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The lichen genus Heterodermia (Physciaceae) in South America – a contribution including five new species

March 2011
Nordic Journal of Botany 29(2):129 - 147

March 2011
29(2):129 - 147

DOI:10.1111/j.1756-1051.2009.00519.x

Authors:

Uppsala University

Thirty-three species of the lichen genus Heterodermia in South America, mainly from Ecuador and Peru, are defined. Morphology, anatomy, chemistry, habitat, distribution and interrelation between the species are discussed. A key to the treated species is presented. Five species are described as new; Heterodermia andina, H. arvidssonii, H. badia, H. fertilis and H. parva. One new combination is proposed; H. spinigera. Two species are reported as new to South America, H. spathulifera and H subcitrina, and H. palpebrata is reported as new to USA.

Heterodermia allardii. Peru, 1981, Santesson, Tehler and Thor P100:17 (in UPS). Scale bar=1 mm.

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Heterodermia andina. Ecuador, 1983, L. and A. Arvidsson 3137a (in GB). Scale bar=2 mm.

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Heterodermia arvidssonii. Equador, 1972, Arvidsson and Nilsson 1200 (in GB). Scale bar=2 mm.

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Heterodermia badia. Peru, 1981, Santesson, Tehler and Thor P90:30 (in UPS). Scale bar=1 mm.

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Heterodermia fertilis. Peru, 1981, Santesson and Moberg P43:8 (in UPS). Scale bar=2 mm.

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The lichen genus

Heterodermia

(Physciaceae) in

South America



a contribution including ﬁve new species

Roland Moberg

R. Moberg (roland.moberg@evolmuseum.uu.se), Museum of Evolution, Uppsala Univ., Norbyva

¨gen 16, SE752 36 Uppsala, Sweden.

Thirty-three species of the lichen genus Heterodermia in South America, mainly from Ecuador and Peru, are defined.

Morphology, anatomy, chemistry, habitat, distribution and interrelation between the species are discussed. A key to the

treated species is presented. Five species are described as new; Heterodermia andina,H. arvidssonii,H. badia,H. fertilis

and H. parva. One new combination is proposed; H. spinigera. Two species are reported as new to South America,

H. spathulifera and H subcitrina, and H. palpebrata is reported as new to USA.

Among the foliose lichens, the genus Heterodermia (Phys-

ciaceae) is one of the most common in tropical and

subtropical regions, with a few species reaching temperate

regions, e.g. H.speciosa. The genus was included in the

genus Anaptychia by Kurokawa (1962) in his monograph,

but Poelt (1965) separated those species having thick-

walled spores and containing the substance atranorin

(Heterodermia) from those with thin-walled spores and

lacking atranorin (Anaptychia). Later studies, mainly on a

regional basis, which include parts of North America

(Culberson 1966, Moberg and Nash 1999), India (Awasthi

1973), east Africa (Swinscow and Krog 1976, 1978), the

Azores (Moberg and Purvis 1997) South Africa (Moberg

2004a), Europe (Moberg 2004b) and Costa Rica (Lu

¨cking

et al. 2008), have accepted this division with only a few

exceptions (Kurokawa 1973).

The genus Heterodermia is recognized by its greyish upper

surface reacting Kyellow (atranorin), a prosoplectenchy-

matous upper cortex (hyphae parallel to the surface), and

thick-walled, 2-celled ascospores, often having sporoblasti-

dia. Pycnidia are usually present when apothecia are present

and often absent in other cases. They are immersed in the

thallus with black weakly protruding tips and pycnoconidia

are bacilliform 461mm. The subdivision of the genus

suggested by Kurokawa (1962) is not used.

Recent DNA studies of the ‘Heterodermia obscurata

group’ (Lu

¨cking et al. 2008) have revealed very interesting

results concerning the species pair concept. Although their

results are very convincing, the present contribution is not

completely adjusted to their results since their study is

limited to Costa Rica and I feel that it is necessary to study

the material used.

This contribution includes thirty-three species, mainly

from Peru and Ecuador, but also some material from

other parts of South America. The aim is to present the

results from the Regnellian expeditions to Peru and

Ecuador collected in the 1980s. In addition, herbarium

material from Argentina, Brazil, Colombia, Uruguay and

Venezuela present at herbarium UPS has been included

in the current study. As this material is limited, the study

of South American Heterodermia is not complete, but all

known species accepted by me are discussed and treated in

alphabetic order. A few recently described species (Sipman

1995, p. 329, Marcelli et al. 2007, p. 221) are only briefly

mentioned as I have seen too few specimens (or none) to

judge their status.

The studied material includes more than 500 specimens

and ‘Specimens examined’are listed below only for the new

and less known species. However, a complete list of

examined specimens is preserved at UPS. In spite of the

number of studied specimens, the distribution given have to

be regarded as incomplete, as mainly material from the

herbaria GB, UPS and S has been studied extensively.

Important characters for separating the different

species are:

1) thallus and lobe shape, 2) presence of cilia, 3) presence

and shape of soralia and isidia/squamules, 4) presence or

absence of a lower cortex, 5) ascospore size and structure,

6) chemistry.

Thallus and lobe shape

Most species can be characterized as rosette-shaped, which

means that the lobes are 9radiating. These include H.

albicans,H. andina,H. badia,H. chilensis,H. corallophora,

H. diademata,H. flabellata,H. granulifera,H. hypoleuca,H.

isidiophora,H. japonica,H. magellanica,H. obscurata,H.

pseudospeciosa,H. spathulifera,H. speciosa,H. squamulosa,

Nordic Journal of Botany 29: 129147, 2011

doi: 10.1111/j.1756-1051.2009.00519.x,

#2011 The Author. Nordic Journal of Botany #2011 Nordic Society Oikos

Subject Editor: Torbjo

¨rn Tyler. Accepted 26 August 2009

129

and H. subcitrina. In some species the lobes are ribbon-like

as in H. allardii,H. arvidssonii,H. fertilis,H. galactophylla,

H. kurokawae,H. leucomela,H. lutescens,H. multiciliata

and H. vulgaris. In a few species the thallus is 9fruticose

and usually short-lobed, as in H. comosa,H. palpebrata,

H. podocarpa,H. spinigera and H. trichophora.

Cilia

Cilia are outgrowths from the upper cortex, in contrast

to rhizines which originate from the lower cortex. If the

lower cortex is lacking, any outgrowths are thus regarded

as cilia. Only H. comosa,H. multiciliata,H. spinigera and

H. trichophora have cilia on the upper surface; H. spinigera

and H. trichophora mainly on the underside of the

apothecia.

Of the species with ribbon-like lobes the following

have dark cilia: H. fertilis,H. leucomela,H. lutescens,H.

multiciliata and H. vulgaris. Pale cilia occur in H. allardii,

H. arvidssonii,H. comosa,H. galactophylla,H. palpebrata,

H. podocarpa and H. spinigera (darkening).

Isidia/squamules

It could sometimes be difficult to separate isidia and

squamules. In this paper, isidia are defined as having cortex

on all sides and squamules as having cortex only on the

upper side. Isidia are present in H. corallophora,H.

granulifera and H. isidiophora. Squamules are regularly

present in H. badia and H. squamulosa.

Lower cortex

A distinct lower cortex occur in H. albicans,H. andina,H.

diademata,H. granulifera,H. isidiophora,H. pseudospeciosa,

H. spathulifera and H. speciosa. All other South American

species lack a lower cortex.

Ascospores

Most ascospores have sporoblastidia, i.e. vacuoles in the

ascospore wall. In general, the sporoblastidia are more

obvious in larger ascospores, and only in a few species

they are lacking. Heterodermia fertilis and H. leucomela

have abundant small sporoblastidia scattered in the ascos-

pore wall.

Chemistry including pigments

Secondary chemistry provides important characters in many

species, but in some the variation is considerable and hardly

correlated with the morphology. In my opinion, the

importance of chemical characters is often overestimated,

an opinion shared by Swinscow and Krog (1976). It seems,

however, that absence or presence of salazinic acid is a

reliable character, while presence or absence of norstictic

acid varies e.g. in H. japonica. This is contradicted by

¨cking et al. (2008) and should be examined further.

In a study of H. japonica from east Africa (unpubl.),

I separated those species with norstictic acid from those

without, but was not successful in separating them

morphologically. Salazinic and norstictic acid does not

occur together in the examined material, and as the K test

have a similar yellowred reaction it seems necessary to

identify them by TLC. Sometimes the terpenoids form

a pattern useful for identification e.g. in H. spathulifera.

Several species have a pigmented lower surface, and

examination by TLC reveals many different pigments.

Kurokawa (1973, p. 575) has presented a table of such

pigments. I will not give a similar table here, but only

a summary of species with pigments. For further informa-

tion, see the appropriate species description. Defined

pigments are present in H. comosa,H. flabellata,H.

lutescens,H. obscurata,H. subcitrina and H. vulgaris.

Many species, such as H. badia,H. chilensis,H. corallophora,

H. fertilis,H. hypoleuca,H. japonica,H. kurokawae,H.

leucomela,H. magellanica and H. squamulosa have a more or

less coloured underside. A detailed study of pigments is in

preparation.

Material and methods

The present study is based on field studies in Peru (1981)

and Chile (1995). Collected material from Peru by myself,

R. Santesson, A. Tehler and G. Thor and from Chile

by myself and A. Tehler was examined together with

herbarium material from FH, G, GB, H, LIS, S, TROM,

TUR UPS, WU. Some 500 specimens were examined

and a complete list of specimens is held at UPS. Chemical

analyses were performed using standardized thin layer

chromatography (TLC) (White and James 1985, Arup

et al. 1993) using solvent system C. Ascospore measure-

ments were made in water mounts and have mostly been

statistically calculated except when limited material was

available. Lists of examined specimens are given only for

new and rare species which hopefully will be more collected.

If no further data is given, the material is preserved at UPS.

Duplicates of some specimens are deposited in CTES and F.

Key to species

1. Thallus mainly consisting of small stipitate apothecia,

sometimes surrounded by a very small lobate thallus ...

.............................................H. parva

Normal thallus developed ..........................2

2. Lobes dichotomously branched, distinctly elongate with

prominent marginal cilia (‘leucomela-group’)..........3

Lobes irregularly branched, not distinctly elongate,

with or without marginal cilia...................... .10

3. Lower side with pigment .............................4

Lower side without pigment ........................8

4. Pigment yellowish (if pinkish K).....................5

Pigment reddish or brownish .......................6

5. Marginal cilia pale, 9abundant; pigment inspersed in

the medulla.............................H. kurokawae

Marginal cilia black, not very abundant; pigment

superficial.................................H. lutescens

6. Pigment reddish to purplish ............... .H. vulgaris

Pigment brownish or pinkish .......................7

7. Pigment Kpurple......................H. arvidssonii

Pigment K...............................H. fertilis

8. With prominent ciliate/lobate apothecia ...............9

Without or rarely with apothecia (cf. also H.

multiciliata)..............................H. leucomela

130

9. Lobe length more than 10 times lobe width; apothecia

lobate......................................H.fertilis

Lobe length less than 10 times lobe width; apothecia

not or sparsely lobate..................H. multiciliata

10. With prominent marginal cilia .....................11

Without marginal cilia, but rhizines may project from

underside of lobes.................................19

11. Laminal cilia 9abundant (few in H. spinigera)......12

Laminal cilia absent (may appear on underside of

apothecia)........................................15

12. Underside with yellow pigment, occasionally with

cyanobacteria..............................H. comosa

Underside without pigment......................13

13. Lobes 9plane, linear; apothecia laminal ............14

Lobes convex, paddle-shaped when young; apothecia

terminal...................................H. comosa

14. Cilia pale to darkening ....................H. spinigera

Cilia black...........................H. multiciliata

15. With terminal, labriform soralia; without apothecia . .16

Without soralia; apothecia 9abundant .......... .17

16. With norstictic acid ........................H. allardii

Without norstictic acid..............H. galactophylla

17. Apothecia terminal, or distinctly stipitate ............18

Apothecia laminal, shortly stipitate .....H. palpebrata

18. Underside of apothecia with short cilia ...H. trichophora

Underside of apothecia without cilia ...H. podocarpa

19. With lower cortex .................................20

Without lower cortex............................27

20. With isidia........................................21

Without isidia.................................. .22

21. Isidia granular; upper surface 9dull; Kyellow 0red,

salazinic acid...........................H. granulifera

Isidia cylindrical; upper surface 9shiny; Kyellow

.......................................H. isidiophora

22. Without soralia.........................H. diademata

With soralia.....................................23

23. Soralia mainly marginal, not labriform; with norstictic

or salazinic acids..................................24

Soralia mainly apical, labriform; without norstictic

or salazinic acids..................................25

24. With salazinic acid, Kyellow 0red ......H. albicans

With norstictic acid, Kyellow....H. pseudospeciosa

25. Lobes flat to concave, rounded and distinctly widening

at tips; underside blackish; rhizines often ‘squarrose’...

...........................................H. andina

Lobes flat to convex, not distinctly widening at tips;

underside not blackish; rhizines mainly simple ......26

26. Soralia often few and large, 9firmly attached, contain-

ing spathulin..........................H. spathulifera

Soralia abundant, 9loosely attached, without spathu-

lin ........................................H. speciosa

27. Lower side with yellow or rusty pigments in the

medulla, not violet in central part ..................28

Lower side not pigmented, but mostly turning violet

or blackish in central parts .........................30

28. Without soralia..........................H. flabellata

With soralia.....................................29

29. With distinct terminal soralia; pigment mostly rusty ...

.........................................H. obscurata

With mainly marginal soralia; pigment mostly yellow

........................................H. subcitrina

30. Without vegetative diaspores, mostly with apothecia. .31

With isidia, squamules or soralia ..................32

31. Ascospores 3746 mm long....H. magellanica-complex

Ascospores 2634 mm..................H. hypoleuca

32. With isidia or squamules ...........................33

With soralia.....................................35

33. With isidia............................H. corallophora

With squamules............................... . .34

34. With norstictic acid; lobes less than 1 mm broad;

marginal squamules rounded................H. badia

Without norstictic acid; lobes more than 1 mm broad;

marginal squamules 9elongate ..........H. squamulosa

35. Lobes narrow, convex......................H. chilensis

Lobes flat to concave, very variable ......H. japonica

Heterodermia albicans

(Pers.) Swinsc. & Krog

(1976, p. 113)

(non Anaptychia albicans Kurok. 1962, p. 80). Basionym:

Parmelia albicans Pers. (1810, p. 17). Holotype: St Domingo

(in L, not seen). Taxonomic synonym: Anaptychia domingensis

(Ach.) Massal. (1853, p. 39).

Literature: Swinscow and Krog (1976, p. 114, 1988,

p. 89), Moberg and Purvis (1997, p. 192), Moberg and

Nash (1999, p. 2).

Thallus orbicular to irregular, small, lessthan 3 cm in diameter,

firmly adnate; upper surface grey to brownish grey, darker at

lobe tips, sometimes weakly pruinose. Lobes narrow, 0.5(1.0)

mm, short, to 3mm, usually richly branched, weakly

convex, 9widening, tips convex, without soralia, not ascend-

ing. Soralia white to bluish grey, arising from small lateral

knob-like structures forming small, 9continuous marginal

soralia towards the thallus centre, sometimes becoming

granular. Lower surface corticate, whitish to pale brownish,

rarely dark grey, weakly rhizinate with usually short (ca 1 mm),

mostly dark brown or black rhizines. Upper and lower cortex

prosoplectenchymatous. Apothecia very rare, to 3 mm in

diameter, ascospores Pachysporaria-type, without sporoblasti-

dia, (18.5)19.528.0(32.5) (8.5)10.513.5(15.0) mm.

M23.8;11.9 mm. SD4.38;1.60 mm. n16. Pycnidia

rare.

Chemistry:Kyellow0red. Atranorin, zeorin, salazinic

acid, unidentified triterpenoids (9).

Habitat and distribution: Mainly on rocks or over mosses

on rocks in open situations, rarely on tree trunks in tropical

to warm temperate areas. I have examined material from

Brazil, Ecuador, Peru and Venezuela. Also known from

east Africa, South Africa, the Azores, Canary Islands,

Madeira and North America (Mexico, Arizona).

Similar species

Heterodermia albicans is characterized by the narrow,

convex, short lobes and the corticate lower surface. The

soralia arise from marginal knob-like structures forming

small, 9continuous marginal soralia towards the thallus

centre. Heterodermia speciosa also has a lower cortex but has

larger, 9flat lobes and lacks salazininc acid (Kyellow

only). Heterodermia albicans differs from H. pseudospeciosa

in the more adpressed thallus, less delimited soralia and

131

the absence of norstictic acid. TLC is recommended to

separate the two species. In contrast to H. speciosa, the

soralia are usually not terminal labriform.

Heterodermia allardii

(Kurok.) Trass (1992, p. 6)

(Fig. 1)

Basionym: Anaptychia allardii Kurok. (1962, p. 98).

Holotype: Peru, Hua

´nuco, San Martin, east of Tingo

Maria, in jungle on ridge, 2500 ft, 19491950 Allard

21408c (in US, not seen).

Thallus irregular, up to 3 cm in diameter, 9loosely attached

to substratum with elongate lobes having prominent pale,

marginal cilia; upper surface pale grey without pruina. Lobes

to 1 mm broad, but widening towards apices and sometimes

reaching 4mm, particularly when soralia appear. Soralia

terminal, labriform on the underside of lobes. Without lower

cortex. Upper cortex prosoplectenchymatous. Apothecia and

pycnidia not seen.

Chemistry:Kyellow. Atranorin, zeorin and norstictic acid.

Habitat and distribution: On branches in fairly open

situations at altitudes between 750 and 1750 m a.s.l.

Known from Peru, Brazil and Bolivia. Also known from

Central America and West Indies.

Similar species

Heterodermia allardii is similar to H. galactophylla and

without a chemical test (TLC), the two species could hardly

be separated; H. allardii contains norstictic acid. Kurokawa

(1962) reports salazinic acid which has not been found by

TLC. The amount of norstictic acid varies and the K

reaction (red) is not easily seen. Regarding the chemical

variation in other species, this species may well be included

in H. galactophylla, but I have seen too little material to

formalize such a treatment. Already in his description,

Kurokawa states that they are hardly separable without

chemical analysis.

Specimens examined (all in UPS): Bolivia. 1934 A. M. Bang.

Brazil.Sa

˜o Paulo, Serra da Mantiqueira. Campos do

Jordao, 1700 m a.s.l. 1975, Schindler 5501; 1750 m

a.s.l. 1977, Schindler 7167. Peru. Cusco, Paucartambo,

road Paucartambo-Pillcopata, ca 34 km (road distance)

from Pillcopata, alt 1500 m a.s.l. 13800?S, 71815?W.

On branches. 1981, Santesson, Tehler and Thor P100: 17.

Heterodermia andina

Moberg sp. nov. (Fig. 2)

Thallus foliaceus,diametro usque ad 4 cm,supra griseus,

pruinosus,subtus corticatus,rhizinis squarrosis.Lobi radiantes,

usque ad 2 mm lati.Soralia marginalia.

Holotype: Ecuador, Imbabura, Quichinche, 8 km west of

Otavalo, 0813?S, 78820?W, 2500 m a.s.l. Agriculture area.

Epiphyte. 17 Jun 1983, L. and A. Arvidsson 3137a (in GB).

Thallus orbicular to irregular, to 4 cm in diameter, with

radiating lobes; upper surface pale grey, dull pruinose. Lobes

to 2 mm broad, rounded and widening at tips, often distinctly

concave. Soralia lip-shaped on lateral lobes, rarely present

on terminal lobe tips. Lower surface corticate, pale at the

very tips, centrally dark brown to black with dark brown

to black, often squarrose, 9abundant rhizines. Upper and

lower cortex prosoplectenchymatous. Only one apothecium

seen, 2 mm in diameter; ascospores of Pachysporaria-type,

without sporoblastidia, 2225812 mm. Pycnidia not seen.

Chemistry:Kyellow. Atranorin, 9zeorin, 9leucotylin,

and a distinct triterpene below atranorin (TLC solvent C).

Habitat and distribution: On trees, rarely on rocks in fairly

open areas as on wayside trees but also in more moist

situations such as along streams. So far only found in

Venezuela and Ecuador in the Andes.

Notes:Heterodermina andina is easily recognized by the

pruinose upper surface, the rounded lobes and the lateral,

labriform soralia.

Figure 1. Heterodermia allardii. Peru, 1981, Santesson, Tehler

and Thor P100:17 (in UPS). Scale bar 1 mm.

Figure 2. Heterodermia andina. Ecuador, 1983, L. and A.

Arvidsson 3137a (in GB). Scale bar2 mm.

132

Specimens examined (paratypes): Ecuador. Azuay, road

Cuenca-Loja, 3 km north of Cumbe, 2600 m a.s.l. 1983,

L. and A. Arvidsson 4403 (in GB); ca 10 km south of Paute

and 6 km north of Gualace

´o, 2300 m a.s.l., 1987, K. and A.

Kalb 19531 (in hb Kalb); Bolivar, 15 km east of Guaranda

on road to Riobamba 1835?S, 78855?W, 3200 m a.s.l.

1985, Arvidsson, Lindqvist and Lindstro

¨m 6319 (in GB);

Imbabura, road Otavalo San Jose

´de Minas, 20 km west

of Otavalo, 3000 m a.s.l. 1983, L. and A. Arvidsson 3197

(in GB); Pichincha, Quito, Univ. de Catolica, 0813?S,

78830?W, 2800 m a.s.l. 1985, Arvidsson, Lindqvist and

Lindstro

¨m 5696a (in GB); the Guayllabamba Valley,

4.5 km north of the village Guayllabamba on the road

Cayambe-Quito, 0804?S, 78820?W, 2300 m a.s.l. 1985,

Lindstro

¨m and Lindqvist 2453 (in GB); Tungurahua

Pastaza Valley, between Banos and Rio Verde, 1500 m

a.s.l. 1983, L. and A. Arvidsson 4088 (in GB). Venezuela.

Lara, Sierra de Barbacoas, entre la quebrada de El Vino y

Barbacoas, 1700 m a.s.l. 1978, Lopez-Figueiras and Smith

16533 (in UPS).

Heterodermia arvidssonii

Moberg sp. nov. (Fig. 3)

Thallus foliaceus,irregularis,dichotome divisus,supra griseus,

subtus decorticatus,granulatussorediatus,roseus pigmentifer,

Kruber;laciniae lineares,margine ciliatae;cilia basi

pallida apice nigrescentia.Apothecia non visa.

Holotype: Ecuador. Napo, ca 5 km west of Pifo, 2700 m

a.s.l. Epiphyte. 12 Mar 1972, L. Arvidsson and D. Nilsson

996 (in GB).

Thallus irregular, to 3 cm, loosely attached to the substrate

or rarely ascending, dichotomously branched; upper surface

pale grey, with pale marginal cilia. Lobes linear, elongate,

12 mm broad, rarely widening at tips, cilia pale at base,

occasionally becoming darker towards tips. Lower surface

ecorticate partly dissolved into granular soredia. Medulla

overlaid with a rose pigment, Kdark red. Apothecia and

pycnidia not seen.

Chemistry: atranorin, zeorin and a rose pigment, Kred.

Habitat and distribution: Epiphytic on twigs in fairly open

situations as roadsides but also in cloud forest. Known only

from a few localities in Ecuador.

Notes:Heterodermia arvidssonii is easily recognized by the

pale marginal cilia and the rose, Kdark red pigment on

the underside.

Specimen examined (paratype): Ecuador. Loja, Malacatos,

1600 m a.s.l. Roadside, epiphyte, 1972, L. Arvidsson and

D. Nilsson 1200 (in GB).

Heterodermia badia

Moberg sp. nov. (Fig. 4)

Thallus foliaceus,irregularis,supra brunneus vel brunneogri-

seus,apicibus loborum pruinosis,subtus decorticatus,rhizinis

sparsis.Lobi 9ascendentes,usque ad 1 mm lati,margine 9

lobulati. Soralia et isidia non visa.

Holotype: Peru. Cusco, Urubamba, Valley of Rio Piri,

1617 km road distance, from (northwest of) Ollantay-

tambo, 13807?S, 72822?W, 3200 m a.s.l. On soil on

rocks. 23 Mar 1981, R. Santesson P85:4 (in UPS, isotypes:

CTES, F).

Thallus irregular, rarely orbicular, centrally sometimes

squamulose; upper surface brownish to brownish grey,

pruinose at lobe tips. At higher altitudes the lobes become

shorter, narrower and darker in colour. Lobes linear, 9

ascending, often disjunct, plane to weakly convex, with

9abundant adventitious marginal lobules, rarely also on

the surface, occasionally central parts covered with lobules;

lower surface white, to dark blue centrally. Soralia and isidia

absent. Upper cortex prosoplectenchymatous, lower cortex

absent; medulla white, Apothecia rare, to 3 mm in diameter

with abundant squamules; ascospores of Pachysporaria-type,

with sporoblastidia, 44541821 mm. Pycnidia not seen.

Chemistry:Kyellow. Atranorin, zeorin, norstictic acid and

leucotylin.

Figure 3. Heterodermia arvidssonii. Equador, 1972, Arvidsson and

Nilsson 1200 (in GB). Scale bar2 mm.

Figure 4. Heterodermia badia. Peru, 1981, Santesson, Tehler and

Thor P90:30 (in UPS). Scale bar1 mm.

133

Habitat and distribution: On rocks or among mosses on

rocks in open situations at high altitude. So far known

only from Peru and Bolivia at altitudes between 2500 and

4100 m a.s.l.

Similar species

Heterodermia badia is recognized by the narrow, 9

ascending lobes with pruinose tips. The marginal lobules

on not ascending lobes are also an important character.

It might be confused with H. corallophora which, however,

has true isidia and lacks norstictic acid. Heterodermia

squamosa has broader, concave lobes and elongate marginal

squamules and lacks norstictic acid.

Specimens examined (paratypes):Bolivia. La Paz, Nonuo, ca

25 km east of La Cumbre, 16819?S, 67858?W, 4100 m a.s.l.

2001, Bjerke 073/01 (TROM). Peru. Cusco, Urubamba,

valley of Rio Piri, ca 30 km road distance from (ca northwest

of) Ollantaytambo, 13805?S, 72822?W, 3700 m a.s.l. 1981,

Santesson, Tehler and Thor P89:33, P89:35 (in UPS);

ca 28 km, road distance, from (northwest of) Ollantay-

tambo, 13806?S, 72822?W, 3600 m a.s.l. 1981, Santesson,

Tehler and Thor P90:5, P90:30 (in UPS); Paucartambo,

road Paucartambo-Pillcopata, 1 km northeast of the turn

to Tres Cruces, 13807?S, 71837?W, 3500 m a.s.l. 1981,

Santesson, Tehler and Thor P106:14 (in UPS); Machu

Piccu, between uppermost ruins and inka bridge, 13809?S,

72831?W, 2900 m a.s.l. 2001, Bjerke 140/01 (TROM).

Hua

´nuco, Dos de Mayo, valley de Rio Maranon, ca

58 km (road distance) westnorthwest of Huanuco, 9848?S,

76840?W, 3900 m a.s.l. 1981, Santesson and Moberg

P50:10 (in UPS); valley of Rio Hirueras, 23 km (road

distance) west of Huanuco, 9855?S, 76824?W, 2500 m a.s.l.

1981, Santesson and Moberg P47:28 (in UPS); 27 km (road

distance) west of Huanuco, 9855?S, 76826?W, 2700 m a.s.l.

1981, Santesson and Moberg P46:7 (in UPS).

Heterodermia chilensis

(Kurok.) Swinsc. & Krog

(1976, p. 115)

Basionym: Anaptychia chilensis Kurok. (1962, p. 65).

Holotype: Chile, Valparaiso, Alto del Puerto, Aug 1940,

Santesson 2919 (in S, isotype: UPS).

Literature: Swinscow and Krog (1976, p. 115, 1988, p. 90),

Moberg (2004a, p. 259).

Thallus irregular to orbicular, up to 5 cm in diameter, 9

loosely adnate, lobe tips not ascending, only weakly

widening towards apices; upper surface pale grey to dark

grey, matt. Lobes radiating, to 2 mm, usually discrete.

Soralia labiate, terminal; soredia 9granular. Lower surface

without cortex, but with thick corticate margins, white

at apices, rarely brownish or bluish black centrally; rhizines

sparse, marginal. Apothecia (in isotype) to 4 mm in

diameter, subsessile with crenulate margins; ascospores of

Pachysporaria-type, remarkably wide and not thickened

at apices, with sporoblastidia, 27401728 mm.

Chemistry:Kyellow. Atranorin and zeorin.

Habitat and distribution: Mainly on rocks in fairly open

situations in subtropical regions. Seen from Chile and Peru

in South America. Also known from east Africa.

Notes:Heterodermia chiliensis is characterised by the narrow,

convex lobes, the white lower side with thick, corticate

margin and terminal, robust soralia. May sometimes be

difficult to separate from the related H. japonica.

Heterodermia comosa

(Eschw.) Follmann & Redon

(1972, p. 446)

Basionym: Parmelia comosa Eschw., in Martius (1828,

p. 26). Type: Crescit ad ramulos arborum in Provincia

Parae

¨nsi, e.g. prope Villam da Porta Mo

ˆz. Icon. Table XII,

Fig. 1.

Epitype: Brazil, Minas Gerais, Caldas, ad arbores, Aug

1873, H. Mose

´n (in UPS, fide Moberg and Nash 1999,

p. 4).

Literature: Kurokawa (1962, p. 103), Swinscow and Krog

(1976, p. 116), Moberg and Nash (1999, p. 4), Moberg

(2004a, p. 259).

Thallus irregular, fruticose, usually forming small tufts of

ascending lobes, to 6 cm wide; upper surface pale grey, mostly

provided with 9abundant cilia. Lobes sometimes short and

erect, when well-developed elongate, to 5 mm wide, convex,

with prominent marginal and laminal, pale cilia. Soralia

and isidia absent. Lower surface without cortex; medulla

lax, when well-developed usually inspersed with a yellow to

rusty pigment. Apothecia common, to 10 mm in diameter,

lobulate, ciliate as in thallus, situated at the end of the

ascending lobes; ascospores Pachysporaria-type, with or with-

out sporoblastidia, (28.0)32.543.5(49.5) (13.0)15.5

22.5(26.0) mm, M38.0; 19.0 mm, SD5.33; 3.32 mm,

n56. Pycnidia 9common.

Chemistry:Kyellow. Atranorin, zeorin and a Kpigment.

Habitat and distribution: On tree trunks and branches

in fairly open situations in tropical and subtropical areas.

Seen from Argentina, Bolivia, Brazil, Colombia, Ecuador,

Paraguay, Peru and Venezuela in South America. Also

known from Africa, Asia and North America.

Similar species

Heterodermia comosa is recognized by its convex ciliate lobes

with apical, ciliate apothecia and the lax medulla inspersed

with a yellow pigment. In young thalli this pigment is

often absent. In some material the pigment is replaced

by cyanobacteria as has been observed in other species

with the same pigment.

Heterodermia corallophora

(Tayl.) Skorepa

(1972, p. 490)

Basionym: Parmelia corallophora (‘coralliphora’) Tayl.

(1847, p. 164). Based on the same type: Anaptychia

hypoleuca ssp. corallophora (Tayl.) Vain. (1890, p. 135).

Holotype: Peru, Casapi, Mathews (in FH, isotype: K).

134

Literature: Kurokawa (1962, p. 62).

Thallus orbicular to irregular, up to 5 cm in diameter with

radiating, isidiate lobes; upper surface grey, sometimes

brownish centrally. Lobes plane to convex, to 2 mm wide

but usually narrower, tips often irregularly crenate. Isidia

cylindrical, often abundant and sometimes covering central

parts, sometimes in tufts, marginal to laminal. Lower

surface without cortex, white at tips, yellowish or purplish

in inner parts. Apothecia uncommon, sessile, with margin

dissolving into isidia, to 5 mm in diameter, ascospores

Pachysporaria-type, with sporoblastidia, (35.5)37.544.5

(46.5)(15.0)17.019.5(20.5) mm, M 40.9; 18.3 mm,

SD3.58; 1.42 mm, n 16. Pycnidia not seen.

Chemistry:Kyellow. Atranorin, zeorin and leucotylin.

Habitat and distribution: Mainly on trees and shrubs in the

tropical and subtropical zones in Central and South

America. Seen from Ecuador, Peru and Venezuela.

Similar species

The abundant cylindrical isidia, and the decorticate lower

side with purplish inner parts makes this species easily

recognized. A similar species is H. isidiophora which differs

in having a lower cortex and often irregularly crenate lobe

tips. H. badia differs in having lobules along the margin

and containing norstictic acid. Taylor (1847) published the

name as Parmelia coralliphora, an epithet which has been

corrected to corallophora. Kurokawa regarded the species as

combined at species level by (Vainio (1890), p. 135), but

in this publication the taxon is treated at subspecific level.

All subspecies are marked with an asterisk in that

publication as was common at that time (see for instance

p. 87, 93).

Heterodermia diademata

(Tayl.) D. D. Awasthi

(1973, p. 113)

Basionym: Parmelia diademata Tayl. (1847, p. 165).

Holotype: Nepal, Wallich (in FH-Taylor, not seen).

Literature: Swinscow and Krog (1976, p. 117, 1988, p. 92),

Kurokawa (1962, p. 28), Moberg (2004a, p. 260).

Thallus orbicular, to 5 cm in diameter or irregular and

confluent with other thalli, 9loosely adnate; upper surface

grey to dark grey, rarely brownish, margins darker. Lobes

to 2 mm wide, flat to concave, often truncate, with marginal,

small, budlike lobules arising from delimited pseudocyphel-

lae. Soralia and isidia absent. Lower surface with cortex,

prosoplectenchymatous; white to pale-grey or brownish

with scattered, pale to almost black rhizines. Apothecia

common, to 5mm in diameter, sessile, margins crenulate to

lobulate; ascospores Pachysporaria-type, without sporoblasti-

dia, (25.0)26.531.5(37.5) (9.5)12.515.0(16.0) mm,

M28.9; 13.8 mm, SD 2.48; 1.15 mm, n48. Pycnidia

common.

Chemistry:Kyellow. Atranorin, zeorin and leucotylin.

Habitat and distribution: Mainly on tree trunks and

branches, rarely on rocks in tropical and subtropical areas.

Seen from Argentina, Brazil, Ecuador, Paraguay, Peru and

Uruguay in South America. Also known from Africa, Asia

and Central America.

Similar species

Heterodermia diademata is characterized by the presence

of apothecia, and by the budlike lobules along the lobe

margins. The pseudocyphellae from which the almost

circular lobules develop is a very distinct feature of this

species. Evidently it is the fertile counterpart of H. speciosa.

Figure 5. Heterodermia fertilis. Peru, 1981, Santesson and Moberg P43:8 (in UPS). Scale bar 2 mm.

135

Heterodermia fertilis

Moberg sp. nov. (Fig. 5)

Heterodermiae leucomelae similis sed apotheciis abundantibus

lobatis,ciliis abundantibus squarrosis differt.

Holotype: Ecuador, Azuay, road Cuenca-Molleturo, 25 km

west of Cuenca. 2846?S, 79812?W, 3600 m a.s.l. Grass

paramo and shrub vegetation, epiphyte. 8 Jul 1983, L. and

A. Arvidsson 4158 (in GB, isotype: UPS).

Thallus irregular with narrow, interwoven lobes, sometimes

several cm long, with long marginal cilia, often richly

branched which gives the thallus a shaggy appearance; upper

surface whitish to cream-coloured, without pruina. Lobes

1.0(1.5) mm wide, parallel-sided, not widening towards

apices, dichotomously branched with long, squarrose

rhizines, black except for the pale base. Soralia and isidia

absent. Lower surface without cortex, white or rarely pinkish.

Apothecia abundant, laminal to apical, to 3 mm in diameter,

stipitate, disc black, 9pruinose, margin with mostly promi-

nent lobules and cilia; ascospores Pachysporaria-type, often

with numerous, spread sporoblastidia, (36.5)45.063.0(

75.5)(20.5)22.028.0(31.0) mm, M53.9; 25.0 mm,

SD9.10; 3.00 mm, n48. Pycnidia rare.

Chemistry:Kyellow. Atranorin, zeorin and sometimes

with a K‘pigment’.

Habitat and distribution: Epiphytic over mosses in both

moist and fairly dry situations in tropical and subtropical

areas. So far known from Bolivia, Brazil, Chile, Ecuador

and Peru.

Similar species

Heterodermia fertilis is characterised by the abundant lobate

and ciliate apothecia and absence of soralia, which separates

it from H. leucomela. Apices are not spirally recurved as in

H. leucomela ssp. boryi.

Specimens examined (paratypes).Bolivia. La Paz, Mapiri.

1893, Bang 1751 (in UPS). Brazil. Minas Gerais, Caldas,

21856?S, 46822?W. 1868, Henschen (in UPS). Chile.

Aconcagua, Matorral Zampallar, 200 m a.s.l. 1965, Foll-

mann 13418 (in UPS); Coquimbo, Matorral Fray Jorge.

1963 Follmann (in UPS). Ecuador. Azuay, road Sayausid-

Molleturo, 3000 m a.s.l. 1979, L. Arvidsson and D. Nilsson

1570 (in GB); Las Cajas, ca 20 km west of Sayausid. 1982,

M. Lindstro

¨m 972 (in GB); road Cuenca-Molleturo, 25 km

west of Cuenca, 3600 m a.s.l. 1983, L. and A. Arvidsson

4158 (in GB); road Sayausid-Molleturo, ca 10 km west

Sayausid, 3250 m a.s.l. 1972, L. Arvidsson and D. Nilsson

1146 (in GB). Cotopaxi, Volcan Cotopaxi, western slope,

34003500 m a.s.l. 1982, M. Lindstro

¨m 923 (in GB);

Parque Nacional Cotopaxi, north side of volcano Cotopaxi

(12 km from Panamerica Hwy), 0840?S, 78830?W, 3600 m

a.s.l. 1985, L. Arvidsson, M. Lindqvist and M. Lindstro

¨m

6288. Loja, ca 5 km south El Limo 800 m a.s.l. 1980, L.

Andersson 935 (in GB); road Loja-Zaruma, near Las

Chinches, 2400 m a.s.l. 1974, L. Andersson 340 (in GB);

road Loja-Catacocha, ca 2 km south of Las Chinches,

2400 m a.s.l. 1977, L. Andersson, U. Molau and M.

Neuendorf 532 (in GB); road Loja-Zaruma, near Las

Chinches, 2400 m a.s.l. 1974, L. Andersson 340 (in GB).

Napo, road Quito-Papallacta, 10 km east of the pass, 3500

m a.s.l. 1983, L. and A. Arvidsson 3538 (in GB). Pichincha,

5 km southeast Nono on road Quito-Nanegal, 0806?S,

78832?W 3250 m a.s.l. 1985, L. Arvidsson, M. Lindqvist

and M. Lindstro

¨m 5490 (in GB); road Pifo-Palallacta,

2900 m a.s.l. 1972, L. Arvidsson and D. Nilsson 1042 (in

GB). Peru. Cusco, Urubamba, Machu Picchu, outside the

ruin town, 13807?S, 72836?W, 2500 m a.s.l. 1981, A. Tehler

and G. Thor P87:13 (in UPS); Hua

´nuco, Cerro Carpish,

north of the tunnel, 9837?S, 76802?W, 2000 m a.s.l. 1981,

Santesson and Moberg P43:8 (in UPS).

Heterodermia flabellata

(Fe´e) D. D. Awasthi

(1973, p. 113)

Basionym: Parmelia flabellata Fe

´e (1837, p. 122). Lecto-

type (Swinscow and Krog 1976, p. 118): [South America]

ad corticem Cinconae lancifoliae (in G).

Taxonomic synonyms: Anaptychia hypoleuca var. fulvescens

Vain. (1913, p. 106), based on the same type: Anaptychia

fulvescens (Vain.) Kurok. (1960a, p. 93).

Literature: Kurokawa (1962, p. 52), Swinscow and Krog

(1976, p. 118, 1988, p. 95), Moberg (2004a, p. 260).

Thallus irregular or 9orbicular, robust, to 4 cm in dia-

meter, 9firmly adnate; upper surface greywhite to rarely

dark grey, 9glossy, not or rarely pruinose. Lobes to 2 mm

broad, radiating, sparsely divided, 9discrete, plane to

9convex. Soralia and isidia absent. Lower surface without

cortex, the lax medulla inspersed with a rust-coloured or

yellowishbrown pigment reacting Kpurple; rhizines

marginal, black apothecia common, to 5 mm in diameter,

substipitate with lobulate margins; ascospores Pachysporaria-

type, with sporoblastidia, (27)3039(45)(13.0)14.5

17.5(19.5) mm, M34.4; 15.9 mm, SD 4.46; 1.57 mm,

n48. Pycnidia 9common.

Chemistry:Kyellow. Atranorin, zeorin and leuco-

tylin, with rust-coloured or yellowish brown, Kpurple

pigment.

Habitat and distribution: Mainly on trees, but also on

rocks, in 9shady positions in tropical to warm temperate

areas. Seen from Argentina, Brazil, Ecuador, Peru and

Venezuela in South America. Also known from Africa, Asia,

North, and Central America.

Similar species

Heterodermia flabellata is characterized by its robust

appearance, the rust-coloured or yellowish brown pigment

(Kpurple) inspersed in the medulla on the lower surface

and the absence of soredia and isidia. A simple way to

observe the coloured lower side is to cut a lobule on the

apothecia. Similar species include the sorediate H. obscur-

ata, which might be regarded as the sorediate counterpart.

Kurokawa has described a similar species, Anaptychia

cyathiformis Kurok. (1973, p. 602) from two localities

in Natal which differs from H. fabellata only in chemistry

(it contains dissectic acid). I have seen just a fragment

136

of that species, and thus, leave it for further studies.

The combination Heterodermia cyathiformis (Kurok.) Trass

(1992, p. 10), is correctly regarded as nom. inval. in ‘Index

of Fungi’(Art. 34.1(b)). Heterodermia flabellata might be

the sorediate counterpart of H. obscurata.

Recent DNA studies of material from Costa Rica

(Lu

¨cking et al. 2008) reveal that H. flabellata and H. obscurata

form a well-supported clade and should thus be regarded

as conspecific. However, they suggest further studies to

solve the taxonomic status.

Heterodermia galactophylla

(Tuck.) W. L. Culb.

(1966, p. 482)

Basionym: Parmelia ciliaris b. galactophylla Tuck. (1847,

p. 224). Based on the same type: Anaptychia galactophylla

(Tuck.) Trev. (1861, p. 52). Holotype: USA, Maine, hb.

Oakes (in FH-Tuck).

Literature: Kurokawa (1962, p. 97), Moberg (1999, p. 4).

Thallus irregular to orbicular, to 5 cm in diameter, with

9elongate, distinctly separate lobes with long, prominent,

pale, marginal cilia; upper surface grey to whitish grey.

Lobes linear, 9dicotomously divided, to 1 mm wide, often

distinctly widening when soredia appear, spreading with

prominent, pale, marginal cilia which vary considerably in

density. Soralia rare to abundant, on the underside of the

lobe tips which thus mostly become distinctly recurved.

Lower surface without cortex. Apothecia and pycnidia

not seen.

Chemistry:Kyellow. Atranorin and zeorin.

Habitat and distribution: Mainly on tree trunk and

branches, often among mosses in tropical to warm tempe-

rate areas. Seen from Ecuador and Peru in South America.

Also known from North and Central America.

Similar species

The distinctive apical soralia and the pale, prominent,

marginal cilia are useful characters to identify H. galacto-

phylla. Very similar to H. allardii from which is differs only

in the presence of norstictic acid.

Heterodermia granulifera

(Ach.) W. L. Culb.

(1966, p. 482)

Basionym: Parmelia granulifera Ach. (1814, p. 212). Based

on the same type: Anaptychia granulifera (Ach.) A. Massal.

(1853, p. 41). Lectotype (as holotype in Kurokawa 1962,

p. 37): North America, Muhlenberg (in H-Ach 1404,

isotype: UPS-Ach).

Literature: Kurokawa (1962, p. 37), Moberg (1999, p. 4).

Thallus irregular, sometimes orbicular, to 3 cm in diameter,

firmly adnate, with isidia on margins and/or on surface; upper

surface whitish grey to grey, usually dull, 9distinctly

pruinose. Lobes radiating, to 1.5 mm broad, adnate to the

tips, 9sinuose, short and overlapping or discrete, flat to

convex, without projecting rhizines. Isidia marginal

and/or laminal, simple, very variable in length from knob-

like to 1 mm. Soredia may sometimes develop from broken

isidia or wartlike projections. Lower side corticate, white

to brown, with grey to brown simple rhizines. Apothecia

9common (not seen in South American material), shortly

stipitate, to 3 mm in diameter with scabrous thalline margin,

isidiose; ascospores Pachysporaria type, without sporoblasti-

dia, (20.5)21.027.0(30.0)(8.5)10.514.0(14.0) mm,

M22.1; 12.1 mm, SD2.99; 1.12 mm, n 24. Pycnidia

9common.

Chemistry:Kyellow0red. Atranorin, zeorin and salazinic

acid.

Habitat and distribution: Mainly on bark of various trees in

moist but open conditions in subtropical to warm tempe-

rate areas. Seen from Ecuador, Peru and Venezuela in South

Americas. Also known from North America.

Similar species

Heterodermia granulifera is recognized by the marginal/

laminal, simple isidia which easily break and sometimes

develop into limited soralia. A closely related species is

H. isidiophora which has mainly cylindrical to coralloid

isidia but lacks salazinic acid.

Heterodermia hypoleuca

(Ach.) Trevis.

(1869a ‘1868’, p. 615)

Basionym: Parmelia speciosa b. hypoleuca Ach. (1814,

p. 211). Based on the same type: Anaptychia hypoleuca

(Ach.) A. Massal. (1860, p. 249). Holotype: Am. Bor.,

¨hlenberg (in H-Ach 1306, isotype: UPS-Ach).

Literature: Kurokawa (1962, p. 42), Swinscow and Krog

(1976, p. 119), Moberg (1999, p. 5).

Thallus irregular to orbicular, to 5 cm in diameter, loosely

adnate with discrete, repeatedly branched lobes and at lobe

tips often small adventitious lateral lobes; upper surface grey

to dark grey or brownish, not or very weakly pruinose.

Lobes to 2 mm broad, usually around 1mm, irregularly

branched, tips mostly weakly pruinose. Soralia and isidia

absent. Lower surface without cortex, white to purple or

almost black, marginal cilia pale to black, 9abundantly

branched. Apothecia 9common, variable in size, to 5 mm

in diameter, sometimes cup-shaped, shortly stipitate or

sessile with lobulate margin; ascospores Pachysporaria type,

with or without sporoblastidia, (22.5)26.533.5(37.5)

(11.0)13.517.5(20.5) mm, M 30.0; 15.3 mm, SD 

3.58; 2.04 mm, n79. Pycnidia 9common.

Chemistry:Kyellow. Atranorin, zeorin, leucotylin and

9norstictic acid.

Habitat and distribution: On bark in fairly moist conditions

in subtropical to temperate areas. Seen from Brazi, Ecuador,

Peru and Uruguay in South America. Also known from

Africa, Asia and North America.

137

Similar species

The ‘woolly’upper lobe tips, the ecorticate lower side,

the discrete lobes with small adventitious lobes and the

lobulate apothecia define H. hypoleuca. It differs from

H. magellanica by having smaller ascospores and smaller,

less distinct sporoblastidia. H. hypoleuca could be charac-

terized as the fertile counterpart of H. japonica. It may

also be related to the isidiate species H. corallophora.

Heterodermia isidiophora

(Vain.) D. D. Awasthi

(1973, p. 114)

(non Anaptychia isidiophora (Nyl.) Vain. used by Kuro-

kawa (1962, p. 33), Swinscow and Krog 1976, p. 121).

Basionym:Anaptychia isidiophora Vain. (1901, p. 409).

Holotype: Angola, Pungo andongo (2400 ad 3800 ped.

s.m.). Ad rupes perpendiculares juxta rivulos ad Barranco

da Pedra Songue frequenter, at raro fruticans, Welwitsch

96, 1857 (in LISU, not seen, isotype in TUR-Vain 7741).

Literature: Swinscow and Krog (1976, p. 121), Moberg

(2004a, p. 261).

Thallus orbicular, to 7 cm in diameter, forming larger

colonies, 9loosely attached, provided with abundant isidia;

upper surface brightly grey, epruinose. Lobes narrow, to

1 mm wide, irregularly branched, 9plane, with short

lateral lobes, sometimes white at margins because of

broken isidia. Isidia cylindrical, simple or branched, mainly

along lobe margins, starting as small lobules, sometimes

abundant on lobe tips, in inner parts appearing laminal.

Lower surface corticate, dirty grey to brownish or almost

black with few, short rhizines. Apothecia sessile, to 4 mm

in diameter, margins isidiate; ascospores Pachysporaria-

type, with few sporoblastidia, (25.0)27.534.0(36.5)

(12.0)13.517.5(21.5) mm, M 30.7; 15.7 mm, SD 

3.18; 2.03 mm, n32. Pycnidia rare.

Chemistry:Kyellow. Atranorin, zeorin and leucotylin

with additional triterpenoids.

Habitat and distribution: Both corticolous and lignicolous,

but apparently preferring trees in open woodland and

wayside trees in subtropical and warm temperate areas. Seen

from Bolivia, Brazil, Colombia, Ecuador and Venezuela in

South America. Also known from Africa.

Similar species

Heterodermia corallophora is usually readily identified by its

cylindrical, usually elongate isidia. It differ from H.

antillarum (not seen from South America) by the 9flattened

isidia and the absence of salazinic acid. Easily separated from

H. corallophora by the corticate lower surface. Anaptychia

isidiophora (Nyl.) Vain. as treated by Kurokawa is Hetero-

dermia antillarum (Vain.) Swinsc. & Krog as it contain

salazinic acid (Swincow and Krog 1976, p. 114).

Heterodermia japonica

(Sato) Swinsc. & Krog

(1976, p. 122)

Basionym: Heterodermia dendritica (Pers.) Vain. var. japonica

(Sato 1936, p. 427). Based on the same type: Anaptychia

japonica (Sato) Kurok. (1960b, p. 353). Holotype: Formosa,

Mt Arisan, Mingetsu, 24 Jan 1936, M. Sato (in TI,

not seen).

Taxonomic synonyms: Heterodermia propagulifera (Vain.)

J. P. Dey (1976, p. 403), based on the same type:

Heterodermia dendritica var. propagulifera (Vain.) Poelt

(1965, p. 31). Anaptychia hypoleuca ssp. dendritica var.

propagulifera Vain. (1913, p. 107). Type: Philippines,

Luzon, Benguet. 2150 m a.s.l., Bur. Sci. 14071 Robinson

(in TUR).

Literature: Kurokawa (1962, p. 58), Swinscow and Krog

(1976, p. 122), Moberg and Purvis (1997), Moberg and

Nash (1999, p. 5), Moberg (2004a, p. 261).

Thallus irregular or rarely orbicular, very variable, to 7 cm,

loosely adnate; upper surface matt greyish to cream-

coloured, rarely brownish, sometimes pruinose at tips,

especially when young. Lobes radiating, to 2 mm wide, to

4 mm at tips, usually discrete, sometimes dissected with

lobules along the margin, occasionally only as scale. Soralia

labiate, on lateral or terminal lobes, sometimes spreading

along lobe margins, soredia farinose to granular. Lower

surface without cortex, white at apices, yellowish brownish

or bluish black centrally, sometimes sparsely dotted with

yellowish or brownish orangered pigment (Kpurple)

towards lobe apices; rhizines marginal, squarrose (or simple),

black. Apothecia sessile, to 4 mm in diameter, known only

from a few specimen; ascospores Pachysporaria-type, with

sporoblastidia, 40452022 mm. Pycnidia rare.

Chemistry:Kyellow or red, Atranorin and zeorin,

sometimes with norstictic, salazinic acid and unidentified

terpenes.

Habitat and distribution: On tree trunks or over mosses on

rocks in both open and shady situations. A pantropical to

subtropical species extending to warm temperate regions of

Europe.

Similar species

Heterodermia japonica is characterized by the dull upper

surface, the 9fan-like lobe apices, the absence of a lower

cortex, and the white to blackish violet lower surface. It is

however an exceedingly variable species in both morphology

and chemistry. The lobes may become more elongate in

shady habitats and then they may have distinct long, black,

marginal rhizines (to 7 mm). Soralia vary from farinose to

granular or may be virtually lacking. Swinscow and Krog

(1976) indicate such a variation in east African populations.

The sparse brownish orangered pigment present on the

lower lobe apices should not be confused with the more

distinct, continuous pigmentation of the lower surface of

H. obscurata.

Heterodermia dendritica var. propagulifera has previously

been discussed by Moberg and Nash (1999, p. 7) and

138

examination of the holotype in TURVain confirms its

inclusion here.

Heterodermia casarettiana (A. Massal.) Trevis. (1869a

‘1868’, p. 624) from South America falls well within the

variation of H. japonica both concerning morphology and

chemistry. As I have not seen the type in VER, I prefer not

to formalize the synonymy.

Heterodermia chilensis is very close to H. japonica and the

two could sometimes be difficult to separate. The only

difference seems to be the lobes; flat in H. japonica and

convex in H. chilensis. They may be conspecific but at

present they are kept separate.

Another member of the H. japonica complex, H. neglecta

Lendemer et al. (2007, p. 490), was described from North

America. It is obvious that it is necessary to study this

complex in the field, not only locally, as the variation both

in morphology and chemistry is considerable. Recent DNA

studies (Lu

¨cking et al. 2008) have also indicated that we

have several taxa within a morphologically very similar

group which needs further studies.

Heterodermia kurokawae

Trass (1992, p. 14)

(Fig. 6)

Nom. nov. for Anaptychia albicans Kurok. (1962, p. 80).

Holotype: Peru, Dept Cuzco, Hda Choquelhuanca,

2200 m a.s.l. Sobre cafe

´viejo, Mar 1919, C. Bues 692 (in

FH).

Thallus irregular, loosely attached to substratum, dichot-

omously branched; upper surface pale grey, with pale,

marginal cilia. Lobes linear, elongate, 12 mm broad, not

widening at tips. Soralia and isidia absent, but the farinose

lower side appears sorediate. Lower surface without cortex,

with yellow pigment (not very obvious). Apothecia to

6 mm broad with lobate margins with the yellow pigment

visible; ascospores (holotype) of Pachysporaria-type, with

sporoblastidia, 35441922 mm.

Chemistry:Kyellow. Atranorin, zeorin and a yellow

pigment (K).

Habitat and distribution: On trees and rocks in open

situations. Seen only from a few localities in Ecuador and

Peru. Also known from Central America.

Similar species

Heteriodermia kurokawae is very distinct by the linear

9elongate lobes with pale 9abundant cilia and the yellow

pigment on the underside. The type material is close to

H. appalachensis (Kurok.) W. L. Culb. (1966, p. 479). It

could be the fertile counterpart of that species which also

occasionally has a yellow pigmentation on the underside.

Another species with yellow pigment on the underside is

H. lutescens but it has much narrower lobes and the

marginal cilia are thin and black. Yet another species with

pigmented underside (reddish) and elongate, linear lobes

is H. vulgaris, but it has much narrower lobes and black

cilia.

Specimens examined:Ecuador. Napo, road Baeza-Teno,

5 km south of Baeza, 1800 m a.s.l. 1983, L. and A.

Arvidsson 3811. (in GB). Pichincha, Alluriquin, the east

side of Rio Pilato

´n, 0819?S, 78859?W, 850 m a.s.l. Pasture

with solitary trees, Saxicolous on exposed rocks 1985,

Lindstro

¨m and Lindqvist 2186, 2217 (in GB). Tungur-

ahua, Rio Negro, 1823?S, 78812?W, 1250 m a.s.l. 1972,

L. Arvidsson and D. Nilsson 696 (in GB); Pastaza Valley,

between Banos and Rio Verde, 1500 m a.s.l. 1983, L. and

A. Arvidsson 4082 (in GB). Peru. Hua

´nuco, Hua

´nuco,

between Mallqui and Puente Cayumba, 9830?S, 75853?W,

1200 m a.s.l. 1981, Santesson and Moberg P37:34 (in

UPS); ca 18 km (road distance) northnortheast of Tingo

Maria, Puente Pumahuasi, 9805?S, 75855?W, 700 m a.s.l.

1981, Santesson and Moberg P38:17 (in UPS).

Heterodermia leucomela

(L.) Poelt (1965, p. 31)

Basionym: Lichen leucomelos L. (1763, p. 1613). Holotype:

America (in LINN 1273.109). Taxonomic synonyms:

Heterodermia leucomela ssp. boryi (Fe

´e) Swinsc. and Krog

(1976, p. 124), based on the same type: Borrera boryi

´e (1824, p. XCVI and Table II, Fig. 23). Anaptychia

neoleucomelaena Kurok. (1961, p. 51).

Literature: Kurokawa (1962, p. 74), Swinscow and Krog

(1976, p. 124, 1988, p. 95), Moberg and Nash (1999,

p. 7), Moberg (2004a, p. 262).

Thallus appearing fruticose of irregular, narrow, interwoven

lobes, sometimes several cm long, with long cilia (rhizines)

along the margins; upper surface whitish to cream-

coloured, shiny without pruina. Lobes to 1.0(1.5) mm

wide, parallel-sided, not widening towards apices, dichot-

omously branched with long, sparsely squarrose rhizines,

usually black except for the pale base. Soralia irregularly

Figure 6. Heterodermia kurokawae. Peru, 1981, R. Santesson and

R. Moberg P37:34 (in UPS). Scale bar2 mm.

139

formed on the underside causing the lobes to widen. Lower

surface without cortex, white or rarely purple, margins

corticate and prominent. Apothecia very rare, 2(3) mm in

diameter, stipitate, disc black,9pruinose, margin crenulate

to lobulate, lobes up to 10 mm; ascospores of Pachyspor-

aria-type, with sporoblastidia, (31.0)34.542.5(49.5) 

(15.0)16.020.0(23.5) mm, M38.5; 18.1 mm, SD

4.10; 1.95 mm, n48. Pycnidia rare.

Chemistry:Kyellow. Atranorin and zeorin.

Habitat and distribution: On trees and over mosses on rocks

in both moist and fairly dry situations. Seen from

Argentina, Bolivia, Brazil, Chile, Ecuador and Peru in

South America. One of the most widespread and abundant

species throughout tropical and subtropical regions extend-

ing also to warm temperate areas and known from Africa,

Asia, Australia and North America.

Similar species

Heterodermia leucomela is easily recognized by the entangled

mats of elongate, linear lobes with long black cilia and the

ecorticate lower surface with 9thick corticate margins. May

be confused with H. lutescens which has a yellow pigment on

the underside and H. vulgaris with a reddish pigment.

Another similar species is H. fertilis which lacks soralia on

the underside but is richly fertile. Specimens with curved lob

tips have sometimes been regarded as a separate taxon (ssp.

boryi), but as I have found all types of transitional states these

are regarded as falling within the range of the variation of the

species. Heterodermia neoleucomelaena was synonymized with

ssp. boryi by both Kurokawa (1973, p. 587) and Swinscow

and Krog (1976, p. 124) and is here treated as a synonym of

H. leucomela.

The use of the epithet lecomelos (leucomelas) is discussed

in Swinscow and Krog (1976, p. 124).

Heterodermia lutescens

(Kurok.) (Follman 1974,

p. 73)

Basionym: Anaptychia lutescens Kurok. (1961, p. 54).

Holotype: Mexico, Monte Ovando, Chipas, 1932, Mat-

suda 46 (in TNS).

Literature: Kurokawa (1962, p. 79), Swinscow and Krog

(1976, p. 129), Moberg and Purvis (1997, p. 191).

Thallus irregular, small, with 9erect lobes with black

marginal cilia; upper surface yellowish white to brownish,

rarely darker brown, 9dull and without pruina. Lobes

narrow, ca 0.5(1.0) mm, 9dichotomously branched, cilia

thin, simple and black, sometimes sparsely squarrose.

Soralia 9terminal, on the underside; soredia granular,

9yellowish. Lower surface without cortex with thick,

corticate and prominent margins, white and mostly with a

K, yellow pigment which sometimes turn reddish in inner

parts. Apothecia rare, subapical, substipitate with lobulate

margins; ascospores of Pachysporaria-type, with sporoblasti-

dia, (37.5)39.547.0(47.0)(20.5)21.024.0(25.0)

mm, M43.4; 22.4 mm, SD 3.72; 1.60 mm, n 8.

Pycnidia rare.

Chemistry:Kyellow. Atranorin, zeorin and a pigment (K).

Habitat and distribution: Corticolous and terricolous over

mosses and soil in fairly shaded positions. Seen from

Bolivia, Brazil, Ecuador, Peru and Venezuela. It is widely

distributed in tropical to subtropical areas, and is known

from the Azores, Africa, Asia and North and Central

America.

Similar species

Heterodermia lutescens is easily distinguised in the field

by the neat mats of small, erect yellow-tinged lobes with

yellow underside and thin, black cilia. Resembles small

individuals of H. leucomela which differ by their grey-white

colour and more robust cilia. Heterodermia kurokawae

also has a yellow underside, but the lobes are usually

much broader and its marginal cilia are pale.

Heterodermia magellanica

(Zahlbr.) Swinsc. &

Krog (1976, p. 130)

Basionym: Anaptychia magellanica Zahlbr. (1917, p. 54).

Holotype: Magellan’s sund, Isla Felix, 24 May1908, C.

Skottsberg (in UPS).

Literature: Kurokawa (1962, p. 66), Swinscow and Krog

(1976, p. 130).

Thallus irregular to orbicular, to 5 cm in diameter, 9loosely

adnate with discrete, repeatedly branched lobes and at lobe

tips small adventitious lateral lobes; upper surface grey to

dark grey or brownish, not or weakly dull pruinose. Lobes to

2 mm broad, usually around 1 mm, irregularly branched,

mostly ‘woolly’tips, in some situations (high altitudes) they

may be short and ascending. Soralia and isidia absent. Lower

surface without cortex, white to purple or rarely black,

rhizines marginal, pale to black, 9abundantly branched.

Apothecia 9common, variable in size, to 5 mm in diameter,

sometimes cup-shaped, shortly stipitate or sessile usually with

distinctly lobulate margins; ascospores of Pachysporaria-

type, with prominent sporoblastidia, (33.5)39.047.5(

54.0)(16.0)17.021.5(25.0) mm, M43.1; 19.3 mm,

SD4.30; 2.14 mm, n64. Pycnidia 9common.

Chemistry:Kyellow. Atranorin, zeorin, leucotylin and

9norstictic acid. The chemistry is variable as in H.

japonica and H. hypoleuca.

Habitat and distribution: On mossy tree trunks in fairly

moist conditions in tropical and subtropical areas usually

at high altitudes. Seen from Argentina, Brazil, Ecuador,

Peru and Venezuela. Also known from east Africa and

Central America.

140

Similar species

Heterodermia magellanica belong to a species complex which

is in need of further field studies. It is characterized by the

‘woolly’upper lobe tips, the ecorticate lower surface, the

discrete lobes with small adventitious lobes and usually

strongly lobulate apothecia. Very similar to H. hypoleuca,

which has smaller ascospores. Without ascospores the two

species can not be separeted with certainty. It could be

characterized as the fertile counterpart of H. japonica. It may

also be related to the isidiate species H. corallophora. Marcelli

et al. (2007, p. 221) published a new species from Brazil,

Heterodermia kalbii M. F. N. Martins & Marcelli, which

belongs to the same complex and they referred to a forth-

coming paper dealing with the H. magellanica-complex.

Heterodermia dactyliza (Nyl.) Swinsc. & Krog (1976,

p. 117) is separated by a distinct cortical margin visible

from the noncorticate lower surface. But in my opinion,

the type material in (G) can hardly be separated from the

variable H. magellanica.

Heterodermia multiciliata

(Kurok.) Trass (1992, p.

16) (Fig. 7)

Basionym: Anaptychia multiciliata Kurok. (1962, p. 72).

Holotype: Chile, Coquimbo, La Serena, Cerro los Loros.

27 Jun 1940, Santesson 2521 (in S, isotype: UPS).

Thallus irregular and very variable with elongate, sparsely

dichotomously divided lobes and abundant black marginal,

9laminal cilia; upper surface grey, often bullate and provided

with abundant pycnidia. Lobes elongate, 12 mm wide,

sometimes wider. Soralia and isidia absent. Lower surface

without cortex, occasionally the medulla develops soredia.

Apothecia subapical, shortly stipitate, 9abundant, sparsely

lobulate with prominent black cilia from small lobules;

ascospores of Pachysporaria-type, with sporoblastidia,

(34.5)35.541.0(45.0) (17.0)18.021.5(23.5) mm,

M38.4; 19.8 mm, SD2.87; 1.70 mm, n 24. Pycnidia

often very abundant.

Chemistry:Kyellow. Atranorin and zeorin.

Habitat and distribution: On shrubs and branches in fairly

open situations, also on open rocks. So far known only from

Chile and Venezuela.

Similar species

The elongate lobes and the black cilia on the upper surface

of the thallus and apothecia make H. multiciliata easily

recognized. May be confused with H. fertilis which also has

black cilia on the apothecial margins, but the latter species

has much longer lobes and lobate apothecia, and the

ascospores are multiblastidiate. It has also some similarities

to the Californian Heterodermia erinacea (Ach.) W. A.

Weber (1987, p. 163), but the the latter has mostly pale

cilia and no cilia on the apothecia and the ascospores are

smaller and without sporoblastidia.

Sipman (1995, p. 329) described two new species,

Heterodermia pinnata Sipman and H. follmannii Sipman

closely related to H. multiciliata. Among the material I have

seen, only one specimen of each species agrees with the

description. They are both from Chile, Maule, between Las

Canas and Punta Santa Ana, 35833?S, 72834?W, 50 m a.s.l.

2001, Moberg 12236a and b (in UPS). Heterodermia

pinnata has narrower, linear lobes and H. follmannii is

smaller with soralia on the underside of the lob tips. I agree

with Sipman that they are very closely related to

H. multiciliata and they might only be modifications.

Specimens examined (selected). Chile. Coquimbo, at the

border of Fray Jorge National Park, 30839?S, 71837?W,

320 m a.s.l. 2001, Moberg 12204 (in UPS); Parc Nacional

Fray Jorge. 1966, Redon (in TUR); 19.2 km north of La

Figure 7. Heterodermia multiciliata. Chile, 1940, Santesson 2918 (in UPS). Scale bar2 mm.

141

Serena, on coastal slopes just north of Punto Arrayan, 50

80 m a.s.l. 1974, Rundel 9858 (Weber, Lich. exs. 489, in

UPS); Santiago, near San Pedro, km-post 115, 33847?S,

71833?W, 300 m a.s.l. 2001, Moberg 12226a (in UPS);

Valparaı

´so, Los Molles, 32815?S, 71832?W, 20 m a.s.l.

2001, Moberg 12202 (in UPS); Cartagena, 33832?S,

71835?W 1940, Santesson 2482 (in UPS). Ecuador. Loja,

7 km from Macara

´along road to Catacocha, 4818?S,

79855?W, 9001000 m a.s.l. 1987, B. Sta

˚hl, M. Neuen-

dorf, J.-E. Bohlin and R. Lundin 436 (in GB). Venezuela.

Bolivar, Macizo del Chimanta, 5816?N, 62809?W, 2100 m

a.s.l. 1985, Ahti, Huber and Pipoly 42239 (in H).

Heterodermia obscurata

(Nyl.) Trevis.

(1869b, p. 114)

Basionym: Physcia obscurata Nyl. (1863, p. 310). Lectotype

(as holotype in Kurokawa 1962, p. 49): Colombia, Lindig

704 (in H-NYL 32570).

Literature: Swinscow and Krog (1976, p. 132), Moberg and

Nash (1999, p. 9), Moberg (2004a, p. 264).

Thallus irregular or 9orbicular, robust, 5 cm in diameter,

9firmly adnate; upper surface greywhite to rarely dark

grey, 9dull, not or rarely pruinose. Lobes to 2 mm, usually

1 mm, radiating, sparsely divided, 9discrete, flat to convex.

Soralia labriform on lateral and terminal lobes, rarely

confluent and forming marginal soralia, sometimes rusty

brown coloured by the medullary pigment; soredia gran-

ular. Lower surface without cortex, the lax medulla

inspersed with rust-coloured or yellowish brown, K

purple pigment; with black marginal cilia. Apothecia not

common, sessile to substipitate, margin 9lobulate, often

with sorediate margins; ascospores of Pachysporaria-type,

with or without sporoblastidia, (28.0)31.034.5(34.5)

(15.0)15.519.5(21.5) mm. M32.8; 17.5 mm. SD 

1.76; 1.87 mm. n 16. Pycnidia rare.

Chemistry: Kyellow. Atranorin, zeorin and a pigment

(Kpurple).

Habitat and distribution: Mostly on mossy rocks but also on

mossy trees in 9sheltered sites. Common and widespread

in tropical and subtropical areas, extending also to warm

temperate regions. Seen from Brazil, Chile, Ecuador, Peru

and Venezuela in South America. Also known from Africa,

Asia, Europe, Central and North America.

Similar species

Heterodermia obscurata is characterised by its robust appear-

ance, the distinctive labriform soredia and the rusty brown

pigmented (Kpurple), ecorticate lower surface. Similar

species include H. japonica which differs by lacking the rusty

pigment in the medulla and by having a bluish underside in

inner parts. Another similar species, H. speciosa, differs by

having a lower cortex. Heterodermia obscurata might be the

sorediate counterpart of H. flabellata.

Recent DNA studies of material from Costa Rica

(Lu

¨cking et al. 2008) reveal that H. obscurata and H.

flabellata form a well-supported clade and should thus be

regarded as conspecific. However, they suggest further

studies to solve the taxonomic status.

Heterodermia palpebrata

(Tayl.) Vain. (1898, p. 38)

Basionym: Parmelia palpebrata Tayl. (1847, p. 173).

Lectotype (Kurokawa 1962, p. 71): Peru, Cuming 1467

(in FH-Tayl, seen as photo).

Thallus irregular, to 7 cm broad, but usually smaller with

ascending lobes but not recurved lobe tips; upper surface

greyish. Lobes 9convex, variable in width, up to 2 mm,

sometimes ascending, with white to dark cilia along margins.

Soralia and isidia absent. Lower surface without cortex and

appearing sorediate with prominent corticate margin.

Apothecia 9abundant, laminal, shortly stipitate, with 9

well-developed ciliate lobules on margins; ascospores of

Pachysporaria-type, with sporoblastidia, (27.0)31.035.5(

36.5)(15.0)16.519.5(20.5) mm. M33.1; 17.8 mm.

SD2.19; 1.56 mm. n 24, Pycnidia not seen.

Chemistry:Kyellow. Atranorin and zeorin.

Habitat and distribution: On branches and twigs in fairly

open situations. Only known from a few localities from

Figure 8. Heterodermia parva. A. Peru, 1981, Santesson, Tehler

and Thor P106:23 (in UPS). B. P106:2 (in UPS). Scale bar

1 mm.

142

Peru in South America and one in USA, Texas (UPS,

already identified by Kurokawa 1961)

Similar species

Heterodermia palpebrata is similar to some modifications of

H. podocarpa (see that species) and Heterodermia erinacea

(Ach.) W. A. Weber (1987, p. 163), but the latter has long,

prominent marginal cilia, apothecial margins that are not

lobate, and much smaller spores.

Specimens examined: Peru. Hampe dedit (in UPS). USA.

Texas, Hampe dedit (in UPS).

Heterodermia parva

Moberg sp. nov. (Fig. 8)

Thallus parvus,12 mm in diametro,saepe absens;apothecia

stipitata,erecta,ad 4 mm elata.

Holotype: Peru, Cusco, Paucartambo, road Paucartambo-

Pillcopata, 1 km southwest of the turn to Tres Cruces,

13808?S, 71838?W, 3500 m a.s.l. On twigs. 29 Mar 1981,

Santesson, A. Tehler and G. Thor P108:2 (in UPS, isotypes

will be distributed in Moberg, Lich. sel. exs. Upsal.).

Thallus very small, 12 mm but often absent and then

consisting of stipitate apothecia only. Upper surface grey.

Lobes 0.10.2 mm. Soralia and isidia absent. Lower surface

without cortex, pale in outer parts, blackening near

apothecia. Apothecia stipitate, erect, often crowded, to

4 mm high, to 3 mm in diameter with smooth to lobulate

margins; ascospores of Pachysporaria-type, without

sporoblastidia, (28.0)29.033.5(33.5)(13.0)15.017.5

(18.5) mm. M31.0; 16.4 mm. SD2.33; 1.27 mm. n 

16. Pycnidia not seen.

Chemistry:yellow. Atranorin and zeorin

Habitat and distribution: Among mosses and other lichens

on twigs and branches but also on thin soil and in cracks on

rocks. So far known only from high altitudes (around 3500

m a.s.l.) in the Dept of Cuzco in Peru.

Notes: The small size makes it difficult to find, but when

observed it is easily identified by the stipitate, often

crowded apothecia.

Specimens examined (paratypes, all in UPS): Peru. Cusco,

Paucartambo, road Paucartambo-Pillcopata, 1 km northeast

of the turn to Tres Cruces, 13807?S, 71837?W, 3500 m

a.s.l. 1981, Santesson, Tehler and Thor P106:2, P106:23; 1

km southwest of the turn to Tres Cruces, 13808?S,

71838?W, 3500 m a.s.l. 1981, Santesson, Tehler and

Thor P108:32; road Paucartambo-Pillcopata, just southwest

of Paso de Tres Cruces, 13808?S, 71838?W, 3450 m a.s.l.

1981, Santesson, Tehler and Thor P96:47, P96:48.

Heterodermia podocarpa

(Be´l.) D. D.Awasthi

(1973, p. 114)

Basionym: Parmelia podocarpa Be

´l. (1840, p. 122, Plate 13,

Fig. 1). Type: Sur les troncs pourrais des arbres, dans les

bois du quartier Sainte-Suzanne (20854?S, 55837?E), a l’ile

de Bourbon (Re

´union) Plate 13, Fig. 1 (sic!). Epitype:

Tanzania, Tanga Prov., Usambara Mts, Amani, in the

surroundings of the Forestry House, 5807?S, 38838?E,

900 m a.s.l., on Mangifera indica by Forestry House,

1971, Moberg 1491e (in UPS).

Taxonomic synonyms: Heterodermia barbifera (Nyl.) Kr.

P. Singh (1979, p. 221). Anaptychia stellata (Vain.)

Kurok. (1962, p. 90).

Literature: Kurokawa (1962, p. 86), Swinscow and Krog

(1976, p. 133), Moberg and Nash (1999, p. 9), Moberg

(2004a, p. 264).

Thallus irregular, to 5 cm broad, but usually smaller with

9ascending lobes terminating in squamulose apothecia;

upper surface grey, sometimes reddish caused by poor drying.

Lobes 9convex, variable in width, up to 2 mm, often

ascending, sometimes imbricate, with whitish cilia along

margins. Soralia and isidia absent. Lower surface without

cortex and appearing sorediate. Apothecia 9abundant,

terminal and stalked, with 9well-developed lobules on

margin; ascospores Pachysporaria-type, with sporoblastidia,

(27.0)32.038.5(41.0)(13.0)15.019.0(21.5) mm.

M35.0; 16.9 mm. SD3.25; 2.08 mm. n 40. Pycnidia

usually abundant.

Chemistry:Kyellow. Atranorin, zeorin, leucotylin and 9

norstictic acid.

Habitat and distribution: Mainly on branches and twigs in

fairly open situations. A widespread but rare species. Seen

from Bolivia, Brazil, Chile, Ecuador, Peru and Venezuela in

South America. Also known from Africa, Asia and Central

and North America.

Similar species

Heterodermia podocarpa is recognized by the lobulate

apothecia on tips of 9erect lobes. It is closely related to

H. palpebrata which has laminal apothecia. Also H.

galactophylla has terminal apothecia but differs by having

labriform soralia. Heterodermia barbifera is here treated

as a synonym because it fits well within the variation range

of H. podocarpa both morphologically and chemically.

The density of rhizines and their branching is not a

reliable character as it can vary within one and the same

thallus.

Following Swinscow and Krog (1976, p, 133) also

Anaptychia stellata (Vain.) Kurok. is here treated as a

synonym. The chemistry agrees and the differing location

of the apothecia is merely a matter of development.

Heterodermia pseudospeciosa

(Kurok.) W. L. Culb.

(1966, p. 484)

Basionym: Anaptychia pseudospeciosa Kurok. (1959,

p. 176). Holotype: Japan, Honshu, Idzu Prov., Suishochi,

Amagi Pass, 20 Aug 1956, Asahina (in TNS, not seen).

Literature: Kurokawa (1962, p. 25), Swinscow and Krog

(1976, p. 134), Moberg and Nash (1999, p. 10).

Thallus orbicular to irregular, small, less than 4 cm in

diameter, loosely adnate, densely lobate; upper surface grey

to brownish grey, darker at lobe tips, sometimes weakly

143

pruinose. Lobes long and narrow, to 1 mm broad, repeat-

edly branched, weakly convex, widening, tips not ascending.

Soralia semi-capitate, white to bluish grey, sometimes at

lobe tips but mainly arising from small lateral knob-like

structures. Lower surface corticate, whitish to pale brown-

ish, rarely dark grey, weakly rhizinate with usually short

(ca 1 mm) pale to dark brown or black rhizines. Apothecia

not seen. Pycnidia 9common.

Chemistry:Kyellow. Atranorin, zeorin, norstictic acid

and 9unidentified terpenes. The records of salazinic acid

are probably misinterpretations of connorstictic acid which

seems to be regularly present. Neither the type material

(Swinscow and Krog 1976, p.134) nor the material I have

examined does not contain salazinic acid.

Habitat and distribution: Mainly on rocks in fairly moist

situations in tropical to warm temperate areas. Seen from

Brazil, Peru and Venezuela in South America. Seemingly

rare and also known from Africa and Central and North

America.

Similar species

Heterodermia pseudospeciosa is characterised by the narrow,

convex, long lobes and the corticate lower surface.

The soralia develop mainly from marginal knob-like

structures forming small, semi-capitate soralia. Heterodermia

albicans also has similar soralia, but they are 9continuously

marginal. Further, H. albicans is more firmly adnate and

contains salazinic acid. The Kyellow 0red reaction in

H. albicans (salazinic acid) can be used to separate

H. pseudospeciosa (norstictic acid) as it gives a Kyellow

reaction only, even though norstictic acid in other genera

reacts Kyellow0red. Heterodermia speciosa is usually

larger, has 9flat lobes and lacks both norstictic and salazinic

acid.

Heterodermia spathulifera

Moberg & Purvis

(1997, p. 192)

Holotype: The Azores, St Michel, Furnas, on a tree in the

park. 3 Apr 1937, H. Persson no. H2 (in UPS).

Literature: Moberg and Nash (1999, p. 11).

Thallus irregular to orbicular, to 7 cm in diameter; upper

surface white to cream-coloured, glossy and without pruina.

Lobes narrow, to 1 mm, discrete to adjacent, usually firmly

adnate, 9plane, bearing 9pale, marginal rhizines visible

from above. Soralia labriform, sometimes becoming very

large (to 5 mm) and spathulate. In moist and shaded

habitats the soredia tend to develop into squamules which

sometimes may cover inner parts of the thallus. Lower

surface semi-corticate, white on outer parts of lobes, pale

brown centrally (appearing corticate, but in sections this

proves to be medulla incrusted by soil particles); rhizines

sparse, simple, white or cream-coloured, sometimes appear-

ing as cilia on the margins. Apothecia rare, only one

seen with either immature or over-mature spores, 3643

1518 mm. Pycnidia not seen.

Chemistry: Kyellow. Atranorin, zeorin, 9lecotylin and

‘spathulin’(16b-acetoxyhopan-22-ol).

Habitat and distribution: Mainly on tree trunks and twigs,

rarely on rocks, in fairly open situations in subtropical areas.

Seen from Brazilia, Ecuador and Peru. Also know from

South Africa, the Azores, Mexico, New Zealand. New to

South America.

Similar species

Heterodermia spathulifera is distinguished by its narrow,

glossy, firmly adnate lobes with pale marginal rhizines

and by containing spathulin. When present soralia are

usually large and spathulate. Might be confused with

H. albicans and H. speciosa which both differ in having

a lower cortex and different chemistry.

Heterodermia speciosa

(Wulf.) Trevis. (1869a ‘1868’,

p. 614)

Basionym: Lichen speciosus Wulf. (1789, p. 119). Lecto-

type (Moberg 2002, p. 71): [Austria] In sylvarum montis

Calvariae saxis, (Wulfen), Collectanea 3, T. 7. Epitype:

Switzerland, Tessin, 5 km south of Bellinzona, 1934,

Degelius (in UPS).

Taxonomic synonym: Heterodermia tremulans (Mu

¨ll. Arg.)

W. L. Culb. (1966, p. 485). Literature: Kurokawa (1962, p.

24), Swinscow and Krog (1976, p. 134), Moberg and Nash

(1999, p. 12), Moberg (2004a, p. 267) (as Heterodermia

tremulans).

Thallus orbicular to irregular, 23(4) cm in diameter,

often coalescing with other thalli, 9firmly adnate; upper

surface white to cream-coloured or brownish grey,

appearing bluish through the dense soralia, 9shiny, the

lobe tips sometimes darkening, very rarely pruinose.

Lobes narrow, ca 1(2) mm, flat to convex, end-lobes

usually without soralia, slightly widening towards

apices. Soralia abundant, labriform on lateral lobes,

sometimes semi-capitate, starting from lateral lobules,

Figure 9. Heterodermia spinigera. Ecuador, 1987, Neuendorf,

Bohlin and Lundin 439 (in GB). Scale bar 2 mm.

144

soredia farinose to 9granular, grey to bluish grey. Lower

cortex present, pale to dark brown with few scattered,

short and robust, usually black rhizines. Apothecia 9

common, to 3(5) mm in diameter and with marginal,

sorediate lobules; ascospores of Pachysporaria-type, with-

out sporoblastidia, (22.5)29.036.5(43.0)(11)12

17(28) mm. M32.7; 14.4 mm. SD3.86; 3.86 mm.

n54. Pycnidia not seen.

Chemistry: Kyellow. Atranorin, zeorin, 9leucotylin,

unidentified triterpene.

Habitat and distribution: In fairly open situations, often

sunny places in mountainous areas, mainly on rocks

but also on tree trunks and twigs. Widely distributed

in subtropical to temperate areas, extending as far north

as northern Scandinavia (Moberg 2002, p. 26). Seen

from Bolivia, Brazil, Chile, Ecuador, Peru and Venezuela.

Also known from Africa, Asia, Europe and North

America.

Similar species

Heterodermia speciosa is characterised by the striking

contrast between the bluegrey soralia and the paler thallus,

and the presence of a lower cortex. The lower surface of

H. obscurata lacks a cortex and has a rusty orange pigment.

Heterodermia albicans is much smaller and contains salazi-

ninc acid (Kyellow0red). Heterodermia tremulans was

earlier treated as a separate species by myself (Moberg

2004a, p. 267), but with experience from South America,

I now agree with Swinscow and Krog (l. c.) that it is

conspecific with H. speciosa.

Heterodermia spinigera

(Kurok.) Moberg comb. nov.

(Fig. 9)

Heterodermia spinigera (Kurok.) Trass cited in Index

Fungorum (nom. inval.) was never made as is stated in

the ‘Abstract of Trass’(1992, p. 2). Basionym: Anaptychia

spinigera Kurok. (1962, p. 66). Holotype: Peru, on rocks

[no precise locality], W. Lobb (holotype: BM).

Thallus foliose, irregular to orbicular or fruticose, to 7 cm in

diameter; upper surface white to cream-coloured, without

pruina, sometimes with a few cilia on surface (not to be

confused with the ciliate primordia of apothecia). Lobes 9

linear, elongate, to 3 mm wide, discrete, losely adnate,

usually plane, not distinctly widening at tips, with long,

pale, darkening marginal cilia. Soralia and isidia absent.

Lower surface without cortex, white and often canaliculate.

Apothecia common, laminal, lobate with marginal cilia, the

underside of apothecia distinctly ciliate, the young, devel-

oping apothecia distinctly and obvious ciliate; ascospores

of Pachysporaria-type, with small sporoblastidia, 3040 

1421 mm. Pycnidia 9common.

Chemistry: Kyellow. Atranorin and zeorin.

Habitat and distribution: On trees and rocks in fairly shaded

situations. Seen from Peru (type) and Ecuador in South

America, the only known localities.

Similar species

Heterodermia spinigera is easily recognized by the

elongate, 9flat, sparsely ciliate lobes and the laminal,

9lobate apothecia which already at young stages are

densely ciliate. The ciliate H. comosa has convex lobes

with apical apothecia and well-developed material has a

yellow pigment on the underside.

Specimen examined: Ecuador. Loja, 7 km from Macara

along road to Catacocha, 4818?S, 79855?W, 9001000 m

a.s.l. 1987, B. Sta

˚hl, M. Neuendorf, J.-E. Bohlin and R.

Lundin 439 (in GB).

Heterodermia squamulosa Degel. (1941, p. 76)

Holotype: USA, North Carolina, Great Smoky Mountains,

near Newfound Gap, 1540 m a.s.l., on Fagus grandifolia,

1939, Degelius (in UPS).

Literature: Kurokawa (1962, p. 60).

Thallus irregular to orbicular, to 5 cm in diameter, 9

loosely adnate, narrow-lobed and squamulose; upper sur-

face grey to cream-coloured or brownish particularly on

lobe tips, 9shiny, without pruina. Lobes narrow, to 1 mm

but usually narrower, repeatedly divided to squamulose,

discrete to overlapping. Squamules at lobe tips, also

marginal, sometimes laminal, occasionally developing gran-

ular soredia particularly in shade. Lower surface without

cortex, white to brownish or bluish centrally, cilia along the

margins, black, simple to squarrose, often forming a dense

felt under the lobes. Apothecia not seen. Pycnidia rare.

Chemistry: Kyellow. Atranorin, zeorin, 9leucotylin and

other triterpenes.

Habitat and distribution: Over mosses on bark and rocks (cf.

also H. japonica). Seen from Argentina, Brazil, Ecuador,

Peru and Venezuela in South America. Also known from

North America.

Figure 10. Heterodermia trichophora. Brazil, 1886, Schenk 4436

(holotype). Scale bar1 mm.

145

Notes: Recognized by its narrow lobes with apical, marginal

and occasionally also laminal squamules. Similar and related

to H. japonica and may be just a modification of that species.

Heterodermia subcitrina Moberg (2004a, p. 266)

Holotype: South Africa. Eastern Cape Province, Soutrivier,

ca 1 km westsouthwest of Nature Valley, 33859?S, 23832?E,

ca10 m a.s.l., on Pterocelastrus tricuspidata on the seashore

in the inlet, 11 Dec 1996, Moberg 11847 (in UPS).

Thallus irregular or 9orbicular, to 5 cm in diameter, 9

loosely adnate; upper surface greywhite to rarely dark

grey, 9dull, not or rarely pruinose. Lobes ca 12 mm wide,

radiating, 9discrete, flat to slightly concave, margins

sometimes folded upwards showing the yellowish lower

surface. Soralia marginal, labriform, sometimes confluent

along the margins, sometimes coloured yellowish by the

medullary pigment, soredia granular. Lower surface without

cortex, the 9lax medulla inspersed with the yellowish

pigment, Kpurple, with black marginal cilia.

Apothecia 9common, to 5 mm in diameter, sessile or

shortly stipitate, margins usually covered by granular

soredia, the yellow pigment sometimes visible; ascospores

of Pachysporaria-type, with sporoblastidia, (27.0)30.0

34.0(37.5)(12.0)13.517.0(19.5) mm. M32.0;

15.2 mm. SD2.24; 1.84 mm. n 34. Pycnidia rare.

Chemistry: Kyellow. Atranorin, zeorin, 9leucotylin and

a pigment (K).

Habitat and distribution: Mainly on trees in fairly moist

situations but also on mossy rocks in subtropical to warm

temperate areas. Seen from Brazil and Uruguay in South

America. New to South America. Elsewhere known from

South Africa, but may well be overlooked and should be

searched for.

Similar species

Heterodermia subcitrina is characterized by the marginal,

labriform, often confluent soralia, and the concave lobes,

usually having a yellowish pigment on the ecorticate lower

surface. The similar H. obscurata differs by having convex

lobes, terminal, labriform soralia and a rusty brown

pigment on the lower surface.

Heterodermia trichophora

(Kurok.) Trass

(1992, p. 21) (Fig. 10)

Basionym: Anaptychia trichophora Kurok. (1962, p. 100).

Holotype: Brazil, Rio de Janerio, Minas, Serra do Picu, bei

det Barra, an Gestra

¨uch, 11 Dec 1886, H. Schenk 4436 (in

UPS).

Literature: Kurokawa (1962, p. 100).

Thallus irregular, fruticose, to 5 cm, but usually smaller with

ascending lobes terminating in squamulose apothecia;

upper surface greyish. Lobes 9erect, convex, variable in

width, up to 2 mm, with whitish cilia along margin. Soralia

and isidia absent. Lower surface without cortex and

appearing sorediate. Apothecia abundant, terminal or

subterminal, 9lobulate on margins, shortly ciliate on

lower side appearing hairy; ascospores of Pachysporaria-

type, with sporoblastidia, 42481922 mm (too few seen

to be statistically analysed). Pycnidia not seen.

Chemistry: Kyellow. Atranorin and zeorin.

Habitat and distribution: On tree trunks and twigs. Only

known from Brazil, Ecuador and Peru in South America.

Notes: Similar to H. comosa but lacks cilia on the upper

surface, Also similar to H. podocarpa but has cilia on the

lower side of the apothecia.

Specimens examined: Brazil.Sa

˜o Paulo, Serra da Manti-

queira; Campos do Jordaoen, Hotel Toriba, 1750 m a.s.l.

1975, H. Schindler 5524 (in UPS). Ecuador. Loja, road

Celica-Alamor, km 1116, 16001700 m a.s.l. 1980, L.

Andersson 957 (in GB). Peru. Cusco, Urubamba, Machu

Picchu, outside the ruin town, 13807?S, 72836?W, 2500 m

a.s.l. 1981, A. Tehler and G. Thor P87:15 (in UPS).

Heterodermia vulgaris

(Vain.) Follman & Redo´n

(1972, p. 447)

Basionym: Anaptychia leucomela var. vulgaris Vain. (1890, p.

128). Lectotype (Kurokawa 1962, p. 81): Brazil, Minas

Gerais, Lafayette, Vainio, Lich. Bras. exs. 289 (in TUR-7855).

Literature. Kurokawa (1962, p. 81), Swinscow and Krog

(1976, p. 137).

Thallus appearing foliose, of irregular, narrow, interwoven

lobes, sometimes several cm long, with long, dark cilia

(rhizines) along the margins; upper surface whitish to

cream-coloured, 9shiny without pruina. Lobes ca 1.0(

2.5) mm wide, parallel-sided, rarely widening towards

apices, dichotomously branched with long, sparsely squar-

rose rhizines, mostly black except for the pale base. Soralia

irregularly formed on the underside causing the lobes to

widen. Lower surface without cortex, white with an

overlying patchy layer of reddish pigment turning deep

purple in K, margins corticate and prominent. Apothecia

laminal to subapical, stipitate, lobulate; ascospores of

Pachysporaria-type, with sporoblastidia, (31.0)34.5

44.0(50.5)(16.0)17.523.0(25.0) mm. M 39.2; 20.2 mm.

SD4.88; 2.78 mm. n24.

Chemistry: Kyellow. Atranorin, zeorin, leycotylin and a

reddish, Kdeep-purple pigment.

Habitat and distribution: On trees and rocks in 9shade.

Seen from Bolivia, Brazil and Peru in South America.

Probably overlooked but common and widespread in

tropical and subtropical areas. Also known from Africa

and North America.

Notes: Easily identified when the reddish patchy pigment on

the lower surface is well developed, but difficult to separate

from H. leucomela when sparse or lacking.

Acknowledgements I am grateful to the directors and curators of

herbaria mentioned in the text for the loan of material. I am also

grateful to colleagues and fellow travellers for letting me use

their material in this publication, especially Lars Arvidsson,

Gothenburg. Anders Nordin, Uppsala, has kindly revised the

Latin diagnoses and given valuable comments on the text and

146

David Galloway, Dunedin, has kindly revised the English text. I

am also grateful to the referees for valuable suggestions.

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Roccellinastrum, Cenozosia and Heterodermia: Ecology and phylogeny of fog lichens and their photobionts from the coastal Atacama Desert

Article

Full-text available

Aug 2023

Some deserts on Earth such as the Namib or the Atacama are influenced by fog which can lead to the formation of local fog oases-unique environments hosting a great diversity of specialized plants and lichens. Lichens of the genera Ramalina, Niebla or Het-erodermia have taxonomically been investigated from fog oases around the globe but not from the Atacama Desert, one of the oldest and driest deserts. Conditioned by its topography and the presence of orographic fog, the National Park Pan de Azúcar in the Atacama Desert is considered to be such a lichen hotspot. Applying multi-gen loci involving phylogenetic analyses combined with intense morphological and chemical characterization, we determined the taxonomic position of five of the most abundant epiphytic lichens of this area. We evaluated Roccellinastrum spongoideum and Hetero-dermia follmannii which were both described from the area but also finally showed that the genus Cenozosia is the endemic sister genus to Ramalina, Vermilacinia, Namibialina and Niebla. As a result, we have described the species Heterodermia adunca, C. cava and C. excorticata as new lichen species. This work provides a comprehensive dataset for common fog lichen genera of the Coastal Range of the Atacama Desert that can be used as a baseline for monitoring programs and environmental health assessments.

Biota liquénica en el monumento arqueológico de Pawkarkancha Santuario Histórico de Machupicchu

Article

Full-text available

Sep 2022

En la presente investigación se analiza la presencia de hongos liquenizados en los muros arqueológicos (recintos y andenes) del monumento de Pawkarkancha, ubicado en la confluencia de los valles Q é sqa y Pampacahuana a 3,131 msnm. de la región Cusco, con la finalidad de evaluar la riqueza liquénica que se establece en los paramentos de este monumento arqueológico. Los registros se realizaron entre los años 2017 y 2020, mediante 2 el método de conglomerados utilizando cuadrantes de 1m . Se llegó a determinar 13 familias, 24 géneros, 44 especies y morfoespecies, siendo las familias Parmeliaceae, Caliciaceae, Lecanoraceae, y Physciaceae las más abundantes. Buellia aethalea. B. stelulata, Lecanora sp. Polycauliona stellata y Verrucaria nigrescens fueron las especies de mayor cobertura (80%) destacando el biotipo crustoso en los muros arqueológicos. Se reportan nuevos registros para Perú: Lepraria tenella, Leptogium burnetiae, Parmotrema crinitum, Polycauliona stellata, Punctelia constantimontium, Pyxine petricola, Sticta albocyphellata, Xanthoparmelia inserta, Xanthoparmelia hypomelaena.

Bacterial microbiome in tropical lichens and the effect of the isolation method on culturable lichen-derived actinobacteria

Article

Full-text available

Apr 2023

Ten samples of tropical lichens collected from Doi Inthanon, Thailand, were explored for the diversity of their bacterial microbiomes through 16S rRNA-based metagenomics analysis. The five predominant lichen-associated bacteria belonged to the phyla Proteobacteria (31.84%), Planctomycetota (17.08%), Actinobacteriota (15.37%), Verrucomicrobiota (12.17%), and Acidobacteriota (7.87%). The diversity analysis metric showed that Heterodermia contained the highest bacterial species richness. Within the lichens, Ramalina conduplicans and Cladonia rappii showed a distinct bacterial community from the other lichen species. The community of lichen-associated actinobacteria was investigated as a potential source of synthesized biologically active compounds. From the total Operational Taxonomic Units (OTUs) found across the ten different lichen samples, 13.21% were identified as actinobacteria, including the rare actinobacterial genera that are not commonly found, such as Pseudonocardia, Kineosporia, Dactylosporangium, Amycolatopsis, Actinoplanes, and Streptosporangium. Evaluation of the pretreatment method (heat, air-drying, phenol, and flooding) and isolation media used for the culture-dependent actinobacterial isolation revealed that the different pretreatments combined with different isolation media were effective in obtaining several species of actinobacteria. However, metagenomics analyses revealed that there were still several strains, including rare actinobacterial species, that were not isolated. This research strongly suggests that lichens appear to be a promising source for obtaining actinobacteria.

NEW OR OTHERWISE INTERESTING RECORDS OF LICHENS AND LICHENICOLOUS FUNGI FROM MAINLAND ECUADOR AND SURROUNDINGS WITH DESCRIPTIONS OF FIVE NEW SPECIES

Article

Nov 2022

Altogether 322 taxa of lichens and lichenicolous fungi are reported from Ecuador of which many are probably new records for the country. These include five species which are described as new to science: Bacidia andina, Bacidina pulverula, Distopyrenis epidiorygma, Pertusaria lucidotetra and P. pseudoparnassia . From the separately treated genera Lepra and Pertusaria , 12 species are recorded for Ecuador and/or other Neotropical countries. Notes on morphology, chemistry and ecology are given.

New Distributional Records of Lichens for the State of Mizoram, Indo-Burma Region of India

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Feb 2022

The present paper enumerated the occurrence of 30 different species of lichens collected from 2 different study sites situated in 2 districts of Mizoram. It included Crustose (12 sp.), Foliose (16 sp.) and Fruticose (2 sp.) along with 9 species named as Diploschistes scruposus (Schreb.) Norman, Diorygma macgregorii (Vain.) Kalb, Staiger & Elix, Diorygma soozanum (Zahlbr.) M. Nakan. & Kashiw., Malmidea granifera (Ach.) Kalb, Rivas Plata & Lumbsch, Letrouitia vulpina (Tuck.) Hafellner & Bellem., Parmotrema austrosinense (Zahlbr.) Hale, Parmotrema crinitoides J.C. Wei, Parmotrema nilgherrense (Nyl.) Hale, Parmotrema rampoddense (Nyl.) Hale reported as a new record for the 1st time to the state of Mizoram, Northeast India. The finding were correlated with other previous reports on lichens and it provided detailed overview of documentation, distribution and diversity of lichens found in the small state like Mizoram under Indo-Burma region. Therefore, the richness of lichen diversity indicates the conditions of good forest health in Mizoram. The study also submits the need of conservation and exploration in the unexplored areas. HIGHLIGHTS The insight of the lichens diversity of the Mizoram state now provides the open platform for Lichenologist as well as Environmentalist The report of few species for the first time from the forest dominated hilly region of Mizoram had shown the richness of lichen distribution and its diversity As urbanisations as well as infrastructures developments are taking places in the remote corner of the state, it is very essential to explore the lichens diversities GRAPHICAL ABSTRACT

New records of lichenized fungi in Sierra Nevada de Santa Marta, Colombia

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Nov 2020

The lichen diversity of the Caribbean region of Colombia is one of the least known in the county. In the Sierra Nevada de Santa Marta (SNSM), whose forests are at risk of disappearing due to deforestation, 114 species of lichens have been recorded. We collected lichens at elevations from 500 to 2500 m on the northwestern slopes of the SNSM, and we report six genera and 19 species from this region for the first time. Our results highlight the importance of exploring new areas to enhance our knowledge of the lichens of SNSM and Colombia.

Highlights from 101 Years of Lichenological Research in Great Smoky Mountains National Park: Celebrating the United States National Park Service Centennial

Article

Nov 2019

000 new field collections generated, the park checklist now includes 920 species, a 129% increase over estimates made two decades ago. Nearly a quarter of the lichens reported in the park are known from only a single occurence whereas only 7% of the lichens are known from 20 or more occurences. An assessment of commonness/rarity for all 920 species indicates that nearly half of the park's lichens should be considered to be infrequent, rare, or exceptionally rare. We assessed the distributions of all 920 species and found that 54 are endemic to the southeastern United States, 30 are endemic to the southern Appalachians, and eight occur nowhere else than within the confines of the national park. We discuss biogeographical affinities of the park's lichen biota as a whole, delimiting six regional "floristic" connections. Our 11 years of research have resulted in the discovery of several species presumed to be extinct or near-extinct. We make one new combination (Fuscopannaria frullaniae) and describe five species as new to science, each commemorating National Park Service staff instrumental to the completion of the study: Heterodermia langdoniana, Lecanora darlingiae, Lecanora sachsiana, Leprocaulon nicholsiae, and Pertusaria superiana.

Key to Heterodermia ( Physciaceae , Teloschistales ) in Brazil, with 15 new species

Article

Full-text available

Feb 2022

The species of Heterodermia in Brazil have been revised based on literature reports, public databases and examination of 500 recent specimens from 15 states. So far, 43 species have been reported in the literature, two of which are not accepted here. We report 10 additional species from Brazil and describe 15 new to science, mostly from the Atlantic rainforest biome, raising the total number to 68, including two still undescribed species from Santa Catarina State. A key to all accepted species is given, keying out more species than represented in any previously published key of the genus. Based on new phylogenetic work, we do not accept the split genera that were recently proposed. We do informally recognize an additional group, the H. dactyliza -group. The following species are described as new: Heterodermia amphilacinulata M. F. N. Martins & M. P. Marcelli, H. caneziae M. F. Souza & Aptroot, H. delicatula M. P. Marcelli & M. F. N. Martins, H. dissecticodiademata M. F. Souza & Aptroot, H. dissecticoflabellata M. F. Souza & Aptroot, H. flavodactyliza M. F. Souza & Aptroot, H. flavulifera M. F. Souza & Aptroot, H. labiata M. P. Marcelli & M. F. N. Martins, H. macrosoraliata M. P. Marcelli & M. F. N. Martins, H. minor M. F. Souza & Aptroot, H. neocrocea M. F. Souza & Aptroot, H. nigromarginata M. F. Souza & Aptroot, H. phyllalbicans M. F. Souza & Aptroot, H. spielmannii M. F. Souza & Aptroot and H. sublinearis M. P. Marcelli & M. F. N. Martins. The following new combinations are also proposed: Heterodermia africana (Kurok.) M. F. Souza & Aptroot and H. borphyllidiata (Kalb & Meesim) M. F. Souza & Aptroot. Dissectic acid was found in species belonging to three different groups and was not restricted to (and characteristic of) one subgroup, as previously reported.

Macrolichens from Chaco National Park (Chaco Province, Argentina)

Article

Full-text available

Sep 2021

A study of the macrolichens diversity from Chaco National Park was conducted. Thirty-six species were identified, three of which are recorded for the first time from Argentina (Dirinaria melanocapa, Pyxine petricola var. convexula, and P. pungens). Moreover, the distribution of 27 species in the country is extended, including new records for Chaco province and, additionally, for other provinces of the region. An identification key for species is here provided and the geographic distribution of each one in Argentina was revised.

Leucodermia guzmaniana sp. nov. (Physciaceae, Lecanorales), a new species from Mexican cloud forest, and a key to Leucodermia in Mexico

Article

Full-text available

Jan 2019

Notes on foliose species of the lichen family Physciaceae in southern Africa 257

Article

Full-text available

Feb 2022

Notes on foliose species of the lichen family Physciaceae in southern Africa Roland Moberg Moberg, R. 2004. Notes on foliose species of the lichen family Physciaceae in southern Africa.-Acta Univ. Ups. Symb. Bot. Ups. 34:1, 257-288. ISBN 91-554-6025-9. Fifty-two species of the lichen family Physciaceae in southern Africa are treated, sixteen Heterodermia, seven Hyperphyscia, eight Phaeophyscia, fifteen Physcia and six Pyxine. Short descriptions of genera and species are given and keys are provided. One species, Heteroder-mia subcinerea, is described as new. Hyperphyscia coralloidea and Physcia jackii are reported as new to Africa. Maps of the known world distribution of Heterodermia spathulifera and Pyxine nubila are presented.

The Genus Heterodermia in the Sonoran Desert Area

Article

Full-text available

Mar 1999

Eighteen species of the lichen genus Heterodermia in the Sonoran Desert and surrounding areas are described. Morphology, anatomy, chemistry, distribution, habitat, relation to other taxa, and biogeography are discussed. A key to the species is presented. One species, H. lineare, is described as new.

Esame comparativo di alcuni generi di licheni

Article