM. =Leprocaulon microscopicum (VILL.) GAMS ex D. HAWKSW. ; 


St. quisquiliare f. pulverulentum (TH. FR.) Z~~~~~.=Leprocaulon microscopicum (VILL.) 

GAMS ex D. HAWKSW.; see LAMB and WARD, 1974, p. 526. 

St. roccella (L.) RAusc~. = Roccella tinctoria (L.) DC. 

St. sertularia ACH. Not a lichen; see ZAHLBR. Cat. Lich. Univ. 4: 673 (1927). 

St. soleirolii DUE. =Lethariella intricata (MORIS) KROG, Norw. J. Bot. 23: 95 (1976). 

St. subalbicans L~~~=Leprocaulon subalbicans (LAMB) LAMB et WARD, 1974, p. 534. 

St. subgracilescens LAMB ex VARGAS, Revista Univ. (Cuzco), 37 (94): 187 (1948) (nomen 

nudum)=Leprocaulon albicans (TH. FR. emend.) NYL., Ch. str. 11; see LAMB and WARD, 

1974, p. 514. 

St. tabulare (THUNB.) Ac~.=Siphula tabularis (THUNB.) NYL. Correct name is Siphula 

verrucigera (GMEL.) SANT. ; see MATHEY, Nova Hedwigia, 22 (314): 805 (1971). 

St. tenellum Tuc~.=Leprocaulon tenellum (TUCK.) NYL.; see LAMB and WARD, 1974, p. 537. 

St. vermiculare (Sw.) RAUSCH. = Thamnolia vermicularis (Sw.) ACH. 

C. Rejected taxa 

Lecidea artyta ACH. 

Lichenographia Universalis : 170 (1810). 

This is listed by ZAHLBRUCKNER, 192627 (1927), p. 633, as a synonym of Stereocaulon 

alpinum LAUR. We were unable to find any type material in H-ACH, but in s there is a small 

specimen without locality, "Herb. Swartzii", labelled as Lecidea artyta, in Ac~~~rus's handwriting; 

it consists of 2 fragments, one a Toninia species, the other Stereocaulon alpinum. 

According to TH. FRIES, 1874, p. 336, the original material of Lecidea artyta was a mixture of 

Stereocaulon tomentosum /3. alpinum (-St. alpinum) and St. denudatum /3. pulvinatum (=St. 

vesuvianum var. nodulosum f. umbonatum), to which was later (after the publication of the 

description) added a specimen of Toninia syncornista (FLORKE) TH. FR. (= T. lobulata (SOMM.) 

LYNGE). Lecidea artyta therefore seems to have been founded on a mixture of 2 Stereocaulon 

species, and the epithet should be discarded in accordance with Art. 70 of the Int. Code. 

Stereocaulon alpinum P. botryosum 2. subcymosum FLOT. 

Bot. Zeitung (Berlin), 8: 538 (1850). 

A nomen nudum; the only observation given is "Ciirie et Breutel Herb. No. 140a. Gron-

land". 

I. MACKENZ~E LAMB: A Conspectus of Stereocaulon 

St. corallinum c. stigmateum RABENH. 

Deutschlands Kryptogamen-Flora, 2, 1. Abt., Lichenen: 112 (1845). 

Typus: Switzerland, Susten, leg. SCHAERER; not seen by us. The description, "mit 

schwarz punktirten, kornigen Schuppchen", indicates a state infested by a parasymbiontic 

fungus. To be rejected (Int. Code Art. 70). 

"Stereocaulon globosus (HuDs.) VAIN." 

This name is given to nos. 27 and 87 of ANDERS'S exsiccat Lich. EXS. Bohem. Bor. The 

former is St. saxatile MAGN., the latter a mixture of St. evolutum GRAEWE and St. tomentosum 

var. compactum FREY. The combination is not mentioned in ANDERS'S book (1928), and 

we cannot trace it to VAINIO. It is probably merely a lapsus due to confusion with a name 

applied to a species of Sphaerophorus, and should be rejected and ignored. 

St. par vifolia 

A nomen nudum without indication of author, mentioned on pages 313 and 480 of H. 

B. DAVIS, "Life and Work of Cyrus Guernsey Pringle" (1936), referring to a specimen numbered 

414 collected by PRINGLE in Mexico. Dr. F. SEYMOUR, Assistant Curator of the PRINGLE 

Herbarium (VT) informed us in litt. 1967 that no specimen with this name could be found in 

their collect ions. 

St. paschale ,B. conglomeratum FR. 

Schedulae Criticae de Lichenibus Exsiccatis Sueciae, Part 111: 20 (1825). 

Typus: Sweden, no exact locality. The isotype material is FRIES, Lich. Suec. Exs. no. 89. 

This is not homogeneous; some examples, as in FH, are St. evolutum GRAEWE, others, as in 

LD and UPS, a mixture of St. evolutum and St. paschale (L.) HOFFM., and yet others, as in 

s, entirely St. paschale. The original description does not give much clue as to which of these 

species FRIES had in mind, but probably mostly St. evolutum. TH. FRIES (1865, p. 181) also 

stated that Lich. Suec. Exs. no. 89 is a mixture, partly consisting of St. evolutum; FREY (1932, 

p. 149) states that it contains also St. subcoralloides (NYL.) NYL. In FRIES, Lichenographia 

Europaea Reformata: 202 (1831) the same taxon is referred to as St. paschale b. minus FR.; 

the word "conglomeratum" forming the description; reference is made to the original Sched. 

Critic. publication. TH. FRIES (1857, p. 17) made the combination St. coraZZoides [var.] P. 

~onglomeratum (FR.) TH. FR., referring to FR. Lich. Europ. Reform. p. 202, and hence to 

St. paschale p. conglomeratum FR. as basionym. In Lichenogr. Scand. (1871) TH. FRIES 

refers to the taxon in 2 places: on p. 44 as St. coralloides P. conglomeratum, using it in the sense 

of St. subcoralloides (NYL.) NYL., and on p. 47 as St. paschale E. conglomeratum. ZAHLBRUCK- 

NER (1926-27, p. 667) lists St. coralloides P. conglomeratum as a synonym of St. subcoralloides. 

Finally, VAINIO (1940, p. 10) made the combination Stereocaulon conglomeratum ("TH. FR.") 

VAIN., and applied it to St. subcoralloides. We subscribe to the opinion of MAGNUSSON 

(1926, p. 51) that the epithet conglorneratum is so thoroughly confused in its application 

that it should be dropped entirely. To be rejected (Int. Code Art. 69). 

St. paschale ,B. conglomeratum 1. robustum FLOT. 

Bot. Zeitung (Berlin), 8: 537 (1850). 

A nomen nudum; the only observation given is "Curie et Breutel Herb. No. 141. Gron- 

land."


St. paschale [var.) E. pulvinatum SCHAER. 

Lichenum Helveticorum Spicilegium, Pars prima, Sect. VI: 274 (1833). - St. corallinum [f.] b. pulvinaturn 

(SCHAER.) RABENH. Deutschlands Kryptogamen-Flora, 2, Abt. 1: 112 (1845). -St. corallinum 

[var.] B. pulvinatum (SCHAER.) SCHAER. Enumeratio Critica Lichenum Europaeorum: 180 (1850). - St. 

denudatum [var.] y. pulvinatum (SCHAER.) FLOT. Bot. Zeitung (Berlin), 8: 539 (1850). - Stereocaulon 

pulvinatum (SCHAER.) NYL. Bull. SOC. Linn. Normandie, ser. 4, 1: 201 (1887) (nom. illegit.; non AcH.). 

- St. vesuvianum var. pulvinatum (SCHAER.) "LAMB"; DUNCAN, A Guide to the Study of Lichens: 79 

(1959) (comb. inval.); P. JAMES, Lichenologist, 3: 98 (1965) (val.). - St. coralloides var. pulvinatum 

"SCH."; (SCHAER.) MALBR. Bull. Soc. Amis Sci. Nat. Rouen, 3: 439 (1868). -St. denudatum var. 

umbonatum f. pulvinata (SCHAER.) L'D~~~~"; RAs. Lich. Fenn. Exs. no. 397 (1939). - St. denudatum 

[subsp.l*pulvinatum (SCHAER.) HUE, NOUV. Arch. Mus. Hist. Nat. ser. 3,2: 247 (sep. 39) (1890). 

Typus: Switzerland, Susten, leg. SCHAERER (date ?) (G-SCHAER, not seen by us). 

The epithet has been widely used to designate St. vesuvianum var. nodulosum f. umbonaturn 

(WALLR.) LAMB, but FREY (1932), who saw the type material, testifies that it is a mixture, 

consisting mainly of St. botryosum ACH. em. FREY, and not including any specimen referable 

to St. vesuvianum PERS. sens. lat. To be rejected as based on discordant elements (Int. Code 

Art. 70). 

St. pileatum f. sorediiferum sub f. terrestre HARM. 

Lichens de France, Fasc. 111: 369 (1907). 

No description given, merely a statement of its occurrence on bare siliceous soil. The 

taxon is therefore not validly published. 

St. pruinosum FLORKE 

Deutsche Lichenen, Lief. IV: 14 (1819). 

A nomen nudum; no description or locality given. 

St. tomentosum [var.] p. alpestre f. stigmatea FLOT. 

Flora, Jahrg. 19, 1, Beiblatt 1: 17 (1836). 

Described from several localities in the Riesengebirge (Polish-Czechoslovakian border), 

no holotype designated. Type material presumably in herb. FLOTOW (B) and therefore 

destroyed in World War 11. 

If, as seems to be the case, it is a state (perhaps not of St. tomentosum) with fruit bodies 

of a parasymbiont, cfr. Catillaria stereocaulorum (TH. FR.) OLIV., it is to be rejected as based on 

disparate elements. KORBER, however (Syst. Lich. German. p. 11, 1855) considered it to represent 

the spermogoniiferous condition of St. tomentosum. 

St. tomentosum var. karelica VAIN. 

ex Rjis. Ann. Bot. Soc. Zool. Bot. Fenn. Vanamo, 12 (1): 54 (1939). 

Typus: U.S.S.R., Karelo-Finnish SSR, Valamo Island in Lake Ladoga, leg. E. A. VAINIO, 1923 

(TUR-VAIN, holotypus; UPS-MAGN, ~SO~YPUS). 

Specimen in poor condition, not with certainty referable to St. tomentosum. The sorediate 

efflorescences belong to some other lichen (Lepraria sp.?) growing epiphytically over it. 

To be rejected as based on discordant elements (Int. Code, Art. 70). 

St. tomentosum B. majus SCHAER. 

Lichenum Helveticorum Spicilegium, Pars prima, Sect. VI: 276 (1833). 

and


St. tomentosum [var.] a. granulosum SCHAER. 

Enumeratio Critica Lichenum Europaeorurn : 181 (1 850). 

Both these taxa were founded on the same type material from Switzerland, Susten, "ad 

torrentes alpinos", leg. SCHAERER and distributed by him in his exsiccat Lich. Helvet. Exs. 

Fasc. XI (1836) no. 262, under the name Stereocaulon tomentosum B. majus. This number is 

however very heterogeneous; copies seen by FREY contained St. alpinum LAUR., St. tomentosum 

FR. and St. tomentosum var. alpestre FLOT. (FREY, 1932), and those seen by us were St. alpinum 

(PC) and St. alpinum var. erectum FREY (FH-TUCK). SCHAERER'S brief and vague descriptions 

do not clearly indicate any particular one of these taxa, and in our opinion both his epithets 

should be discarded as nomina confusa (Int. Code, Art. 70). 

APPENDIX 3 

AK: Natural History Museum, Akureyri, Iceland. 

ALA: University of Alaska, College, Alaska. 

ASAH: Herbarium of Y. ASAHINA (died 1975), Tokyo, Japan. 

AWAS: Herbarium of D. D. AWASTHI, Lucknow, India. 

B: Botanisches Museum, Berlin-Dahlem, Germany. 

BA: Museo Argentino de Ciencias Naturales "Bernardino Rivadavia", Buenos Aires, Argentina. 

B. DE LESD: Herbarium of M. BOULY DE LESDAIN, Dunkerque, France (destroyed in World 

War 11). 

BFU: Institut fiir Systematische Botanik der Freien Universitat, Berlin, Germany. 

BG : Universitetets Botaniske Museum, Bergen, Norway. 

BIRM: Department of Botany, University of Birmingham, Birmingham, England. 

BM: British Museum (Natural History), London, England. 

BM-BAB: Herbarium of CHURCHILL BABINGTON (formerly in CGE). 

BOL: Bolus Herbarium, University of Cape Town, South Africa. 

BP: Department of Botany, Museum of Natural History, Budapest, Hungary. 

BRATT: Herbarium of G. C. BRATT, Department of Botany, University of Tasmania, Hobart, 

Tasmania. 

BRLU: Laboratoire de Botanique SystCmatique et de PhytogCographie, Universitt Libre de 

Bruxelles, Bruxelles, Belgium. 

BRNU: Institutum Botanicum Acad. Scient. i'echoslovacae, Brno, Czechoslovakia. 

BRSL: Herbarium, Botanical Institute of the University, Wroclaw (formerly Breslau), Poland. 

c: Botanical Museum and Herbarium, University of Copenhagen, Copenhagen (K0benhavn), 

Denmark. 

CAN: National Museum of Canada, Ottawa, Ontario, Canada. 

CANE: Commonwealth Scientific and Industrial Research Organization, Division of Plant 

Industry, Canberra, Australia. 

CGE: Botany School, University of Cambridge, Cambridge, England. 

CHR: Botanical Division of the Department of Scientific and Industrial Research, Christchurch, 

New Zealand. 

COLO: University of Colorado Museum, Boulder, Colorado, U.S.A. 

cuw: Department of Botany, Clark University, Worcester, Massachusetts, U.S.A.


DAO: Plant Research Institute, Department of Agriculture, Ottawa, Ontario, Canada. 

DEGEL: Herbarium of G. N. DEGELIUS, Askim, Sweden. 

DODGE: Herbarium of C. W. DODGE, C/O University of Vermont, Burlington, Vermont, 

U.S.A. 

DSIR: Botany Division, Department of Scientific and Industrial Research, Wellington, 

New Zealand. 

DUKE: Department of Botany, Duke University, Durham, N. Carolina, U.S.A. 

E: Royal Botanic Garden, Edinburgh, Scotland. 

EDWIL: Canadian Wildlife Service, Edmonton, Alberta, Canada. 

EYERD: Herbarium of W. J. EYERDAM, Seattle, Washington, U.S.A. 

FH: Farlow Herbarium of Cryptogamic Botany, Harvard University, Cambridge, Massachusetts, 

U.S.A. 

FH-RIDD: Herbarium of L. W. RIDDLE, in Farlow Herbarium. 

FH-TAYL: Herbarium of T. TAYLOR, in Farlow Herbarium. 

FH-TUCK: Herbarium of E. TUCKERMAN, in Farlow Herbarium. 

FI: Herbarium Universitatis Florentinae, Istituto Botanico, Firenze, Italy. 

FREY: Herbarium of E. FREY (died 1974), Miinchenbuchsee, Bern, Switzerland (now in Botanisches 

Institut der Universitat, Bern). 

G: Conservatoire et Jardin Botaniques, Genkve, Switzerland. 

G-MULL: Herbarium of J. MULLER (ARC.), in Conservatoire et Jardin Botaniques. 

G-SCHAER: Herbarium of L. E. SCHAERER, in Conservatoire et Jardin Botaniques. 

GALLOWAY: Herbarium of D. J. GALLOWAY, C/O Department of Scientific and Industrial 

Research, Palmerston North, New Zealand. 

GB: Goteborgs Botaniska Tradgird, Goteborg, Sweden. 

czu: Institut fur Systematische Botanik, Universitat, Graz, Austria. 

H: Botanical Museum, University of Helsinki, Helsinki, Finland. 

H-ACH: Herbarium of E. ACHARIUS, in Botanical Museum, University of Helsinki. 

H-NYL: Herbarium of W. NYLANDER, in Botanical Museum, University of Helsinki. 

H-RAS: Herbarium of V. RASANEN, in Botanical Museum, University of Helsinki. 

HAK: Herbarium of R. HAKULINEN, Kyro, Finland. 

HBG: Staatsinstitut fur Allgemeine Botanik, Hamburg, W. Germany. 

HKU: Department of Botany, University of Hong Kong, Hong Kong. 

HUN: Herbarium of S. HUNECK, Halle-Neustadt, German Democratic Republic. 

K: Royal Botanic Gardens Herbarium, Kew, Surrey, England. 

KASSEL: Naturkundemuseum der Stadt Kassel, Kassel, W. Germany. 

KERSH: Herbarium of K. KERSHAW, Imperial College of Science and Technology, London, 

England. 

KIENER: Herbarium of W. KIENER (died 1959), Game, Forestation and Parks Commission, 

Lincoln, Nebraska, U.S.A. 

KLEM: Herbarium of 0. KLEMENT, Kreuzthal-Eisenbach, Allgau, W. Germany. 

L: Rijksherbarium, Leiden, Netherlands. 

L-PERS: Herbarium of C. H. PERSOON in Rijksherbarium. 

LAE: Department of Forests, Papua and New Guinea, Lae, New Guinea. 

LD: Botanical Museum and Herbarium, Lund, Sweden. 

LE: Botanical Institute of the Academy of Sciences of the USSR, Leningrad, U.S.S.R.


LEPAGE: Herbarium of E. LEPAGE, Ecole #Agriculture, Rimouski, Quebec, Canada. 

LG: Institut et Jardin Botanique de I'UniversitC, Litge Belgium. 

LE GALLO: Herbarium of P. C. LE GALLO, Guadeloupe, Leeward Islands, West Indies. 

LIL: Instituto Miguel Lillo, Tucumin, Argentina. 

LSU: Herbarium, Louisiana State University, Baton Rouge, Louisiana, U.S.A. 

LWG: National Botanic Gardens, Lucknow, India. 

M: Botanische Staatssammlung, Miinchen, W. Germany. 

M-ARN: Herbarium of F. ARNOLD, in Botanische Staatssammlung. 

M-KREMP: Herbarium of A. v. KREMPELHUBER, in Botanische Staatssammlung. 

MAASS: Herbarium of W. MAASS, Atlantic Regional Laboratory, Halifax, Nova Scotia, 

Canada. 

MB: Botanisches Institut der Universitat, MarburgILahn, W. Germany. 

MC CULLOUGH: Herbarium of H. A. Mc CULLOUGH, Samford University, Birmingham, 

Alabama, U.S.A. 

MEL: Botanic Garden and National Herbarium, Melbourne, Victoria, Australia. 

MEXPOL: Instituto PolitCcnico Nacional, Mexico City, Mexico. 

MICH: University Herbarium, University of Michigan, Ann Arbor, Michigan, U.S.A. 

MO: Missouri Botanical Garden, Saint Louis, Missouri, U.S.A. 

MSC: Department of Botany, Michigan State University, East Lansing, Michigan, U.S.A. 

MSK: Biological Institute of the Academy of Sciences of the Byelorussian SSR, Minsk, 

U.S.S.R. 

NEBC: Herbarium of New England Botanical Club, c/o Gray Herbarium, Haward University, 

Cambridge, Massachusetts, U.S.A. 

NFLD: Department of Biology, Memorial University of Newfoundland, Saint John's, Newfoundland, 

Canada. 

NICH: Hattori Botanical Laboratory, Nichinan, Japan. 

NY: New York Botanical Garden, Bronx Park, New York, New York, U.S.A. 

o: Botanisk Museum, Oslo, Norway. 

oc: Department of Botany, Oberlin College, Oberlin, Ohio, U.S.A. 

ORE: Herbarium of the University of Oregon, Eugene, Oregon, U.S.A. 

PC: Laboratoire de Cryptogamie, Museum National d'Histoire Naturelle, Paris, France. 

PC-HUE: Herbarium of A. M. HUE, in Laboratoire de Cryptogamie. 

PC-MONT: Herbarium of J. F. C. MONTAGNE, in Laboratoire de Cryptogamie. 

PH: Academy of Natural Sciences, Philadelphia, Pennsylvania, U.S.A. 

PR: Botanical Department of the National Museum, Praha, Czechoslovakia. 

QMP: MusCe de la Province, QuCbec City, Quebec, Canada. 

RB: Jardim Botfinico, Rio de Janeiro, Brazil. 

REN : Laboratoire de Botanique, UniversitC de Rennes, Rennes, France. 

s: Botanical Department, Naturhistoriska Riksmuseum, Stockholm, Sweden. 

SASK: W. P. FRASER Memorial Herbarium, Department of Plant Ecology, University of 

Saskatchewan, Saskatoon, Saskatchewan, Canada. 

SCHINDL: Herbarium of H. SCHINDLER, Karlsruhe, W. Germany. 

SGO: Museo Nacional de Historia Natural, Santiago, Chile. 

SOVMONG: Soviet-Mongolian Complex Biological Expedition, Bolshaya Ordynka 21/16, 

Moskva, U.S.S.R. 

SOWTER: Herbarium of F. A. SOWTER (died 1972), Stoughton, Leicestershire, England.


STR: Institut de Botanique de la Facult6 des Sciences de 11Universit6, Strasbourg, France. 

SUND: Herbarium of S. W. SUNDELL, Munkfors, Sweden. 

SWINSC: Herbarium of T. D. V. Swr~scow, Topsham, Exeter, Devon, England. 

TAI: Department of Botany, National Taiwan University, Taipei, Taiwan, China. 

TENN: Herbarium, Department of Botany, University of Tennessee, Knoxville, Tennessee, 

U.S.A. 

TI: Botanical Institute, University of Tokyo, Tokyo, Japan. 

TNS: Botanical Section, National Science Museum, Tokyo, Japan. 

TRH: Botanical Department, Museum of the Royal Norwegian Society for Science and Letters, 

Trondheim, Norway. 

TRTC: Cryptogamic Herbarium, Department of Botany, University of Toronto, Toronto, 

Ontario, Canada. 

TU: Department of Taxonomy and Geobotany of Tartu State University, Tartu, Estonian 

SSR, U.S.S.R. 

TUR: Botanical Institute of the University, Turku, Finland. 

TUR-VAIN: Herbarium of E. A. VAINIO, in Botanical Institute of the University. 

u: Botanical Museum and Herbarium, University of Utrecht, Utrecht, Netherlands. 

UBC: Department of Botany, University of British Columbia, Vancouver, British Columbia, 

Canada. 

uc: Herbarium of the University of California, Berkeley, California, U.S.A. 

UPS: Institute of Systematic Botany, University of Uppsala, Uppsala, Sweden. 

UPS-MAGN: Herbarium of A. H. MAGNUSSON, in Institute of Systematic Botany. 

UPS-TH FR: Herbarium of TH. FRIES, in Institute of Systematic Botany. 

UPS-THUNB: Herbarium of C. P. THUNBERG, in Institute of Systematic Botany. 

us: Department of Botany, Smithsonian Institution, U. S. National Museum, Washington, 

District of Columbia, U.S.A. 

USGO: Departamento de Botanica, Universidad de Chile, Santiago, Chile. 

usw-MAHU: Herbarium of M. MAHU, in Departamento de Botanica. 

v: Provincial Museum, Victoria, British Columbia, Canada. 

VARGAS: Herbarium of C. VARGAS, Universidad de Cuzco, Cuzco, Peru. 

VORAG: Institute of Agronomy, Voronesh (Woronesch), U.S.S.R. 

VT: Pringle Herbarium of the University of Vermont, Burlington, Vermont, U.S.A. 

w: Botanische Abteilung, Naturhistorisches Museum, Wien, Austria. 

: Botanisches Institut der Universitat, Wurzburg, W. Germany. 

WELC: Department of Botany, Wellesley College, Wellesley, Massachusetts, U.S.A. 

WIS: University of Wisconsin Herbarium, Madison, Wisconsin, U.S.A. 

WTU: Herbarium, Botany Department, University of Washington, Seattle, Washington, 

U.S.A. 

YU: Herbarium of Osborn Botanical Laboratory, Yale University, New Haven, Connecti- 

cut, U.S.A. 

z: Botanischer Garten und Museum der Universitat, Zurich, Switzerland. 

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SUMMARY 

An emended subgeneric classification of the genus Stereocaulon (SCHREB.) HOEEM. is presented. 

123 species are recognized, with 47 varieties and 40 forms. Literature references, synonymy, exsiccata, 

useful illustrations, chemical constituents and general geographical distribution are listed for each 

species. All taxa are typified, 51 lectotypes and 1 neotype being newly selected. The following taxa are 

described as new: 

Species: St. austroindicum LAMB, St. brassii LAMB, St. crambidiocephalum LAMB, St. didymicurn 

LAMB, St. dusenii LAMB, St. fibrillosum LAMB, St. fronduliferum LAMB, St. halei LAMB, St. himalayense 

AWAS. et LAMB, St. novogramtense LAMB, St. papuanum LAMB, St. paradoxurn LAMB, St. procerum 

GROENH. ex LAMB, St. rugulosum LAMB, St. sibiricum LAMB, St. togashii LAMB, St. uliginosum LAMB, 

St. wadei LAMB, St. weberi LAMB. 

Varieties: St. colensoi var. caliginosum LAMB, St. cornutum var. corallizans LAMB, St. corticatulum 

var. cornplanaturn LAMB, St. myriocarpum var. altaicum LAMB, St. pileatum var. nipponicum LAMB, St. 

ramulosum var. perpumilum LAMB, St. ramulosum var. simplicius LAMB, St. sasakii var. tomentosoides 

LAMB, St. sorediiferum var. Ieprosolingulatum LAMB, St. staufferi var. bornefnse LAMB, St. tennesseense


var. nigrofastigiatum LAMB, St. tomentosum var. capitatum LAMB, St. verruculigerum var. subfurfurascens 

LAMB, St. vesuvianum var. thyrsoideum LAMB. 

Forms: St. botryosum f. subglobuliferum LAMB, St. dactylophyllum f. sessile LAMB, St. intermedium 

f. compactum LAMB, St. meyeri f. tucumanum LAMB, St. nunodes f. schadeanum LAMB, St. nigromaculatum 

f. fastigiato-pulvinatum LAMB, St. ramulosum f. subcompressum LAMB, St. ramulosum f. tomentosulum 

LAMB, St. ramulosum var. pulvinare f. crebratum LAMB, St. subcoralloides f. sorediascens LAMB, St. 

vesuvianum var. nodulosum f. verrucosum LAMB. 

New name: St. melanopotamicum LAMB. 

The following new combinations are made: St. arcticum var. turjadnse (Us.) LAMB, St. atlanticum 

(LAMB) LAMB, St. corticatulum var. capense (LAMB) LAMB, St. foliolosum var. botryophorum (MULL. 

ARC.) LAMB, St. glabrum f. jfabellans (LAMB) LAMB, St. glaucescens var. caespitosulum (NYL.) LAMB, 

St. meyeri f. acuminatum (MULL. ARC.) LAMB, St. pomiferum f. mikenoense (DUVIGN.) LAMB, St. ramulosum 

var. exalbidum (NYL.) LAMB, St. ramulosum var. gracilius (M~~LL. ARC.) LAMB, St. ramulosum f. 

nudatum (MULL. ARC.) LAMB, St. ramulosum var. pilophoroides (TUCK.) LAMB, St. ramulosum var. 

pulvinare (DODGE) LAMB, St. ramulosum var. submollescens (NYL.) LAMB, St. rivulorum f. pulvinarium 

(SAV.) LAMB, St. sasakiivar. simplex (RIDD.) LAMB, St. saxatile f. sorediatum (MAGN.) LAMB, St. spathuliferum 

f. dissolutum (MAGN.) LAMB, St. spathuliferurn f. pygmaeum (MAGN.) LAMB, St. stenospermum 

(LAMB) LAMB, St. verruciferum var. surreptans (LAMB) LAMB, St. vesuvianum var. nodulosum f. umbricolum 

(FREY) LAMB, St. vesuvianum var. nodulosum f. depressum (MAGN.) LAMB, St. vesuvianum var. 

obscurum (MULL. ARC.) LAMB, St. vulcani f. maunae-loae (MAGN.) LAMB. 

New taxonomic positions (n. stat.) are given to the following 4 infrageneric taxa: subgen. Stereocaulon, 

sect. Denudata, subsect. Botryoideum (G. T. JOHNS. emend.) LAMB; sect. Lobophoron (DUVIGN.) 

LAMB; subgen. HoIostelidium, sect. Holostelidium, subsect. Dactyloideum (LAMB) LAMB; and subgen. 

Pilophoropsis (LAMB) LAMB. 

91 doubtful, excluded or rejected taxa are also listed. 

Argopsis cymosa (CROMB.) STIZENB. 21 1 Lecidea artyta ACH. 328 

Baeomyces denudatus (FL~RKE) HEPP 244 Leprocaulon Nn. 193 

-paschalis (L.) WAHLENB. 200 

- - 8. nanum (AcH.) WAHLENB. 326 

- delisei (BORY) OLIV. 291 

Lichen sect. Stereocaulon SCHREB. 200 

Cereolus condensatus (HOFFM.) BOIST. 209 - condensatus (HOFFM.) ACH. 209 

-- subvar. acaulon (NYL.) BOIST. 209 - crustaceus GILIB. 200, 309 

-- subvar. curtulus (NYL.) BOIST. 322 

-paschalis L. 200 

- pileatus (AcH.) BOIST. 226 - ramulosus Sw. 278 

-- var. sorediifer (NYL.) BOIST. 226 - salazinus BORY 302 

Chlorocaulum CLEMENTS 196, 199 - vulcani BORY 257 

- salazinum (BORY) CLEMENTS 302, 303 

Patellaria paschalis (L.) WALLR. 200 

Cladonia paschalis (L.) WIGG. 200 -- var. bracteata WALLR. 244, 31 5 

Coralloides HOFFM. 195 -- var. nodulosa WALLR. 244 

- WOLF 195 -- var. umbonata WALLR. 246 

- paschale (L.) HOFFM. 200 

-pileata var. crustacea WALLR. 209, 310 

Corynophoron NYL. 197 -- var. thamnodes WALLR. 209, 310 

- colensoi (BAB.) NYL. 287 

Gymnocaulon DUVIGN. 198 

- tomentosa var. decumbens WALLR. 236, 314 

- - p. erecta WALLR. 221, 312 

- caespitosum (RED.) DUVIGN. 306 

Phyllocaulon (TUCK.) VAIN. 196 

- gregarium (RED.) DUMGN. 307 - wrightii (TUCK.) VAIN. 259 

Lecanocaulon NYL. 196 

Pilophoron colensoi (BAB.) KNIGHT 287 

Synonyms and doubtful, excluded or rejected taxa are in italics.


-- var. reagens (RAs.) ZAHLBR. 218 

Stereocaulomyces paschalis (L.) THOMAS 200 

Stereocaulon (SCHREB.) HOFFM. 192, 195 

- sect. Botryoideum G. T. JOHNS. 196 

- subgen. Cylindrocladium DUVIGN. 197 

-- sect. Corynophoron (NYL.) DUVIGN. 198 

-- sect. Lecanocaulon (NYL.) DUVIGN. 196 

-- sect. Rhopalophoron DUVIGN. 197 

- subgen. Enteropodium LAMB 195 

-- sect. Chondrocaulon TH. FR. 193 

-- sect. Eustereocaulon ser. Granulata 

DUVIGN. 195 

--- ser. Umbilicata DUVIGN. 196 

-- sect. Leucocheilon LAMB 195 

--- subsect. Peltophyllum (Du RIETZ) 

LAMB 196 

- sect. Eustereocaulon KORB. 195 

- subgen. Eustereocaulon JATTA 195 

-- sect. Paschalia JATTA 195 

- subgen. Holostelidium LAMB 197 

-- sect. Dactyloideum LAMB 198 

-- sect. Ecephalodiata LAMB 196 

-- sect. Holostelidium 197 

--- subsect. Aciculisporae Du RIETZ 197 

--- subsect. Dactyloideum (LAMB) LAMB 

198 

--- subsect. Holostelidium 197 

-- sect. Pilophoropsis LAMB 197 

-- sect. Redingeria LAMB 198 

-- sect. Sacculata subsect. Ascaridisporae 

DU RIETZ 197 

- subgen. Lecidocaulon sect. Verruculata 

SAT6 196 

- subgen. Lecanocaulon (NYL.) VAIN. 196 

-- DUVIGN. (non NYL., nec VAIN.) 197 

- subgen. Lobophoron DUVIGN. 197 

- subgen. Phyllocaulon TUCK. 196 

- sect. Phyllocaulon (TUCK.) Us. 196 

- subgen. Pilophoropsis (LAMB) LAMB 197 

- subgen. Pilophoropsis DUVIGN. 195 

- subgen. Platycladium DUVIGN. 195 

-- ser. Granulata DUVIGN. 195 

-- ser. Umbilicata DUVIGN. 196 

- subgen. Rhopalophoron DUVIGN. 197 

-- ser. Euclavulifera DUVIGN. 197 

-- ser. Protoclavulifera DUVIGN. 197 

- sect. Sacculata SATB 197 

- subgen. Stereocaulon 195, 200 

-- sect. Denudata JATTA 195 

--- subsect. Botryoideum (G. T. JOHNS.) 

LAMB 196 

--- subsect. Denudata 196 

--- subsect. Phyllocaulon (TUCK.) LAMB 

196 

-- sect. Lobophoron (DUVIGN.) LAMB 197 

-- sect. Stereocaulon 195, 200 

-- sect. Stereocladium (NYL.) DODGE 195 

- abduanum ANZI 222, 3 12 

- acaulon NYL. 209, 3 10 

- aciculare (AcH.) TUCK. 326 

- aggregatum (Sw.) RAUSCH. 326 

- albicans TH. FR. 326 

- alpinum LAUR. 202 

-- f. adpressum MAGN. 202, 309 

-- f. alpestre (FLoT.) TH. FR. 238 

-- var. botryosum (AcH.) SCHAER. 207 

-- f. botryosum (AcH.) FR. 207 

-- p. botryosum 2. subcymosum FLOT. 328 

-- var. erectum FREY 203 , 

-- var. glabrum MULL. ARG. 21 8 

-- var. gracilentum (TH. FR.) MAGN. 204 

-- f. gracilentum (TH. FR.) LAMB 204 

-- var. grande (MAGN.) FREY 220 

-- var. Janii LYNGE 250 

-- f. platycladum FREY 203 

-- f. pulvinarium SAV. 229 

-- var. tyroliense (NYL.) ARN. 240 

-- var. vesuvianum (PERs.) LINK 242 

-- f. walamo&nse (NYL.) FREY 325 

- andinum B. DE LESD. 326 

- anomalum LAMB 249 

- antarcticum VAIN. 204 

- aogasimense ASAH. 264 

- apocalypticum NYL. 259 

- arbuscula (NYL.) NYL. 326 

-- var. aberrans ASAH. 326 

- arcticum LYNGE 249 

-- var. turjaense (Us.) LAMB 250 

- arenarium (SAV.) LAMB 250 

- argodes NYL. 286, 3 19 

- argus HOOK. F. et TAYL. 286 

-- var. stenospermum LAMB 304 

- armatulum ZAHLBR. 263, 317 

- assimile NYL. 302, 321 

- atlanticum (LAMB) LAMB 286 

- austroindicum LAMB 205 

- azoreum (SCHAER.) NYL. 205 

- botryocarpum MAGN. 238 

- botryophorum M~~LL. ARC. 267 

- botryosum ACH. 207

-- f. confluens (MAGN.) FREY 216 

-- f. congestum (MAGN.) FREY 208 

-- f. depressum FREY 216 

-- var. dissolutum (MAGN.) MAGN. 233 

-- f. dissolutum (MAGN.) FREY 233 

-- f. finmarkicum (MAGN.) FREY 208 

-- f. globuliferum (MAGN.) FREY 234 

-- f. irregulare (MAGN.) FREY 208 

-- subsp. islandicum DUVIGN. 208, 309, 321 

-- f. pygmaeum MAGN. 234 

-- f. sirnplicior (MAGN.) FREY 208 

-- var. spathuliferum (VAIN.) LAMB 233 

-- f. spathuliferum (VAIN.) FREY 233 

-- f. subglobuliferurn LAMB 208 

-- var. subincrustaturn FREY 202, 309 

-- subsp. vesuvianum (PERs.) ACH. 242 

- brassii LAMB 271 

- buchanani KNIGHT 327 

- caespitosurn RED. 306 

- capense LAMB 289 

- capitellaturn MAGN. 209 

- carinthiacum FREY 241 

- cereolinum ACH. 327 

-- var. cereolus (AcH.) TH. FR. 327 

-- f. pileatum (AcH.) TH. FR. 226 

- cereolus ACH. 327 

- chlorellum TUCK. 327 

- chlorocarpoides ZAHLBR. 272 

- cinchonae (WILLD.) RAUSCH. 327 

- claviceps TH. FR. 271 

-- var. pomiferum (DUVIGN.) LAMB 276 

-- var. tuberaturn LAMB 276 

-- var. yunnanense HUE 276, 319 

- colensoi BAB. 287 

-- var. caliginosum LAMB 288 

-- var. reagens RAS. 218, 311 

- cornrnixtum (ASAH.) ASAH. 260 

- condensatum HOFFM. 209 

-- var. acaulon (NYL.) OLN. 209 

-- f. acaulon (Nn.) MAGN. 209 

-- f. athallinurn SANDST. 210, 3 10 

-- subsp. condyloideum (AcH.) NYL. 210 

-- var. condyloideum (AcH.) NYL. 210 

-- f. condyloideum (AcH.) MAGN. 210 

-- f. crustaceum (WALLR.) RABENH. 209 

-- var. minor EGELING 210, 310 

-- f. septentrionale MAGN. 210, 310 

-- var. sorediatum HARM. 226, 31 2 

- condyloideum ACH. 210, 310 

- confine (0. M~L.) HOFFM. 327 


- confluens MULL. ARC. 248, 316 

-- f. fastigiata DUVIGN. 321 

-- var. fuscescens MULL. ARC. 248, 316 

-- f. gracilius DUVIGN. 321 

-- f. persquamosa DUVIGN. 248 

-- f. ramosa DUVIGN. 321 

-- f. typica DUVIGN. 321 

-congestumN~~. 327 

- conglomeratum (FR.) VAIN. 235, 329 

- coniophyllurn LAMB 268 

- coralligerum MEYER 274, 321 

- corallinoides HOFFM. 327 

- corallinum (L.) SCHRAD. 212, 327 

-- var. pulvinatum (SCHAER.) SCHAER. 330 

-- f. pulvinatum (SCHAER.) RABENH. 330 

-- c. stigmateum RABENH. 329 

- coralloides FR. 212, 310 

-- var. conglomeratum (FR.) TH. FR. 329 

--- f. pumilum NYL. 227 

-- var. dactylophyllum (FLORKE) TH. FR. 21 1 

-- subvar. dactylophyllum (FLORKE) BOIST. 

211 

-- f. dactylophyllum (FLORKE) TH. FR. 21 1 

-- subsp. delisei (BORY) NYL. 291 

-- f. expansum MAGN. 213 

-- var. flabellaturn FREY 214 

-- f. intermedium SAV. 223 

-- var. japonicum (TH. FR.) NYL. 263 

-- var. occidentale MAGN. 214 

-- var. orientalis RAS. 212, 310 

-- var. pulvinatum (SCHAER.) MALBR. 330 

-- var. pumilum (NYL.) OLIV. 227 

-- var. pygmaeum FREY 21 5 

-- var. spissum (NYL.) LAMB 215 

-- f. subbotryosum FREY 214 

- cornutum M ~L. ARC. 251 

-- var. corallizans LAMB 251 

- corticatulum NYL. 288 

-- var. capense (LAMB) LAMB 289 

-- var. complanatum LAMB 289 

-- subsp. detergens NYL. 288, 320 

-- var. detergens (NYL.) MULL. ARC. 288 

-- var. humile (MULL. ARC.) LAMB 289 

-- var. procerum LAMB 296 

-- var. subcorticatum (RAs.) LAMB 290 

- crambidiocephalum LAMB 290 

- cupriniforme NYL. 236, 314 

- curtatoides ASAH. 261 

- curtaturn NYL. 261 

- curtulum NYL. 322

3 50 Joum. Hattori Bot. Lab. No. 43 

- curtum (RAs.) LAMB 269 

- cyrnosum CROMB. 21 1 

- dactylophyllum FLORKE 21 1 

-- f. expansum (MAGN.) GRUMM. 213 

-- var. flabellatum (FREY) GRUMM. 214 

-- var. mujor Som. 212, 310 

-- var. occidentale (MAGN.) GRUMM. 214 

-- var. pygmaeum (FREY) GRUMM. 215 

-- f. sessile LAMB 214 

-- var. spissum (NYL.) LAMB 215 

-- f. subbotryosum (FREY) GRUMM. 214 

- delisei BORY 290 

- dendroides ASAH. 261 

- denudatulum NYL. 322 

- denudatum PERS. 283, 319 

- denudatum FLORKE 244, 3 15 

-- f. bracteata (WALLR.) VAIN. 244 

-- var. caespitosulum NYL. 220 

-- f. capitatum FLOT. 244, 315 

-- f. commune TH. FR. 244, 315 

--- subf. digitatum (LAuR.) TH. FR. 244 

--- subf. tenue (LAuR.) TH. FR. 244 

-- f. compacturn FLOT. 246, 315 

-- var. depressum MAGN. 247 

-- status digitatum LAUR. 244, 31 5 

-- f. digitatum (LAuR.) RABENH. 244 

-- var. genuinum TH. FR. 244, 315 

-- f. nodulosa (WALLR.) VAIN. 244 

-- var. platycladurn LAMB 327 

-- subsp. pulvinatum (SCHAER.) HUE 330 

-- var. pulvinatum (SCHAER.) FLOT. 330 

--- f. arenarium SAV. 250 

--- f. saxicolum SAV. 322 

-- subsp. santorinense J. STEIN. 243 

-- status tenue LAUR. 244, 31 5 

-- f. tenue (LAuR.) LAUR. 244 

-- subsp. turja2nsis RAs. 250 

-- f. umbonata (WALLR.) VAIN. 246 

-- var. umbonatum (WALLR.) ZAHLBR. 246 

--- f. pulvinata (SCHAER.) Ris. 330 

-- var. umbricolum FREY 246 

-- status validum LAUR. 244, 315 

-- var. validum (LAuR.) OLIV. 244 

-- f. validum (LAuR.) LAUR. 244 

-- subsp. vesuvianum (PERs.) TH. FR. 242 

-- var. vesuvianum (PERs.) LAUR. 242 

-- var. vulcani @ORY) NYL. 257 

- depreaultii DEL. 21 5 

-- var. caespitosulum (NYL.) LAMB 220 

-- f. corallinum HUE 21 1 

- depressum (FREY) LAMB 216 

- didymicum LAMB 291 

- dusenii LAMB 292 

- esterhuysenae LAMB 216 

- etigoense (ASAH.) LAMB 262 

- evolutoides (MAGN.) FREY 23 1 

-- f. glabrescens LAMB 23 1 

-- var. paschaleoides HAV. 232 

-- var. saxatile (MAGN.) LAMB 230 

- evolutum GRAEWE 217 

-- var. fastigiatum (ANZI) TH. FR. 207 

-- f. fastigiatum (ANZI) MIG. 207 

-- var. laxum (FREY) LAMB 21 8 

-- f. laxurn (FREY) LAMB 217 

-- f. planum MAGN. 217, 3 11 

-- f. sterile SAV. 234 

-- var. typicum TH. FR. 217, 31 1 

- exalbidum NYL. 285 

- exile ASAH. 262 

- explanaturn LAMB 306 

- exutoides LAMB 264 

- exutum NYL. 260 

-- f. approximans (HUE) DODGE 260 

-- f. cornplanaturn (HUE) DODGE 260 

-- var. humile (MULL. ARG.) DODGE 263 

-- var. nigrum (HUE) SAT6 264 

-- var. sorediata RKs. 270, 31 8 

- farinaceum MAGN. 209, 310 

- farinosum (TH. FR.) VAIN. 297 

- fastigiatum ANZI 207, 309 

-- f. confluens MAGN. 216, 31 1 

-- f. congestum MAGN. 208, 310 

-- f. depressum MAGN. 216 

-- var. dissolutum MAGN. 233 

-- f. finmarkicum MAGN. 208 

-- f. globuliferum MAGN. 234 

-- f. irregulare MAGN. 208 

-- f. simplicior MAGN. 208 

-- f. spathuliferum (VAIN.) MAGN. 233 

- fibrillosum LAMB 251 

-fibula TUCK. 327 

-flavireagens GYELN. 257, 31 7 

-- f. cinerascens MAGN. 257, 317 

-- f. densum MAGN. 257, 317 

- foliiforme HUE 259, 3 17 

- foliolosum Nn. 267 

-- var. botryophorum (MULL. ARG.)~LAMB 

267 

-- var. stricturn (BAB.) LAMB 268 

- formosanum ASAH. 267

-fragile (L.) HOFFM. 327 

- fronduliferum LAMB 293 

- fungiforme SCHAER. 327 

- furcaturn FR. 278, 319 

- furfuraceum DUVIGN. 252 

- giltayi DUVIGN. 257, 31 6 

- glabrum ( M~L. ARG.) VAIN. 21 8 

-- f. flabellans (LAMB) LAMB 218 

- glareosum (SAV.) MAGN. 218 

-- var. brachyphylloides LAMB 219 

-- f. congesturn MAGN. 219, 312 

- glaucescens TUCK. 21 9 

-- var. caespitosulum (NYL.) LAMB 220 

- globiferum HOFFM. 327 

- "globosus (HuDs.) VAIN." 329 

- gracilescens NYL. 327 

- gracilius (MULL. ARG.) DUVIGN. 284 

- gracillimum MULL. ARG. 322 

- graminosum SCHAER. 252 

- grande (MAGN.) MAGN. 220 

- granulosum LAUR. 206, 313 

- gregarium RED. 307 

- groenlandicum (DAHL) LAMB 221 

- halei LAMB 294 

- haumanianum DUVIGN. 296, 320 

- himalayense AWAS, et LAMB 269 

- hokkaidense ASAH. et LAMB 263 

- humbertii DUVIGN. 269 

- humile MULL. ARG. 289 

- hypopetraeum VAIN. 240, 3 15 

- implexum TH. FR. 294 

-- f. sorediosa VAIN. 297, 320 

- incogniturn SCHADE 242 

- incrustatum FLORKE 221 

-- var. abduanum (ANZI) FREY 222 

-- var. elatum OKSN. 222, 312 

-- f. gracile FREY 222 

- intermedium (SAV.) MAGN. 223 

-- f. compactum LAMB 223 

- intricatum MORN 327 

- japonicum TH. FR. 263 

-- var. aogasimense (ASAH.) LAMB 264 

-- var. commixturn ASAH. 260 

-- subsp. etigoense ASAH. 262 

-- var. subfastigiatum ASAH. 263 

-- var. tokioense LAMB 263 

- karisimbiense D ~IGN. 296, 320 

- kotiluotot?nse VAIN. 322 

- laccatum FR. 327 

- laseroni DODGE 327 

I. MACKENZ~E LAMB: A Conspectus of Stereocaulon 

- lavicola MAGN. 252 

- lecanoreum NYL. 305, 321 

- leporinum TH. FR. 206, 313 

-- var. elatum TH. FR. 206, 313 

-- var. pumilum TH. FR. 206, 313 

- leprocauloides LAMB 223 

- leprocephalum VAIN. 253 

- leptaleum NYL. 288, 320 

- loricatum LAMB 307 

- lusitanicum MAGN. 21 7, 31 1 

- macquariense DODGE 283, 323 

- macrocarpoides (NYL.) KREMP. 283 

-macrocarpurn RICH. 282 

-- var. argus (HOOK. F. et TAYL.) TH. FR. 

286 

-- var. curta RAs. 269 

-- f. genuinum TH. FR. 282 

- macrocephalum MULL. ARG. 272 

-- var. stricturn (BAB.) DODGE 268 

-- var. yunnanense (HUE) DODGE 276 

- maderense TUCK. 206, 3 13 

- madreporiforme HOFFM. 327 

- magellanicum (TH. FR.) ZAHLBR. 236 

- mamiNosum DUVIGN. 276, 3 18 

- massartianum HUE 272 

-- var. chlorocarpoides (ZAHLBR.) LAMB 

272 

- Maunae Loae MAGN. 258 

- meissnerianum FLORKE 323 

- melanocarpum (Sw.) RAUSCH. 327 

- melanopotamicum LAMB 296 

- meyeri B. STEIN 296 

-- f. acuminatum (MULL. ARG.) LAMB 297 

-- subsp. atlanticum LAMB 286 

-- var. bornmiilleri B. STEIN 296, 320 

-- var. farinosum (TH. FR.) LAMB 297 

-- var. tucurnanum LAMB 297 

-- f. tucumanum LAMB 297 

- microcarpum MULL. ARG. 297 

- microscopicurn (VILL.) FREY 327 

- microthuja DUVIGN. 253, 316 

-- fac. abietina DUVIGN. 253 

-- fac. fastigiato-pulvinata DUVIGN. 254 

- mikenoense DUVIGN. 277 

- mixtum NYL. 278, 319 

-- subsp. assimile (NYL.) NYL. 302 

-- var. denudatum (PERs.) MULL. ARG. 284 

-- subsp. nesaeum (Nn.) NYL. 272 

-- sorediatum NYL. 305, 321 

-- var. sorediatum (NYL.) HUE 305


-- subsp. strictum (BAB.) NYL. 268 

-- var. tenellum MULL. ARG. 297, 320 

- rnontagneanurn LAMB 274 

- myriocarpoides (NYL.) HUE 224 

- myriocarpum TH. FR. 224 

-- var. altaicum LAMB 225 

-- var. orizabae TH. FR. 224, 312 

-- f. typicum TH. FR. 224 

- nabewariense ZAHLBR. 244, 31 5 

- nanodes TUCK. 240 

-- f. carinthiacum (FREY) LAMB 241 

-- f. schadeanum LAMB 242 

-- f. tyroliense (NYL.) LAMB 240 

- nanum (AcH.) ACH. 327 

-- f. mundum FR. 323 

-- f. pulverulenturn TH. FR. 327 

- nesaeum NYL. 273, 318 

- nesaeum Auctt. (non NYL.) 272, 273 

-- var. lecideoides VAIN. 272, 3 18 

-- var. zeorina VAIN. 272, 318 

- nigromaculatum DUVIGN. 253 

-- f. fastigiato-pulvinatum LAMB 254 

- nigrum HUE 264 

- novo-arbuscula ASAH. 328 

- novogranatense LAMB 298 

- obesum TH. FR. 254 

-- var. egorescens RAS. 248 

- obscurutn MULL. ARG. 249 

- obtusatum (VAHL) ACH. 328 

- octomerelloides ASAH. 264, 3 18 

- octomerellum MULL. ARG. 264 

-- var. sandwicense (MAGN.) LAMB 264 

- octomerum MULL. ARC. 265 

-- f. robustior ASAH. 262, 31 8 

- orizabae (TH. FR.) VAIN. 224 

- ostrobottniae MAGN. 241, 315 

- pachycephalum VAIN. 299 

- papuanum LAMB 225 

- paradoxum LAMB 275 

-parvifolia 329 

- paschale (L.) HOFFM. 200, 307 

-- f. alpinum (LAuR.) MUDD 202 

-- var. alpinum (LAuR.) DU RIETZ 202 

--- f. flabellans LAMB 218 

-- var. cereolinum (AcH.) SCHAER. 328 

-- f. colligatum LAMB 200, 309 

-- var. condensatum (HOFFM.) SCHAER. 209 

-- f. congestum LAMB 200 

-- p. conglomeratum FR. 329 

--- 1. robustum FLOT. 329 

-- E. conglomeratum (FR.) TH. FR. 329 

-- subsp. coralloides (FR.) BRANTH et 

ROSTR. 212 . 

-- subsp. dactylophyllum (FLORKE) BRANTH 

et Rosm. 211 

-- var. denudatum (FLORKE) SCHAER. 244 

-- f. denudatum (FL~RKE) MUDD 244 

-- subsp. depreaultii (DEL.) LAMB 215 

--- var. caespitosulum (NYL.) LAMB 220 

-- var. erectum (FREY) LAMB 203 

-- var. evolutoides MAGN. 231, 313 

-- subsp. evolutoides (MAGN.) LAMB 231 

--- f. laxatum LAMB 231, 313 

-- var. gracilenturn TH. FR. 204 

-- f. gracilentum (TH. FR.) TH. FR. 204 

-- var. grande MAGN. 220 

-- var. magellanicum (TH. FR.) NYL. 236 

-- f. paschale (L.) MUDD 200 

-- var. pileatum (AcH.) WAHLENB. 226 

-- var. pulvinatum SCHAER. 330 

-- f. ramulferum NYL. 219, 311 

-- var. serpens TH. FR. 200, 308 

-- f. serpens (TH. FR.) TH. FR. 200 

-- f. sorediatum MAGN. 232 

-- var. squamescens (NYL.) RAS. 323 

-- f. squamescens NYL. 323 

-- f. subcoralloides NYL. 235 

-- f. subcrustosum FR. 324 

-- f. taeniarum MAGN. 231, 313 

-- f. thyrsoideum (SCHAER.) RABENH. 202 

-- subsp. tomentosum (FR.) BRANTH et 

ROSTR. 236 

-- var. tomentosum (FR.) DUBY 236 

-- f. typicum Du RIETZ 200, 308 

-- f. velutinum FREY 324 

-- var. vulgare TH. FR. 200, 308 

-- f. vulgare (TH. FR.) TH. FR. 200 

-patagonicum LAMB 218, 31 1 

-- f. reagens (RiCs.) LAMB 21 8 

-- f. subirregulare LAMB 21 8, 3 1 1 

- peladense VAIN. 305 

- pendulum ASAH. 265 

-penicillium DUVIGN. 296, 320 

- philippinense RAs. 299 

- pileatum ACH. 226 

-- f. macrum MAGN. 226, 312 

-- var. nipponicum LAMB 227 

-- f. pumilum (NYL.) LAMB 227 

-- f. ramificans MAGN. 227, 313 

-- f. sessile MAGN. 246

-- var. sorediiferum NYL. 226, 312 

-- f. sorediiferum (Nn.) HARM. 226 

--- subf. terrestre HARM. 226, 330 

-pilophoroides TUCK. 283 

- piluliferum TH. FR. 275 

-- var. sinense (HUE) LAMB 275 

- pityrizans NYL. 254 

- platycladum (LAMB) LAMB 327 

- porniferum DUVIGN. 276 

-- f. mikenoense (DUVIGN.) LAMB 277 

- procerum GROENH. 277 

- prostratum ZAHLBR. 266 

- proximum NYL. 278, 319 

-- var. compactius ZAHLBR. 294, 320 

-- var. compressurn NYL. 306 

-- subsp. exalbidum (NYL.) NYL. 285 

-- status ferruginascens MULL. ARG. 297, 

320 

-- var. gracilius MULL. ARG. 284 

-- subsp. macrocarpoides NYL. 282, 319 

-- var. macrocarpoides (NYL.) NYL. 282 

-- var. nudatum M ~L. ARG. 281 

-- f. sorediata NYL. 296, 324 

-- f. traversii HUE 28 1, 3 19 

-prrrinosum FL~~RKE 330 

- pseudoarbuscula ASAH. 328 

- pseudodepreaultii ASAH. 228 

- pseudomassartianum LAMB 300 

-pseudosasakii ASAH. 223 

- pulvinare DODGE 283 

- pulvinatum ACH. 328 

-pulvinatum (SCHAER.) NYL. 330 

- pygmaeum VAIN. 328 

- quisquiliare (LEERS) HOFTM. 328 

-- f. pulverulentum (TH. FR.) ZAHLBR. 328 

- ramulosum (Sw.) RAUSCH. 278 

-- var. acuminatum MULL. ARG. 297 

-- var. assimile (NYL.) LAMB 302 

-- var. bornmulleri (B. STEIN) MULL. ARG. 

296 

-- subsp. claviceps (TH. FR.) NYL. 271 

-- var. compacturn MULL. ARG. 278, 319 

-- var. compressurn BAB. 293, 320 

-- f. compressum NYL. 306, 321 

-- f. elegans TH. FR. 281 

-- var. exalbidum (NYL.) LAMB 285 

-- var. farinosum TH. FR. 297 

-- f. farinosum (TH. FR.) HUE 297 

-- var. furcaturn (FR.) NYL. 278 

-- f. genuinum TH. FR. 278 


-- var. gracilius (MULL. ARG.) LAMB 284 

-- var. implexum (TH. FR.) NYL. 294 

-- var. macrocarpoides (NYL.) MULL. ARG. 

283 

-- subsp. macrocarpum (RICH.) HUE 282 

--- f. compressurn (BAB.) HUE 293 

-- var. macrocarpum (RICH.) BAB. 282 

-- var. microcarpoides MULL. ARG. 278, 

319 

-- var. nudatum (MULL. ARG.) MULL. ARC. 

28 1 

-- f. nudatum (MULL. ARG.) LAMB 281 

-- var. perpumilum LAMB 284 

-- var. pilophoroides (TUCK.) LAMB 283 

-- f. proximum (NYL.) TUCK. 278 

-- var. pulvinare (DODGE) LAMB 283 

--- f. crebratum LAMB 283 

-- subsp. roccelloides TH. FR. 300, 321 

-- var. simplicius LAMB 284 

-- var. stricturn BAB. 268 

-- var. strigosum (TH. FR.) HUE 278 

-- f. strigosum TH. FR. 278, 319 

-- f. subcompressum LAMB 282 

-- var. sublecanoreum LAMB 279 

-- subsp. submollescens Nn. 285 

-- var. submollescens (NYL.) LAMB 285 

-- f. tomentosulum LAMB 282 

-- var. vimineum (TH. FR.) NYL. 279 

-- f. vimineum (TH. FR.) HUE 279 

- richardianum (MoNT.) ZAHLBR. 255 

- rivulorum MAGN. 228 

-- var. groenlandicum DAHL 221 

-- f. nana CENGIA SAMBO 324 

-- f. pulvinarium (SAV.) LAMB 229 

- roccella (L.) Rjiusc~. 328 

- roccelloides (TH. FR.) LAMB 300 

- rubiginosum PERS. 300 

- rugulosum LAMB 301 

- ruwenzoriense DUVIGN. 253, 316 

- salazinum (BORY) F~E 302 

- sandwicense MAGN. 264, 318 

- sanguineum DEL. 300 

- sasakii ZAHLBR. 229 

-- var. simplex (RIDD.) LAMB 230 

-- var. tomentosoides LAMB 230 

- saviczii Du RIETZ 266 

- saxatile MAGN. 230 

-- f. paschaleoides (HAv.) LAMB 232 

-- f. sorediatum (MAGN.) LAMB 232 

- saxonicum BACHM. 226, 31 2


- scutelligerum TH. FR. 254 

- sertularia ACH. 328 

- sibiricum LAMB 232 

- sinense HUE 275, 318 

- soleirolii DUF. 328 

- solutum LAMB 324 

- sorediatum (NYL.) VAIN. 296, 324 

- sorediiferum HUE 270 

-- f. esorediosum ASAH. 270 

-- var. leprosolingulatum LAMB 271 

- spathuliferum VAIN. 233 

-- f. dissolutum (MAGN.) LAMB 233 

-- f. globuliferum (MAGN.) LAMB 234 

-- f. pygmaeum (MAGN.) LAMB 234 

- speciosum LAMB 256, 316 

-- var. surreptans LAMB 256 

- sphaerophoroides TUCK. 206, 257 

-- f. elatum (TH. FR.) TH. FR. 206 

-- f. pumilum (TH. FR.) TH. FR. 206 

- spissum NYL. 214, 31 1 

-- f. dactylinum FREY 21 5, 31 1 

-- f. granuliferum FREY 324 

-- var. laxum FREY 217 

- squamescens (NYL.) FUs. 323 

- stalagrnitiferum DWIGN. 272, 31 8 

- staufferi LAMB 303 

-- var. bomdnse LAMB 304 

- stenospermum (LAMB) LAMB 304 

- sterile (SAV.) LAMB 234 

- strictum (BAB.) MULL. ARG. 268 

- strictum TH. FR. 305 

-- var. compressum (Nn.) LAMB 306 

-- var. lecanoreum (NYL.) LAMB 305 

- subalbicans LAMB 328 

- subcoralloides (Nn.) NYL. 234 

-- f. pumilum (NYL.) ZAHLBR. 227 

-- f. sorediascens LAMB 235 

- subcorticatum Us. 290 

- subdenudatum HAV. 255 

- subgracilescens LAMB 328 

- subintricans NYL. 216, 324 

- submollescens (NYL.) ZAHLBR. 285 

- subramulosum MULL. ARG. 260, 317 

-- f. approximans HUE 260, 317 

-- f. complanatum HUE 260, 317 

-- var. humile MULL. ARG. 263, 317 

- substrictum HUE 305 

- supervestiens MAGN. 235 

- symphycheilum LAMB 255 

- tabulare (THIJNB.) ACH. 328 

- tenellum TUCK. 328 

- tenerrimum LAMB 262 

- tennesseense MAGN. 236 

-- var. nigrofastigiatum LAMB 236 

- tenuissimum DEL. 325 

- togashii LAMB 278 

- tomentosum FR. 236 

-- var. abduanum (ANZI) OLIV. 222 

-- var. alpestre FLOT. 238 

--- f. stigmatea FLOT. 330 

-- subsp. alpestre (FLoT.) HUE 238 

-- f. alpestre (FLoT.) NYL. 238 

-- var. alpinum (LAuR.) TH. FR. 202 

-- subsp. alpinum (LAuR.) TH. FR. 202 

--- f. verrucosum TH. FR. 202, 309 

-- var. azoreum SCHAER. 205 

-- f. botryocarpum (MAGN.) LAMB 238 

-- var. botryosum (AcH.) NYL. 207 

-- var. campestre KijRB. 236, 314 

-- var. capitatum LAMB 239 

-- var. compactum FREY 239 

-- var. cupriniforme (NYL.) OLIV. 236 

-- f. cupriniforme (NYL.) VAIN. 236 

-- f. flabelliforme OHL. 325 

-- f. glareosum SAV. 219 

-- var. granulosum SCHAER. 206, 331 

-- var. granulosum (LAuR.) OLIV. 206 

-- var. graphiticola US. 231, 313 

-- var. hercynicum FREY 325 

-- var. inciso-crenatum SCHAER. 236, 314 

-- f. incisocrenatum (SCHAER.) FREY 236 

-- subsp. incrustatum (FL~RKE) NYL. 221 

-- var. incrustatum (FLORKE) SCHAER. 221 

-- var. karelica VAIN. 330 

-- f. Ieptothamnium FREY 325 

-- subsp. magellanicum TH. FR. 236, 314 

-- B. rnajus SCHAER. 330 

-- subsp. myriocarpoides NYL. 224, 312 

-- subsp. myriocarpum (TH. FR.) NYL. 224 

--- var. orizabae (TH. FR.) LAMB 224 

-- var. simplex RIDD. 230 

-- f. tectorum TOM. 239, 314 

-- var. thyrsoideum SCHAER. 202, 309 

-- f. truncatum FREY 237, 314 

-- var. tyroliense (NYL.) OLIV. 240 

-- var. walamoCnse (Nn.) NYL. 325 

-- f. walamot?nse NYL. 325 

- tortuosum NYL. 216, 326 

- trachyphloeum LAMB 304 

- turfosum BORY 326

- turgescens Nn. 252 

- tyroliense (NYL.) LETT. 240 

-- var. lapponicum MAGN. 240, 314 

- uliginosum LAMB 240 

- uvuliferum MULL. ARG. 263, 3 17 

- valamo5nse (Nn.) RXs. 325 

- vanoyei DUVIGN. 256 

- vermiculare (Sw.) KUSCH. 328 

- verruciferum NYL. 256 

-- var. surreptans (LAMB) LAMB 256 

- verruculigerum HUE 267 

-- var. formosanum ASAH. 267 

-- var. subfurfurascens LAMB 267 

- vesuvianum PERS. 242 . 

-- var. arcticum (LYNGE) LAMB 249 

-- var. capitatum PLOT.) JAMES 244 

-- var. confluens (MULL. ARG.) B. STEIN 

248 

-- var. denudatum (FL~~RKE) LAMB 244 

--- f. capitatum (FLoT.) POELT 244 

-- var. depressum (MAGN.) LAMB 247 

-- var. emorescens (Ks.) LAMB 248 

-- var. glaucescens (TUCK.) LAMB 219 

-- var. kilimandscharoense B. STEIN 248 

-- f. nabewariense (ZAHLBR.) ASAH. 244 

-- var. nodulosum (WALLR.) LAMB 244 

--- f. depressum (MAGN.) LAMB 247 

--- f. sessile (MAGN.) LAMB 246 

--- f. umbonatum (WALLR.) LAMB 246 

--- f. umbncolum (FREY) LAMB 246 

--- f. verrucosum LAMB 247 

A Conspectus of Stereocaulon 355 

-- var. obscurum (MULL. ARG.) LAMB 249 

-- var. pulvinatum (SCHAER.) JAMES 330 

-- f. santonnense (J. STEIN.) LAMB 243 

-- var. surugaense ASAH. 248 

-- var. symphycheileoides LAMB 247 

-- f. tenue (LAuR.) OKSN. 244 

-- var. thyrsoideum LAMB 249 

-- var. umbonatum (WALLR.) LAMB 246 

-- var. umbricolum (FREY) LAMB 246 

-- var. violascens (MULL. ARG.) LAMB 248 

- vimineum TH. FR. 279,3 19 

- violascens MULL. ARG. 248, 3 16 

- virgatum ACH. 257 

-- f. achariana VAIN. 257, 316 

-- f. applanata VAIN. 257, 316 

-- f. primaria VAIN. 257, 316 

- vulcani (BORY) ACH. 257 

-- f. maunae-loae (MAGN.) LAMB 258 

-- subsp. richardianum MONT. 254, 316 

- wadei LAMB 305 

- weberi LAMB 258 

- wrightii TUCK. 259 

- yunnanense (HUE) LAMB 276 

Stereocladium Nn. 195 

- apocalypticum (NYL.) NYL. 259 

- saxonicum (BACHM.) SCHADE, STOLLE et 

RIEHM. 226 

- tyroliense Nn. 240, 3 14 

- wrightii (TUCK.) Nn. 259 

Verrucaria paschalis (L.) HUMB. 200

A CONSPECTUS OF THE LICHEN GENUS STEREOCAULON ...

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