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A review of the genus Hypogymnia (Parmeliaceae) in ChileAuthor(s) :Arve ElvebakkSource: The Bryologist, 114(2):379-388. 2011.Published By: The American Bryological and Lichenological Society, Inc.DOI: http://dx.doi.org/10.1639/0007-2745-114.2.379URL: http://www.bioone.org/doi/full/10.1639/0007-2745-114.2.379

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A review of the genus Hypogymnia (Parmeliaceae) in Chile

Arve Elvebakk1

Tromsø University Museum, University of Tromsø, N-9037 Tromsø, Norway

ABSTRACT. Three sorediate and four primarily fertile species of the lichen genus Hypogymnia are

accepted for Chile in the present study, and only two of these correspond to species listed in

current checklists. Hypogymnia antarctica, H. lugubris and H. subphysodes are considered to be

common; the former has previously been poorly understood. Hypogymnia pulverata and H. bitteri

are reported here as new to Chile, the latter also as new to austral areas. The former is not

convincingly well separated from H. subphysodes. Hypogymnia enteromorphoides and H.

solidepedicellata are also accepted here, based on literature reports, whereas six species reported

rather recently from southern South America are not accepted here for this area. Sorediate species

are much more widely distributed than primarily fertile ones, not only in the present study area,

but also on a global scale. Vittatolic, 2’-O-methylphysodic, and 3-hydroxyphysodic acids were

found to be reliable differential compounds between some of the species studied. The superficially

similar genera Hypogymnia and Menegazzia are of strong biogeographical interest, as they show

parallel distribution patterns, although they have probably originated in different hemispheres.

KEYWORDS. taxonomy, distribution, lichens, biogeography, Southern Hemisphere.

¤ ¤ ¤

The lichen genus Hypogymnia consists of foliose to

fruticose species, mostly with inflated, hollow lobes,

which have a black lower side devoid of rhizines.

Apothecia are rare in many species, common in

others, and are lecanorine, stipitate, with concave,

red-brown discs. While most species are grey due to

the presence of atranorin in their cortex, a few are

yellowish from usnic acid or brownish from

melanins. Members of Hypogymnia resemble species

of two other genera with hollow lobes. Cavernularia

is an oceanic Northern Hemisphere genus of two

small species with pitted lower sides, and Menegazzia

is a very species-rich genus in panaustral areas,

mostly with conspicuous perforations through the

epicortex. These genera are sometimes

accommodated within the Hypogymniaceae (e.g. Elix

1992; Karnefelt et al. 1992), but this concept has been

abandoned, and these taxa are now included in the

Parmeliaceae. Although rather few samples of these

genera have been analyzed phylogenetically, Crespo

et al. (2007), defines a hypogymnioid group,

comprising Hypogymnia and some other genera and

species with loose medullas, whereas Menegazzia

remains isolated and monophyletic, although weakly

associated with Cetrariella delisei (Bory ex Schaer.)

Karnefelt & Thell in a later study (Crespo et al. 2010).

The genus Hypogymnia has its highest

biodiversity in the Northern Hemisphere, e.g. with 42

species known from China (Wei & Wei 2005) and 31

species from North America (Esslinger 2010; H.

lugubris (Pers.) Krog, now a pan-austral species,

excluded). The total number of species was estimated

as ‘c. 86’ a few years ago by B. McCune, cited by

1 Author e-mail: arve.elvebakk@uit.no

DOI: 10.1639/0007-2745-114.2.379

The Bryologist 114(2), pp. 379–388 0007-2745/11/$1.15/0Copyright E2011 by The American Bryological and Lichenological Society, Inc.

Galloway (2007), and should by now be close to 100.

Only a few species are known from tropical Asian

mountains, and its species diversity increases in the

Southern Hemisphere, where an austral element

dominates. At present, five species are accepted from

Chile (Feuerer 2009), and seven from Argentina

(Calvelo & Liberatore 2002). However, only two of

these are in common. During own fieldwork in Chile

focusing on the lichen family Pannariaecae, some

collections of a Hypogymnia species new to austral

areas were also made. The aim of the present paper is

to report on the occurrence of Hypogymnia species in

Chile based on collections housed in TROM, review the

diverging reports on the genus from southern South

America in the literature and to include the species in

a phytogeographical discussion.

MATERIALS AND METHODS

The collections in the herbarium TROM were

examined, and the descriptions are based on this

material. All specimens were studied by thin-layer

chromatography of acetone extracts using

standardized procedures and solvents A and C

(Culberson 1972; Orange et al. 2001). The samples

were run on Merck TLC Silica Gel 60 F254 plates and

the identification also involved the use of a Camag

UV lamp, operating at 366 nm. A total number of

138 samples were analyzed, including references from

other areas, and six reference species not present in

the study area. A number of samples were analyzed

more than once, to obtain reliable interpretations.

Several unidentified compounds with scattered

occurrences and terpenoids in low quantities were

not included among the results.

RESULTS

Hypogymnia antarctica (Bitter) Dodge Fig. 1

Morphology. Thallus foliose, 3–10 cm wide,

corticolous, appressed to the surface. Central lobes

contiguous, 1–4 mm wide; marginal lobes mostly

discrete and ramified, 1–2 mm broad, mostly

dichotomously branched; all lobes hollow. Upper

surface glossy and rugose, grey, young lobe tips

brown. Lower surface black, glossy, strongly faveolate

and wrinkled, without perforations, extending up

along margins of the upper surface. Apothecia very

common, often in groups, large and up to 15 mm

broad, stipitate and 5 mm tall; pedicel urn-shaped

when young, funnel-shaped when old, faveolate and

with distinct longitudinal furrows; thalline

excipulum thin, c. 0.2 mm wide, even. Disc dark

brown, becoming flattened in the largest apothecia.

Ascospores hyaline, non-septate and ellipsoid, 6.5–

8.5 3 4.5–6.0 mm in size. Pycnidia common, forming

numerous black dots on the upper surface.

Chemistry (Fig. 2). Chemotype I: atranorin

(incl. chloroatranorin) (trace to major), physodic, 3-

hydroxyphysodic acids (major), 29-O-

methylphysodic (minor). Chemotype II: atranorin

(incl. chloroatranorin) (trace to major), physodic, 3-

hydroxyphysodic and physodalic acids (major), 29-

O-methylphysodic and protocetraric acids (trace/

minor).

Distribution. The species has been collected

from latitudes 38 to 53 uS in Chile. It is particularly

common in very humid areas in regions XI and XII

in Chile, and in places the species can be dominant

on smooth bark of light-exposed trees, particularly

near bogs.

Discussion. The species was described based on

its holotype specimen only, which was collected by

Lechler at ‘Sandy Point’ (5 Punta Arenas). There are

scattered reports in the literature, but the species has

not been studied since its description, except for the

following information. Although Elix (1979) did not

discuss the species in his monograph on Hypogymnia

in Australasia, he obviously considered it different

from his newly described species. Moreover, he

reported its type at BM to contain atranorin/

Figure 1. Hypogymnia antarctica (Bitter) Dodge, Elvebakk

99:758A.

380 The Bryologist 114(2): 2011

chloroatranorin, physodic, 3-hydroxyphysodic, 29-O-

methylphysodic and alectorialic acids. Calvelo (1998)

included the species in a key of four Hypogymnia

species in Argentina. Goward & McCune (2007)

briefly indicated that H. antarctica belongs in a

widely distributed group named after H.

metaphysodes (Asah.) Rass. Goward et al. (2010)

revised H. metaphysodes in North America and

suggested that H. antarctica instead has affinities with

the H. austerodes group, due to a white medullary

ceiling and black mottling on the upper surface. They

also cited three collections of H. antarctica.

The material studied here shows that specimens,

which contain the physodic acid complex and those

which in addition contain the physodalic acid

complex are morphologically similar. Thus,

Hypogymnia antarctica is chemically a parallel to H.

enteromorphoides, in including two similar

chemotypes. However, the central lobes of the former

are less robust, lower side perforations are lacking,

and the upper surface is distinctly rugose, all

characters agreeing with the thorough description of

H. antarctica by Bitter (1901). Morphologically, it

appears to be most similar to H. australica Elix,

which also has urn-shaped pedicels and narrow lobes

(Elix & Jenkins 1989). However, H. antarctica has a

rugose upper surface, white medullary ceiling, black

upper side margins, larger apothecia and apothecia

often in groups. No species resembling H. antarctica

have been described with the physodalic acid

complex. Concerning the upper surface, it is

concluded here that it rarely has a black mottling,

apart from the black margins and dots around the

pycnidia. The character separating H. antarctica and

H. lugubris in the key by Calvelo (1998) is the

presence of black mottling in the latter and absence

in the former. This view is supported here.

Selected specimens examined. Chemotype I

(from a total of 8). CHILE: IX REGION DE LOS LAGOS,

Provincia de Llanquihue, Parque Nacional Vicente

Perez Rosales, southern slope of Volcan Osorno,

41u069S, 72u279W, 780 m, Bjerke 676/01; XII REGION DE

MAGALLANES Y DE LA ANTARTICA CHILENA, Provincia de la

Ultima Esperanza, Parque Nacional Torres del Paine,

1.5 km SE of Cascada Pingo, 51u049S, 73u139W,

150 m, Elvebakk 98:550 & Bjerke. Chemotype II

(from a total of 11). CHILE: IX REGION DE LA ARAUCANıA,

1.2 km S of SE boundary of Reserva Nacional

Malalcahuello, 38u249S, 71u309W, 1400 m, Elvebakk

07:383; Parque Nacional Vicente Perez, lago Todos

los Santos, Petrohue, 41u089S, 72u249W, 70 m,

Elvebakk 00:584; XI REGION AISEN DEL GENERAL CARLOS

IBANEZ DEL CAMPO, Provincia de Aisen, lago Riesco near

Puerto Aisen, 45u309S, 72u419W, Elvebakk 06:425;

Parque Nacional Queulat, Rıo Ventisquero, 1 km E of

Carretera Austral, 44u289S, 72u349W, 20 m, Elvebakk

06:534A; XII REGION DE MAGALLANES Y DE LA ANTARTICA

CHILENA, Provincia de la Ultima Esperanza, Puerto

Eden, 49u089S, 74u249W, Tømmervik s.n.; Provincia de

Magallanes, Laguna el Parrillar, 40 km SW of Punta

Arenas, 53u229S, 71u179W, 300 m, Elvebakk 99:758A.

Hypogymnia bitteri (Lynge) Ahti

Nomenclatural note: Parmelia physodes var.

obscurata Ach. (Acharius 1814) is a synonym of

Parmelia austerodes Nyl., now known as Hypogymnia

austerodes (Nyl.) Ras., a neighbouring species of H.

bitteri. Bitter (1901) introduced the names Parmelia

obscurata (Ach.) Bitter and Parmelia farinacea var.

obscurascens Bitter, for what is now these two

Figure 2. Chromatogram showing TLC-detectable com-

pounds in solvent A in I) Hypogymnia antarctica (Bitter)

Dodge, Chemotype II (n512), II) H. antarctica (Bitter) Dodge,

Chemotype I (n58), H. austerodes (Nyl.) Rasanen (n520). H.

pulverata (Nyl. ex Cromb.) Elix (n56), H. subphysodes

(Kremp.) Filson (n513) and H. tubulosa (Schaer.) Hav.

(n54), III) H. vittata (Ach.) Parrique (n54), IV) H. bitteri

(Lynge) Ahti (n532). Compound abbreviations: ‘at’ 5

atranorin, ‘2-O’ 5 29-O-methylphysodic acid, ‘pdi’ 5 physodic

acid, ‘3-h’ 5 3-hydroxyphysodic acid, ‘pda’ 5 physodalic acid,

‘pc’ 5 protocetraric acid, ‘v’ 5 vittatolic acid, and as additional

references ‘ns’ 5 norstictic acid, ‘cs’ 5 connorstictic acid, u 5

usnic acid, ‘al’ 5 alectorialic acid.

Elvebakk: Hypogymnia in Chile 381

Hypogymnia species. However, he mixed them up, so

that Parmelia obscurata became a heterotypic

synonym of H. austerodes instead. This was pointed

out by Lynge (1921), who therefore renamed Bitter’s

Parmelia farinacea var. obscurascens as Parmelia

bitteri (Lynge), a species later transferred to

Hypogymnia by Ahti (1964).

For illustrations see http://www.nhm.uio.no/

botanisk/lav/ and Brodo et al. (2001: 348).

Morphology. Thallus 3–10 cm wide, corticolous,

rarely saxicolous; lobes appressed to the substrate,

flattened, 1–2 mm wide, inflated, irregularly divided

and contiguous in central parts. Upper surface

glabrous and glossy, maculate in young parts, weakly

rugose in older parts, greenish grey to brown, brown

pigment dominant or only present in young lobe

tips, lower surface black. Black margins visible from

above, particularly on light-exposed specimens.

Soralia capitate, farinose, developing along lobe

margins or on short, erect secondary lobules.

Apothecia rare, not seen in Chilean material.

Chemistry (Fig. 2). Atranorin (incl.

chloroatranorin) (trace to major), physodic and

vittatolic acids (majors).

Distribution. New to Chile and to austral areas.

Hypogymnia bitteri is a very distinct species and

represents the only documented report of a Northern

Hemisphere Hypogymnia species occurring in austral

areas. In Chile this species extends further south than

the other sorediate species and is very common in

central parts of Torres del Paine National Park. Its

occurrence on rocks in Morro Chico close to the

Argentina boundary in central parts of the

Magellanic region is remarkable. This predominantly

treeless area has mostly been referred to as

‘Patagonian steppe’. Although it was characterized as

‘antiboreal Chiliotrichum-Empetrum heathland’ by

Elvebakk & Moberg (2002), as opposed to the drier

and warmer temperate steppe present 30–40 km

away, this is still a dry zone very different from the

moist forests. Also the localities near Malalcahuello

National Reserve further north in Region IX are in

dry forests, here dominated by Araucaria araucana.

Discussion. In shaded habitats it lacks the strong

brown pigment which dominates in light-exposed

sites. However, even in shaded environments young

lobe tips are always brown.

Hypogymnia bitteri is otherwise widespread, and

locally common, although mostly scattered in the

Northern Hemisphere. It is strongly northern in

North America, but also occurs further south in arid

areas, and extends into Mexico (Brodo et al. 2001;

Thomson 1984). It has isolated occurrences in

Colombia and Venezuela (Feuerer 2009). It has a

parallel pattern in the Old World, widespread in

northern, continental areas, but extending

southwards into countries such as Morocco, Turkey,

Ethiopia, Kenya and Uganda (Feuerer 2009). The

pattern is shared by H. austerodes, the other

widespread species in the group. The third species of

this group, H. subobscura (Vainio) Poelt, is restricted to

dry and cold areas, particularly in the High Arctic.

Another three melanin-brown species, also terricolous

and saxicolous, have similar ecological deviations from

most other Hypogymnia species. Hypogymnia castanea

McCune & Krog and H. fistulosa McCune & Krog are

arctic species (McCune 2008) and H. kosciuscoensis Elix

is high-alpine in Australia and New Zealand. From a

habitat and ecological point of view, these six species,

as opposed to H. antarctica, could be hypothesized to

form the H. austerodes group, although McCune

(2008) suggested that the resemblance of some of them

may be superficial.

Selected specimens examined (from a total

of 14). CHILE: IX REGION DE LA ARAUCANıA, Provincia de

Malleco, near SE boundary of Reserva Nacional

Malalcahuello, near Rıo Cautın, 38u269500 S,

71u319W, 1180 m, Elvebakk 09:158; 09:188; XI REGION

AISEN DEL GENERAL CARLOS IBANEZ DEL CAMPO, Cochrane,

Reserva Nacional Tamango, Quilhot 3001; XII REGION

DE MAGALLANES Y DE LA ANTARTICA CHILENA: Provincia de

la Ultima Esperanza, Parque Nacional Torres de

Paine; 1 km SW of Refugio Zapata, 51u049300S,

73u169W, 170 m, Elvebakk 98:269 ; Provincia

Magallanes, Morro Chico, 52u039S, 71u249W, 200 m,

Elvebakk 99:1235.

Hypogymnia enteromorphoides Elix

The species was described from Australia and

New Zealand by Elix (1979). Elix (1992) later

reported it from southernmost South America with

no further information, except the inclusion of an

older synomym at variety level, collected from

Hogget Bay, Tierra del Fuego. No own specimens

could with certainty be identified as this species.

382 The Bryologist 114(2): 2011

Hypogymnia lugubris (Pers.) Krog

A very common species primarily on the ground

in steppes and bogs, partly also on twigs, in

southernmost South America. Four Chilean samples

studied chemically showed the same compounds as

in chemosyndrome II of H. antarctica, except 29-O-

methylphysodic acid being absent, and 3-

hydroxyphysodic acid lacking from three of the

samples.

Hypogymnia pulverata (Nyl. ex Cromb.) Elix

For synonyms, illustrations and detailed

description, see Elix (1979).

Morphology. Thallus 3–15 cm wide, corticolous;

primary lobes appressed, flattened and 2–3 mm wide,

developing ‘secondary’ lobes which are more or less

ascending, sparingly branched, elongated, only 1–

2 mm wide, flattened, rigid from a well-developed

medulla, and mostly without a central cavity. Upper

surface greenish grey without brown melanins,

smooth, glossy in young parts, lower surface black,

irregularly pitted, brown near apices. Soredia farinose

and laminal, starting in distal parts of ascending

lobes, gradually becoming diffuse and covering large

parts of the lobes. Apothecia not seen in Chilean

material.

Chemistry (Fig. 2). Atranorin (incl.

chloroatranorin) (trace to major), physodic and 3-

hydroxyphysodic acids (majors), 29-O-

methylphysodic acid (trace/minor). In addition,

physodalic acid is present in the southernmost

specimen.

Distribution. New to Chile. Hypogymnia

pulverata is otherwise common in southeastern

Australia and in New Zealand, and is also known

from Japan, China and easternmost Russia (Elix

1979, Galloway 2007). The species is very rare in

North America and is known from only four coastal

sites (McCune & Geiser 2009). The species was

reported from a single locality in the Argentinean

province of Rıo Negro by Calvelo & Lorenzo (1989).

The Argentinean province of Chubut was added in

the checklist by Calvelo & Liberatore (2002)

Specimens examined. CHILE: IV REGION DE

COQUIMBO, Provincia de Limarı, Parque Nacional

Fray Jorge, 30u409S, 71u409W, c. 500 m, Elvebakk

00:417 ; IX REGION DE LA ARAUCANıA, Provincia de

Malleco, Parque Nacional Nahuelbuta, 0.5 km N of

Centro de Informaciones, 37u509S, 73u019W,

1300 m, Bjerke 980/01 ; X REGION DE LOS LAGOS,

Provincia de Llanquihue, 15 km W of Puerto

Montt, Puente Taylor, 41u319S, 73u049W, 70 m,

Elvebakk 00:422 ; Parque Nacional Vicente Perez

Rosales, S slope of Volcan Osorno, 41u079S,

72u259W, 315 m, Bjerke 649/01 ; XII REGION DE

MAGALLANES Y DE LA ANTARTICA CHILENA, Puerto Eden,

49u089S, 74u249W, Tømmervik s.n.

Hypogymnia solidepedicellata (Bitter) Dodge

The species was described from Punta Arenas

(Bitter 1901), based on the holotype specimen only.

It has later been reported from Isla Navarino by Du

Rietz (1926) and from Punta Arenas and Rıo Grande

(Argentina) by Zahlbruckner (1917). Elix (1979)

discussed his new species Hypogymnia turgidula Elix

vs. H. solidepedicellata, particular in view of the solid

apothecium pedicel of the latter. He also reported its

holotype to contain atranorin/chloroatranorin,

physodic, 3-hydroxyphysodic, and 29-O-

methylphysodic acids. No own specimens studied

here. For distinguishing features, see the key below.

Hypogymnia subphysodes (Kremp.) Filson

For synonyms, illustrations and detailed

description, see Elix (1979).

5 Hypogymnia subphysodes (Kremp.) Follmann &

Redon, Willdenowia 6: 441, 1972. nom. inval.,

later homonym.

Morphology. Thallus 3–15 cm wide, corticolous;

primary lobes appressed, flattened and 2–3 mm wide,

developing ‘secondary’ lobes which are more or less

ascending, sparingly branched, elongated, only 1–

2 mm wide, isodiametric in cross-section, brittle

from the lack of a well-defined medulla and mostly

with a central cavity. Proportions of primary vs.

secondary lobes variable. Upper surface greenish grey

without brown melanins, smooth, glossy in young

parts; lower surface black, irregularly pitted, brown

near apices. Soredia farinose and laminal, starting in

distal parts of ascending lobes, gradually becoming

diffuse and covering large parts of the lobes.

Apothecia rare, disc chestnut brown up to 5 mm

wide, stipitate with a ribbed, funnel-shaped

receptacle which is sorediate in upper parts, spores 8,

hyaline, ellipsoid, 7–10 3 4–6 mm.

Elvebakk: Hypogymnia in Chile 383

Chemistry (Fig. 2). Atranorin (incl.

chloroatranorin) (trace to major), physodic and 3-

hydroxyphysodic acids (majors), 29-O-

methylphysodic acid (trace/minor).

Distribution. Hypogymnia subphysodes is

‘common in South America’ (Elix 1979), but only

one locality from Chilean Tierra del Fuego was cited.

It had first been reported from Chile (‘Valdivia’) by

Bitter (1901), who stated that it had previously been

labeled as ‘Parmelia vittata’ or ‘P. lugubris’. It was

also reported from the very isolated cloud forest Fray

Jorge by Zahlbruckner (1926) and Follmann &

Redon (1972), and from Laguna San Rafael by

Quilhot et al. (2002). The species was also reported

from three provinces in Argentina by Calvelo &

Liberatore (2002), but no localities appear to have

been published from this country. Hypogymnia

subphysodes is common in southern and south-

eastern Australia and in New Zealand according to

Elix (1979). In the Northern Hemisphere it is known

from a single locality in Oregon, USA (McCune &

Rosentreter 1997).

The present study indicates that Hypogymnia

subphysodes is the most common sorediate species in

Chile. It is very common in humid Valdivian forests,

is also common in the Aysen Region, but has not

been seen in the Magellanic forests, although it may

be present as less common there. Hypogymnia

pulverata has the same distribution pattern, but

appears to be less frequent. Actually, it could not be

distinguished from H. subphysodes on Chilean

material by any other character than its more or less

solid and flattened lobes. Besides, this is not a

constant character, and a small sample such as Bjerke

980/01 could not be determined with full certainty.

As they shared the same two chemosyndromes in

Australasia, Elix (1979) suggested that they might not

represent taxa that are well separated at species level.

The need to study this hypothesis is supported here,

and a molecular approach could resolve this

question.

Selected specimens examined (of a total of 9).

CHILE: IX REGION DE LA ARAUCANıA, Provincia de Cautın,

N of Melipeuco, 5 km S of Parque Nacional

Conguillıo, 38u429S, 71u349W, 600 m, Tronstad 11

Nov. 1996 ; X REGION DE LOS LAGOS, Provincia de

Llanquihue, 1 km W of Lago Chapo, 41u259500S,

72u239W, 270 m, Bjerke 840/01 ; Provincia de Chiloe:

Cucao, Parque Nacional de Chiloe, near Guarderıa at

Tepual path, 42u379S, 74u069W, 30 m, Elvebakk

00:496 ; XI REGION AISEN DEL GENERAL CARLOS IBANEZ DEL

CAMPO, Parque Nacional Queulat, Rıo Ventisquero,

1 km E of Carretera Austral, 44u289S, 72u349W, 20 m,

Elvebakk 06:533.

DISCUSSION

Distribution of sorediate vs. primarily

fertile species. The three sorediate species present in

the study area all have amphi-tropical (lacking from

tropical areas, but occurring both to the north and

the south) distribution patterns. Their distributions

are clearly much more extensive in one of the

hemispheres, and much more limited in the other.

The latter can be interpreted as secondary

distribution areas, originating from probably rare

cases of long-distance distribution across the

equatorial area. It is noteworthy that the two

primarily Southern Hemisphere species and the

single Northern Hemisphere species all have parallel

patterns in this respect. Among the remaining

Northern Hemisphere species, sorediate ones have

the widest distribution areas. Only species with

vegetative dispersal propagules are present in Europe

and Africa. Thus, species such as Hypogymnia

austerodes (Nyl.) Rasanen, H. farinacea Zopf, H.

physodes (L.) Nyl., H. tubulosa (Schaer.) Hav., and H.

vittata (Ach.) Parrique are very widespread in the

Northern Hemisphere and extend marginally into

equatorial areas.

This pattern is in strong contrast to the

distributions of the primarily fertile species. Except

for the eastern North American H. krogiae Ohlsson,

they are restricted to four areas, East Asia, western

North America, Australia/New Zealand and southern

South America. Hypogymnia lugubris has the widest

distribution, but probably disperses by thallus

fragments. A few species cross the Bering Strait, and

H. enteromorphoides has a South American-

Australasian disjunction. The remaining primarily

fertile species all have limited distributions, even the

very common H. antarctica. The conclusion is that

spores, although smaller than soredia, are obviously

much less effective and viable long-distance diaspores

than soredia. The same has been shown, e.g. for

384 The Bryologist 114(2): 2011

related species of Menegazzia (Bjerke 2003; 2005) and

Pannaria (Elvebakk et al. 2007).

Chemistry. Four of the species studied here (H.

antarctica, enteromorphoides, pulveracea and

subphysodes) produce chemotypes defined by

compounds solely of the physodic acid complex or of

both the physodic and the physodalic acid

complexes. In addition, also single substances in the

physodic acid complex are shown to be

taxonomically important. All the 12 samples of

Hypogymnia antarctica, chemotype II, included 29-O-

methylphysodic acid, whereas the same substance

was lacking from the otherwise identical

chemosyndrome of H. physodes, tested on 17

reference samples (Norway, arctic Canada).

Hypogymnia bitteri is another example on

taxonomic importance of compounds in this

complex. When comparing this species with H.

austerodes, McCune (2002) found them to differ

chemically only in the frequency of 3-

hydroxyphysodic acid (in 25% of the samples in the

former, 75% in the latter). In the present study, this

substance was lacking from all 32 samples of the

former, and present in all 26 samples of the latter.

Vittatolic acid was named after H. vittata,

although not always cited for this species. It was

present in all 11 reference samples studied here of H.

vittata. However, in this species 3-hydroxyphysodic

acid was less diagnostic. It was equally often present,

lacking or replaced by a possible satellite, or even

artefact, named ‘cf. 3-hydroxyphysodic acid’ in

Table 1. Vittatolic acid has rarely been cited to occur

also in H. bitteri (Krog et al. 1994), but was present in

all 32 H. bitteri samples analyzed here. Conversely, it

was lacking from all H. austerodes samples. Thus,

these two species are chemically distinct, and the

chemosyndrome of Chilean material of H. antarctica

(chemosyndrome I), H. pulveracea and H.

subphysodes is shared by the reference samples of H.

austerodes and H. tubulosa (Fig. 2).

However, there might be geographical differences.

Five High Arctic samples of H. austerodes from

Svalbard were homogeneous, but six High Arctic ones

from Devon Island, Canada (not included in Fig. 2),

were heterogeneous. Øvstedal et al. (2009) recently

reduced H. subobscura to a synonym of H. austerodes,

indicating that they have identical chemistry. Three

samples of H. subobscura sampled for the present study

(one from Novaya Zemlya, two from Devon Island)

show, like in Elix (1979), that 3-hydroxyphysodic acid

was absent, which does not support lumping these two

species into one.

Rejected species from southern South America.

Three widely distributed Northern Hemisphere

sorediate Hypogymnia species have been reported

from southern South America: Hypogymnia farinacea

Zopf from Chile (Feuerer 2009; Pereira & San Martın

1998; Redon 1974, as H. bitteriana (Zahlbr.) Ras.), H.

physodes (L.) Nyl. from Argentina (Calvelo &

Liberatore, 2002; Grassi 1950) and H. vittata (Ach.)

Parr from Argentina (Calvelo & Liberatore 2002;

Cenga Sambio 1930). The latter is known otherwise

as far South as the very high mountains of Papua

New Guinea (Elix 1979) and H. physodes is reported

as far south as Kenya (Swinscow & Krog 1988,

Feuerer 2009). These species have not been reported

from any austral areas, and their presence in

Southern South America should be considered

doubtful, until vouchers can be located and critically

compared to other sorediate species.

Hypogymnia billardieri (Mont.) Filson, reported

by Rasanen (1932, as Parmelia conferta Tayl.), H.

turgidula (Bitter) Elix, reported by Rasanen (1932, as

P. turgidula Bitter), Galloway & Quilhot (1998) and

Feuerer (2009), and H. mundata (Nyl.) Oxner ex

Rassad., reported by Galloway & Quilhot (1998),

Galloway (2007) and Feuerer (2009) are not accepted

here, as they might refer to H. antarctica. The report

by Calvelo & Lorenzo (1989) of H. mundata from

Argentina cited by Galloway (2007) instead refers to

H. pulverata, which has also been considered a

variety of H. mundata. Hypogymnia deusta (Hook. F.

& Taylor) Dodge listed from the Cape Horn area

(Dodge 1965), has now been transferred to Nimisia

(Fryday 2005).

No phylogenetic studies have been published on

Hypogymnia yet. The affinity of the well-defined

‘Hypogymnioid group’ with three distinct Northern

Hemisphere genera (Crespo et al. 2007; 2010) could

indicate a Laurasia origin, although more studies will

be needed to confirm this. Menegazzia shares three of

the four centers of species diversity with Hypogymnia,

but with much higher species numbers in the two

centers in the Southern Hemisphere.

Elvebakk: Hypogymnia in Chile 385

As a conclusion, seven species of Hypogymnia

are now accepted for southern South America in the

present study, as a contrast to species included in

recent checklists (Table 1). Although the South

American Hypogymnia species, and primarily the

fertile ones, are still in need of a taxonomic revision,

the currently accepted species can be distinguished by

the following key.

Key to South American species of Hypogymnia.1 Soredia present, apothecia uncommon .............................. 2

1 Soredia absent, apothecia common .................................... 4

2 Soredia capitate and powdery, developing primarily at

lobe tips. Lobes appressed to the substrate ...... H. bitteri

2 Soredia coarser, laminal, gradually covering large parts

of lobes. Secondary ascending lobes common ........... 3

3 Lobes hollow and broadly elliptic to isodiametric in

circumscission ................................................. H. subphysodes

3 Lobes solid and flattened ................................ H. pulveracea

4 Thallus fruticose, mostly terricolous, upper side of lobes

with distinct black mottling ........................ H. lugubris

4 Thallus foliose, corticolous, upper side of lobes

sometimes with dark margins and black pycnidia, but

rarely with black mottling ............................................ 5

5 Lobes 3–6 mm broad, secondary extending lobes common,

perforations present on the lower side H. enteromorphoides

5 Lobes 1–2 mm broad, secondary lobes uncommon, perfo-

rations on the lower side absent ......................................... 6

6 Apothecia large and hollow, with dark brown discs,

upper surface rugose ................................. H. antarctica

6 Apothecia smaller and solid, with pale brown discs,

upper surface smooth ...................... H. solidepedicellata

ACKNOWLEDGMENTS

The author acknowledges permission to collect in Chile by

Corporacion Nacional Forestal de Chile (CONAF), the

company in the field by Dr. J. W. Bjerke and E. H. Robertsen,

University of Tromsø, and to comments to the manuscript by

the former and by B. McCune, Oregon State University.

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