A preliminary World-wide Key to the Lichen Genus



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A Preliminary World-wide Key to the Lichen Genus Pertusaria


A.W. Archer & J.A. Elix
The lichen genus Pertusaria (Pertusariaceae) is widely distributed throughout the world, from equatorial to polar regions (Dibben 1980; Lumbsch & Nash 2001). Species may grow on bark, rock, soil, plant débris and mosses and are differentiated by the apothecial structure (disciform or verruciform), the number and structure of the ascospores (1, 2, 4 or 8 per ascus, smooth- or rough-walled ascospores) and the chemistry (Dibben 1980; Archer 1997). Chemistry has been recognised as an important taxonomic tool in the identification of species in the genus Pertusaria (Lumbsch 1998).
The chemistry of the genus Pertusaria has been reported in many publications. Oshio (1968) reported the colour reactions of Japanese species and the compounds producing these colours were subsequently identified by Dibben (1975) who later published the chemistry of North American Pertusaria (1980). Similarly, Poelt and Vĕzda (1981) described the colour reactions of European species of Pertusaria and the identity of these compounds was later determined by Hanko (1983). Additional synonymy and chemical data for a range of European taxa was reported by Niebel-Lohmann and Feuerer (1992). Chemical data on many type specimens was reported by Archer (1993, 1995) and the chemistry of Australian Pertusaria published (Archer 1997). Additional type specimens hace since been examined and their chemistries determined. A current Key to European Pertusaria (Sipman, www.bgbm.org/BGBM/Staff/Wiss/Sipman/keys/perteuro.htm) contains much chemical information on European taxa, which has been included in this Key. Modern descriptions of new taxa in the genus would now be regarded as incomplete without a report of the chemistry.
The total number of species in the genus is not accurately known. Nash et al. (2001) suggest ca. 350 species and American Record Express (http://americanrecordexpress.com/fungi/p/) suggests ca. 920 species. Archer & Messuti (2009) calculated a possible 1550 species out of a theoretically possible 3500 chemical and morphological combinations (Archer 1997) while the Index Fungorum (www.indexfungorum.org) lists 1770 taxa, which, however, include many forms and varieties. For example, P. coccodes (Ach.) Nyl. has 17 forms and 10 varieties.
This Key includes 794 taxa, together with 482 synonyms or possible synonyms which are listed following the Key.
Please send any comments, corrections and additions to:

john.elix@anu.edu.au

alanw.archer@bigpond.com

Chemistry

The compounds found in the genus Pertusaria fall into 8 categories:
Xanthones such as lichexanthone, 4,5-dichlorolichexanthone and thiophaninic acid.
Orcinol p-depsides such as lecanoric and gyrophoric acid.
Homologues of orcinol p-depsides such as perlatolic acid and its derivatives.
Orcinol depsones such as picrolichenic acid and its homologues.
-Orcinol m-depsides such as hypothamnolic and thamnolic acids.
-Orcinol p-depsides such as squamatic and barbatic acids.
-Orcinol depsidones such as stictic, norstictic, psoromic and protocetraric acids.
Aliphatic acids such as allopertusaric acid and lichesterinic acid.

In addition, some taxa lack lichen compounds.


References:
Archer, A.W. (1997) A chemical and morphological arrangement of the lichen genus Pertusaria in Australia. Bibliotheca Lichenologica 53:1-17.
Archer, A.W. (1995) A chemical and morphological arrangement of the lichen genus Pertusaria in Australia;: additional data and corrections. Mycotaxon 55:385-389.
Archer, A.W. (1997) The lichen genus Pertusaria in Australia. Bibliotheca Lichenologica 69:5-249.
Dibben, M.J. (1975) The Chemosystematics of the lichen genus Pertusaria in North America north of Mexico. Appendix II, Exotic species, pp. 492-519, Ph.D. thesis, Duke University.
Dibben, M.J. (1980) The Chemosystematics of the lichen genus Pertusaria in North America north of Mexico. Milwaukee Publications in Biology and Geology 5: 1-162.
Hanko, B. (1983) Die Chemotypen der Flechtengattung in Europa. Bibliotheca Lichenologica 19:3-296.
Lumbsch, H.T. (1998) Taxonomic use of metabolic data in lichen-forming fungi. In: Frisvald, J.C., Bridge, P.D. & Arora, D.K (eds.) Chemical fungal taxonomy: 345-387. Marcel Dekker, New York.
Lumbsch, H.T. & Nash, T.H (2001) Pertusaria in Nash, H.T., Ryan, B.D., Gries, C & Bungartz, F. Lichen Flora of the Greater Sonoran Desert Region 1: 341-357
Messuti, M.I. & Archer, A.W. (2009) ¿Cuántos taxones pueden incluirse teóricamente en el género Pertusaria? Glalia 2(1): 1-9.
Niebel-Lohmann, A. & Feuerer, T. (1992). Die Gattung Pertusaria DC (Lichenes) in Schleswig-Holstein: Anatomie, Morphologie Taxonomie und Verbreitung 1. Mitt. Inst. Allg. Bot. Hamburg. 24: 199-252.
Oshio, M (1968) Taxonomical studies on the family Pertusariaceae of Japan. Journal of Science of the Hiroshima University, B(2), 12: 81-163.
Poelt, J. & Vezda, A. (1981). Pertusaria in Bestimmungsschlüssel europäischer Flechten. Ergänzungheft II. Bibliotheca Lichenologica 16: 1-390.

Notes:


The key uses apothecial structure and number and type of ascospores to place specimens in appropriate Groups (vide infra). Within the Groups chemical constituents, and size and arrangement of ascospores, are used to distinguish species. Synomyms and possible synonyms are indicated.

Only major chemical constituents are referred to and there remain species whose chemistry is unknown; these species are omitted until their chemistry is determined and published.

Where the results of chemical tests (K, C, etc.) are reported and give a strong indication of the substance(s) present, these species are tentatively included.
Rock includes species growing on soil and plant débris, and mosses.
Disciform species include species where the apothecia is disciform and either sessile or raised in protuberances.
Sterile sorediate or isidiate species include species that may also occur as fertile specimens; sorediate or isidiate species which are always fertile are included in the appropriate fertile Group. Taxa which are reported to occur on both wood and rock are included twice.
Species with 2-3, or 2-4 ascospores per ascus are included in the 4-spored species; species with 6, 7 or 8 ascospores are inclcuded in 8-spored species.
Group 1. Apothecia absent; isidia present; on rock

Group 2. Apothecia absent; isidia present; on wood

Group 3. Apothecia absent; soredia present; on rock

Group 4. Apothecia absent; soredia present; on wood


Group 5. Apothecia present, verruciform; ascospores 1 per ascus; on rock

Group 6. Apothecia present, verruciform; ascospores 1 per ascus; on wood

Group 7. Apothecia present, disciform; ascospores 1 per ascus; on rock

Group 8. Apothecia present, disciform; ascospores 1 per ascus; on wood


Group 9. Apothecia present, disciform; ascospores 2 per ascus; on wood

Group 10 Apothecia, present disciform; ascospores 2 per ascus; on rock


Group 11. Apothecia present, verruciform; ascospores 2 per ascus, smooth; on rock

Group 12. Apothecia present, verruciform; ascospores 2 per ascus, rough; on rock

Group 13. Apothecia present, verruciform; ascospores 2 per ascus, smooth; on wood

Group 14. Apothecia present, verruciform; ascospores 2 per ascus, rough; on wood


Group 15. Apothecia present, verruciform; ascospores 4 per ascus, smooth; on rock

Group 16. Apothecia present, verruciform; ascospores 4 per ascus, rough; on rock

Group 17. Apothecia present, verruciform; ascospores 4 per ascus, smooth; on wood

Group 18. Apothecia present, verruciform; ascospores 4 per ascus, rough; on wood


Group 19. Apothecia present, disciform; ascospores 8 per ascus, on rock

Group 20. Apothecia present, disciform; ascospores 8 per ascus, on wood

Group 21. Apothecia present, verruciform; ascospores 8 per ascus, on rock

Group 22. Apothecia present, verruciform; ascospores 8 per ascus, on wood




Sterile, lacking isidia, soredia and apothecia.


P. shenandoensis Hale & Dibben; saxicolous; norstictic acid present; N.Am

P. scepusiensis (Gyeln.) Erichs.; saxicolous; P-ve; Eur.
Group 1. Apothecia absent; isidia present; on rock or plant débris
1. Xanthones present 2

1a. Xanthones absent 4


2. UV+ yellow; lichexanthone present; SAm. P. subcorallina Nyl.

2a. UV+ orange; thiophaninic acid present 3

2b. Thiophanic acid preent; Qld P. hiatensis A.W. Archer & Elix
3. Thiophaninic acid alone; UK, Sweden P. flavocorallina Coppins & Muhr

3b. Thiophaninic acid and stictic acid present; Mediterranean region



P. rupicola (Fr.) Harm. var. coralloides (Anzi) Croz
4. Thallus K+ red or yellow 5

4a. Thallus K-ve 9


5 K+ yellow; thamnolic or baeomycesic acid present 6

5a. K+ red; norstictic or salazinic acid present 7


6. Thamnolic acid present; Eurasia; also fertile with 2 spored asci P. corallina (L.) Arn.

6a. Baeomycesic and squamatic acids present; Australia P. nerrigensis A.W. Archer & Elix


7. Norstictic acid present 8

7a. Salazinic acid present; Tasmania P. pseudodactylina A.W. Archer


8. East Asia P. corallina var. minor (Yas. ex Räs.) Oshio

8a. Tasmania P. dactylinella Kantvilas & Elix

8b. N.Am, Eur; China; fertile in Eur. P. pseudocorallina (Lilj.) Arnold

8c. Bipolar; also on wood; also 1/ascus 120-130 µm P. coccodes (Ach.) Nyl.


9(4a). Pd+ red; fumarprotocetraric or protocetraric acid present 10

9a. Pd-ve 14


10. Protocetraric acid present 11

10a. Fumarprotocetraric acid present 13


11. Protocetraric acid present 12

11a. Protocetaric and hypothamnolic acids present; SAm. P. pachythallina (Räs.) Messuti


12. Isidia to 1 mm tall; Antarctic; also fertile, 1 per ascus P. corallophora Vain.

12a. Isidia coralloid, to 20 mm tall; SAm P. acroscyphoides Sipman


13. Isidia 1-3 mm tall; N. hemisphere; also fertile, 8 per ascus P. oculata (Dickson) Th. Fr.

13a. Protocetraric acid present P. dealbata (Ach.) Cromb


14. Confluentic or 2'-O-methylperlatolic acids present 15

14a. Fatty acids present 16


15. Confluentic acid present; Australia; Hafellner 15754a Pertusaria sp.

15a. 2'-O-Methylperlatolic acid present; also fertile, 2 /ascus; Antarctic P. signyae Øvstedal


16. Isidia densely branched in upper parts; Antarctica P. peudoculata Øvstedal

16a. Isidia to 1 mm tall; bipolar P. isidiodes (Schaer.) Arnold


Group 2. Apothecia absent; isidia present; on wood

1. Xanthones present 2

1a. Xanthones absent 13
2. Lichexanthone or derivatives present 3

2a. Norlichexanthone derivatives or thiophaninic acid present 12


3. Lichexanthone and salazinc acid present; PNG P. parmatica A.W. Archer & Elix

3a. Lichexanthone and stictic acid; Thailand; P. flavodigitata Jariang.

3b. 4,5-Dichlorolichexanthone or 2-chlorolichexanthone present 4

3c. Lichexanthone, 2-chlorolichexanthone and superlatolic acid present; Qld. P. glebulosa A.W. Archer & Elix
4. 4,5-Dichloro- & 2,4,5-trichlorolichexanthone present; Australia

P. palumensis Elix & A.W. Archer

4a. 2-Chlorolichexanthone, stictic and 2'-O-methyl- & 2-O-methylperlatolic acids present;

Australia

P. bagoensis Elix & A.W. Archer

4b. 4,5-Dichlorolichexanthone and depsides or depsidones present 5



4c. 4,5-Dichlorolichexanthone and chiodectonic acid present; isidia red inside; Thailand

[Jariangprasert 4933.1] “P. rufodigitata”
5. 4,5-Dichlorolichexanthone and stictic or alectoronic acid present 6

5a. 4,5-Dichlorolichexanthone and depsides present 10


6. 4,5-Dichlorolichexanthone and stictic acid present 7

6a. 4,5-Dichlorolichexanthone and alectoronic acid present 11


7. 4,5-Dichlorolichexanthone and stictic acid present; 8

7a. 4,5-Dichlorolichexanthone, stictic acid and depsides present 9


8. Sterile; isidia 0.4-1 mm tall, 0.25-0.5 mm diam; Australia, PNG

P. montpittensis A.W.Archer

8a. Also fertile [Group 17]; isidia 0.1-0.5 mm tall; to 0.1 mm diam.; Eur



P. coronata (Ach.) Th.Fr.
9a. 4,5-Dichlorolichexanthone, stictic and 2'-O-methylperlatolic acid acids present; Australia

P. pilosula A.W. Archer & Elix

9b. 4,5-Dichlorolichexanthone, stictic acid and methyl barbatate present; PNG P. angabangensis A.W. Archer & Elix


10. 2-O-Methylperlatolic acid present; Australia P. georgeana var. georgeana A.W. Archer & Elix

10a. 2-O-Methylperlatolic and 2-O-methylstenosporic acids present; Australia P. georgeana var. methylstenosporica A.W. Archer & Elix

10b. 2,4-Di-O-methylolivetoric acid present; Australia P. georgeana var. goonooensis A.W. Archer & Elix

10c. Planaic acid present; Australia P. georgeana var. victoriana A.W. Archer & Elix

10d. 2-O-Methylconfluentic acid present; Australia P. georgeana var. occidentalis Elix & A.W. Archer
11. Alectoronic acid present; Australia P. alectoronica Elix & A.W. Archer

11a. Alectoronic and thiophanic acids present; Australia P. alectoronica var. thiophanica Kantvilas, Elix & A.W. Archer


12(2a).Thiophaninic and stictic acids present; Australia & PNG P. flavoisidiata A.W. Archer & Elix

Rarely fertile, spores 8/ascus; Europe P. flavida (DC) J.R. Laundon

12a. Arthothelin and 6-O-methylarthothelin present; New Caledonia P. simoneana A.W. Archer & Elix

12b. Thiophanic acid present; Qld. P. lumbschii A.W. Archer & Elix


13(1a).Depsidones present 14

13a. Depsides or depside esters present 21

13b. Lichesterinic acid present; Australia P. ambigua A.W. Archer & Elix

13c. Lichen compounds absent; West Indies P. griseola Vain.


14. Stictic acid or derivatives present 15

14a. Hypoprotocetraric, neotricone, norstictic, protocetraric, psoromic or salazinic acids present

17
15. Stictic acid present; on wood; isidia 0.2-0.8 mm tall, 0.05-0.1 mm diam., Mauritius, Australia

P. muricata J.C. David

NZ; on plant débris; isidia 0.5-1.5 mm tall, 0.15-0.25 mm diam.



P. dennistonensis Elix & A.W. Archer

15a. Stictic and 2'-O-methylperlatolic acids present; Thailand, PNG P. pilosula var. abditiva Jariangprasert

15b. Hypostictic and stictic acids present; Thailand P. hypostictica Jariangprasert

15c. Norstictic acid present 16


16. Isidia to 3 mm tall; Hawaii P. ramulifera H. Magn

16a. Isidia 0.1-0.25 mm tall; Australia, PNG P. roseola A.W. Archer & Elix

16b. Isidia 0.5-1.0 mm tall; bipolar; also on rock P. coccodes (Ach.) Nyl.
17(14a). Protocetraric or hypoprotocetraric acids present 18

17a. Neotricone, norstictic, psoromic or salazinic acids present 20


18. Hypoprotocetraric acid present; PNG P. hypoprotocetrarica A.W. Archer & Elix

18a. Protocetraric acid present 19


19. Protocetraric acid present; Australia P. umbricola A.W. Archer & Elix

19a. Protocetraric and norstictic acids present; Australia P. wallamanensis Elix & A.W.Archer


20(17a)Psoromic acid present; PNG P. wauensis Elix & A.W. Archer

20a. Neotricone present; Australia P. neotriconica Elix & A.W. Archer

20b. Norstictic and salazinic acids present; PNG; HS 19405 Pertusaria sp.
21(13a).Methyl 3--hydroxy-4-O-demethylbarbatate; PNG P. mankiensis A.W. Archer & Elix

21a. Other depsides present 22


22. Thamnolic acid present; Eur; also fertile; 2 spores/ascus; isidia short and thick,

0.1-0.3 mm diam P. corallina (L.) Arn.

Isidia very thin, 0.03-0.05 mm diam.; Australia Pertusaria tricosa Elix & A.W. Archer

22a. 2'-O-methylperlatolic, barbatic or divaricatic acids or derivatives present 23


23. 2'-O-Methylperlatolic acid present; PNG, Australia P. burburana Elix & A.W. Archer

23a. Barbatic or divaricatic acid present 24


24. Barbatic acid present; Australia P. barbatica A.W. Archer & Elix

24a. Divaricatic acid present; PNG P. labuensis A.W. Archer & Elix

Group 3. Apothecia absent; soredia present; on rock or plant débris.
1. Xanthones present 2

1a. Xanthones absent 15


2. Lichexanthone or chlorolichexanthones present 3

2a. Thiophaninic acid or thiophanic acids present 11


3. Chlorinated lichexanthones present 4

3a. Lichexanthone present 5


4. 2,4- and 2,5-Dichlorolichexanthone, 2,4,5-trichlorolichexanthone and stictic acid present

Australia P. puffina A.W. Archer & Elix

4a. 4,5-Dichlorolichexanthone and stictic acid present; PNG; also corticolous

P. balekensis A.W. Archer & Elix
5. Lichexanthone and p-depsides or -orcinol m-depsides or stictic acid present 6

5a. Lichexanthone and picrolichenic or stictic acid acid present 8


6. Lichexanthone and methyl 2'-O-methylmicrophyllinate present; Australia

P. variabilis Elix & A.W. Archer

6a. Lichexanthone and hypothamnolic, haemathamnolic or stictic acid present 7


7. Hypothamnolic acid present P. amaroides H. Magn.

Hypothamnolic and barbatic acids present; Uruguay; 7031 Pertusaria sp.

Stictic acid present; Aus; JAE 43109 Pertusaria sp.

7a. Haemathamnolic acid present; Brazil P. xantholeucoides Müll. Arg.


8. Lichexanthone and picrolichenic acid alone ± superpicrolichenic acid present 9

8a. Lichexanthone, picrolichenic acid and -orcinol m-depsides 10


9. Lichexanthone and picrolichenic acid present; Australia & S.Am.

[rarely with additional norstictic acid] P. subventosa var. deficiens A.W. Archer & Elix

9b. Lichexanthone, picrolichenic and superpicrolichenic acids present; Norfolk Island, Australia

P. verdonii A.W. Archer
10. Lichexanthone, picrolichenic and thamnolic acid present; S. Am, Australia and NZ;

very rarely fertile P. subventosa var. subventosa Malme

10a. Lichexanthone, picrolichenic and hypothamnolic acid present; Australia & PNG

P. subventosa var. hypothamnolica A.W. Archer & Elix

Lichexanthone, picrolichenic and barbatic acid present;S.Am; Ahti 31.i.1989(H) Pertusaria sp.


11(2a).Thiophaninic acid, confluentic and planaic acids present; N.Am; ± fertile P. tejocotensis de Lesd.

11a. Thiophaninic acid and norstictic acid (major) and stictic acid (minor) present; Aus



Pertusaria rogersii A.W. Archer & Elix

11b. Thiophanic acid and arthothelin present; on plant deebris; Qld P. "finniganensis"

11c. Thiophaninic acid and stictic acid present 12


12. In Australia 13

12a. Elsewhere 14


13. Thallus bright yellow; hypostictic acid absent; E Australia, NZ & PNG; ± fertile

P. xanthoplaca Müll. Arg.

13a. Thallus dull yellow; hypostictic acid present; NW Australia; P. remota A.W. Archer


14. In Europe, Asia, Africa P. amarescens Nyl

In Japan P. leucosoroides Nyl.

14a. In N.Am; ± fertile P. flavicunda Nyl.
15(1a) Picrolichenic acid or p-depside present 16

15a. Depsidones present 21

15b. Lichen compounds absent; Canada P. pruinifera Erichs.

Philippines; P. subvaginata var. orientalis Räs.


16. Picrolichenic acid present P. amara (Ach.) Nyl.

Additional protocetraric acid present; Eur. P. amara var. flotowiana (Flörke) Erichs.

16a. Depsides present 17
17. Haemathamnolic acid present; Central Africa P. kinigiensis A.W. Archer et al.

17a. p-Depsides present 18


18. Lecanoric acid present 19

18a. Other p-depsides present 20


19. Lecanoric acid present, variolaric acid absent; Philippines

P. subvaginata var. orientalis Räs.

19a. Lecanoric and variolaric acids present; Europe; ± fertile P. lactea (L.) Arnold


20. Squamatic and baeomycesic acids present; Australia P. nerrigensis A.W. Archer & Elix

20a. 2'-O-Methylisohyperlatolic and 2'-O-methylsuperlatolic acids present; Australia



P. salebrosa A.W. Archer & Elix
21(15a).Norstictic or stictic acid present 22

21a. Protocetraric or fumarprotocetraric acid present 24


22. Stictic acid present; soralia ochre coloured; S. Georgia Pertusaria sp. A Øvstedal

22a. Norstictic acid present 23


23. Thallus off-white; Europe, China, N.Am, P. excludens Nyl.

23a. Thallus olive green; Australia, Lord Howe Is P. miniatescens A.W. Archer & Elix

23b. Norstictic and picrolichenic acids present; Teneriffe P. teneriffensis Vain.
24. Protocetraric acid present; Europe, N.Afr., Japan P. leucosora Nyl

Canary Islands P. inconveniens Vain.

24a. Fumarprotocetraric acid present 25
25a. Atranorin present; Australia; soralia off-white; subtropical P. sordida A.W. Archer

25a. Atranorin absent; ± succinprotocetraric acid present; Eur 26


26. Thallus with flattened white soralia P. aspergilla (Ach.) J.R. Laundon

26a. Thallus with sorediate papillae P. mammosa Harm.

26b. Thallus on limestone P. gypsicola Erichs.

Group 4. Apothecia absent; soredia present; on wood


1. Xanthones present 2

1a. Xanthones absent 9


2. Lichexanthone or 4,5-dichlorolichexanthone present 3

2a. Arthothelin, thiophaninic acid or thuringione present 5



2b. Arthothelin, thiophaninic acid and barbatic acid present;Thailand “P. flavosorediata”

[Jariangprasert 5512.2]
3. 4,5-Dichlorolichexanthone present 4

3a Lichexanthone present with depsone, depsidone, depside or fatty acid 7


4. 4,5-Dichlorolichexanthone present alone; PNG; also saxicolous

P. balekensis A.W. Archer & Elix

4a. 4,5-Dichlorolichexanthone and stictic acid present; N.Am P. expolita Harris


5. Thiophaninic acid and depsidones present 6

5a. Arthothelin and thuringone present; Australia P. flavopunctata A.W. Archer & Elix


6. Thiophaninic acid and stictic acid present; Australia P. maritima A.W. Archer & Elix

6a. Thiophaninic acid and norstictic acid present; N.Am P. azulensis de Lesd.


7. Lichexanthone, picrolichenic and superpicrolichenic acids present; Norfolk Island, Australia P. verdonii A.W. Archer

7a. Lichexanthone and confluentic, thamnolic, or stictic acid present 8



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