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Studies in the lichen family Pannariaceae X. The lichen genus Protopannaria in the subantarctic islands.
Cryptogamie, Mycol., 2001, 22 (1): 67-72
67
© 2001 Adac/l~litions scientifiques et m6dicales Elsevier SAS. Tous droits r6serv6s S0181158401010594/FLA
Studies in the fichen family Pannariaceae X. The lichen genus Protopannaria in the subantarctic islands. Per Magnus JORGENSEN* Dept. of Botany, University of Bergen, All~gt.41, N-5007 Bergen, Norway
Abstract - - The lichen genus Protopannaria is defined as different from Pannaria, being a bipolar taxon with three (of four) species in the subantarctic region: P. austro-orcadensis(13vst.) comb. nov. in the western part and P. alcicornis P.M. JCrg. & R.S. Poulsen sp. nov. as well as P. azorellae P.M. J~rg. & R.S. Poulsen sp. nov. in the eastern one (Kerguelen), showing the greatest diversity there. © 2001 Adac / l~ditions scientifiques et m6dicales Elsevier SAS lichen taxonomy / Pannariaceae / phytogeography / subantarctic region
R ~ s u m 6 - Le genre Protopannaria est accept6 et defini comme distinct de Pannaria. C'est un genre bipolaire dont trois (des quatre) esp~ces sont de la r6gion subantarctique: P. austro-orcadensis (13vst.) comb. nov. dans les zones occidentales; P. alcicornis P.M. JCrg. & R.S. Poulsen, sp. nov. et P. azoreUaeP.M. JCrg. & R.S. Poulsen, sp. nov., dans les zones orientales (Kerguelen), oh se trouve la plus grande diversit6. © 2001 Adac / Editions scientifiques et m6dicales Elsevier SAS
taxonomy des lichens / Pannariaceae / phytog~ography / r~gion sub-antarctique
INTRODUCTION W h e n JCrgensen (1994) remodelled the genus Pannaria, he discussed the problematic position o f Pannaria pezizoides (G.H. Weber) Trevis., and confirmed a placement in subg. Protopannaria Gyeln., but also repeated an earlier idea (JCrgensen, 1978) that it m a y belong to Psoroma. Molecular studies (Ekman & J~rgensen, 2000) clearly put this species outside both genera, and as a consequence this subgenus was accepted as a genus (JCrgensen, 2000a), Protopannaria (Gyeln.) P.M. JCrg. & Ekm. The genera can be distinguished as shown in Tab. 1.
* Correspondence and reprints.
68
M. JCrgensen
Tab. 1. Comparison of characters between PannarialProtopannaria.
Thallus Chemistry Apothecia Hymenium Asci Distribution
Pannaria
Protopannaria
Foliose to squamulose Pannarin and related subst. Squamulose margin I + clear blue in vicinityof asci Without amyloid apical structure Pantropical to warm temperate
Squamulose to crustose No substances Granular margin Totally I + blackish blue Without arnyloid apical structure Bipolar, arctic/alpine
When revising material from the subantarctic islands additional species where found. They are described below.
T H E SUBANTARCTIC SPECIES These species can be distinguished in the following way: la Thallus mostly crustose-granular, richly fertile, in dying Azorella-cushions ............................................................................................. P. azorellae lb Thallus small-squamulose,sparingly fertile, on mosses or soil ................................. 2 2a Squamules coralloid, dark, matt brown, western .................. P. austro-orcadensis 2b Squamules like small antlers of moose, shining brown, eastern ...... P alcicornis 1. Protopannaria austro-orcadensis (Ovst.) P.M. Jc~rg., comb. nov. Pannaria austro-orcadensis Ovst., Crypt., Bryol. Lich6nol.14(4):337 (1993). Type: SOUTH ORKNEY ISLANDS, Signy Isl., Moraine Valley, Jan.1967, R.I.L. Smith 985 (AAS, holotypel). (Fig.l) Thallus squamulose to coralloid, dark brown, forming circular patches up to 5 cm on bryophytes. No lichen acids found. Apothecia rare, to 2.5 mm diam, with dark brown disc and nodular thaUine margin. Hymenium I + blackish blue with asci without amyloid apical apparatus. Spores simple, colourless, broadly ovate 16--18 x 12-13 larn Notes: As pointed out by Ovstedal & Smith (1993: 339) this species is closely related to P. pezizoides with the same ascus structure, so it must be referred to Protopannaria. It differs, however, clearly from P. pezizoides in the darker brown, coralloid thallus which has a lower cortex, and the smaller, smooth spores. It is closer related to P. alcicornis which has a different thallus with more elongated, branched squamules, which looks like small moose antlers, with a different anatomy (thicker upper cortex, no lower one) and different (narrower, rugulose) spores. Habitat and distribution: This is a species of moist moss-tufts on weakly alkaline, mineral soil in solifluction streams (Ovstedal & Lewis Smith, 1993: 328-329). It is often associated with Pannaria caespitosa P.M. J¢rg. and occasionally with Placopsis contortuplicata Lamb. P austro-orcadensis is as yet oialy known from the South Orkney and South Shetland islands, and may prove to be endemic to this region of subantarctic islands. Other material examined: SOUTH SHETLAND, Signy Isl., Robin Peak Plateau, alt.180 m, 20.1.85, R.I. Lewis Smith (AAS).
The lichen genus Protopannaria
69
Fig. 1. a. Protopannaria austro-orcadensis from Signy Isl., showing the coralloid thallus; b. Protopannaria alcicornis, detail of type, showing the characteristic, antler-like squamules.
70
M. Jergensen
Fig. 2. Pannaria azorellae, detail of type specimen, showing the numerous, small apothecia.
2. Protopannaria alcicornis EM. Jerg. & R.S. Poulsen, sp. nov. Protopannariae austro-orcadensis similis sed thallo squamis alcicornis et apotheciis magnis. Typus: KERGUELEN, Peninsule Courbet, just below the summit of Mt.Crozier, alt. 950 m, 18.12 1998 R.S. Poulsen 301 (C, holotype) Thallus squamulose, shining brown, effuse to 5 cm; individual squamules elongated, incised, often with isidioid branchlets, rather like antlers of moose (Alces alces); upper cortex thick to 50 grn, of irregular structure, no lower cortex. No chemical reactions, and without lichen substances. Apothecia irregularly expanding, often with sinuose thalline margin, to 2 mm broad. Hymenium to 100 ~rn, I + dark blue; asci without apical amyloid structures; spores, coloudess, pyriform with apical thickening of perispore, 15-17 x 9 larn. Notes: P. alcicornis is a most characteristic species by the very unusual squamules which in parts appear coralloid, like P. austro-orcadensis which has a darker brown, dull thallus of different anatomy, and smaller, more regular apothecia. Habitat and distribution: This species appears to be confined to naked soil or moss tufts, occasionally onto basaltic rocks in moist places, such as snow beds, at high altitudes in Kerguelen, and gives the impression of being a local endemic. Additional specimens examined: KERGUELEN, La Mortadelle, SE of the summit of La Mortadelle, upper S-exposed side of mountain, sloping scree, on damp gravelly soil, alt. 650 m, R.S. Poulsen 712 (C). 3. Protopannaria azorellae P.M. Jerg. & R. S. Poulsen, sp. nov. Protopannariae pezizoidis similis, sed thaUo atrissimo, granuloso, fertilissimo et apotheciis et sporis minutis.
The lichen genus Protopannaria
71
Fig. 3. Distribution of the genus Protopannaria in the Antarctic region.
Typus: KERGUELEN, Ile Foch, SE side of Baie Phillips, 2.5 km from W end of bay, 0.5 km from the coast, alt. 30 m, R.S. Poulsen 976 (C, holotype). Thallus crustose of granulose, dark brown squamules growing effusely over the substrate, to several centimetres diam. No chemical reactions or substances found. Apothecia plentiful, up to 1 m m diam., with distinct granular thalline margin. Hymenium up to 100 pm, I + dark blue; asci without amyloid apical structures; ascospores simple, colourless, ellipsoid, rugulose, 12-14 x 5-7 jam. Notes: P. azorellae is superficially rather like P. pezizoides, but has a darker, more granular thallus covered in numerous small apothecia with distinctly smaller spores which at most are mgulose, not regularly warted. Habitat and distribution: As yet only known from Kerguelen and there in a very specialized habitat, inside dying tufts of Azorella selago Hook. ill. in exposed coastal habitats, finding protection in these cushions. It remains to be seen whether it is confined to this habitat, and follows the circumantarctic distribution of its host. One collection, however, is on stems in carpets of Acaena magellanica, indicating a less species specific dependance, but one to hosts protecting it from great climatic changes.
72
M. JOrgensen
Other specimens examined: KERGUELEN, Ile Foch, Phillips, ca 2 km from W end of Bay, on dying Azorella Acaena magellanica, 19.2.1999 R.S. Poulsen 968 (C); He Mayes, corner, 100 m from the sea, on moribund Azorella selago, alt. Pouisen 202 (C).
along SE side of Baie selago and stems of Golfe du Morbihan, SE 10 m, 17.12. 1998 R.S.
CONCLUSIVE REMARKS The discovery of three further species in the genus Protopannaria in the subantarctic region is of considerable taxonomic and phytogeographic interest. It proves that this genus is not monospecific and that it most possibly originated in the cold parts of the Southern Hemisphere, far away from the tropical region in which the genus Pannaria s. str. has its main distribution (JCrgensen, 1994). Since P. pezizoides still has not been discovered in the Antarctic region, its origin remains obscure. It is, however, most closely related to the other species, particularly P. azorellae from Kerguelen which is an ancient island where the genus has its greatest species-diversity (50 %, Fig.3), just as in Pannaria subgen. Chryopannaria P.M. J~rg.(J¢rgensen, 2000b). The presence of P. pezizoides in the East African mountains (J~rgensen, 1978), suggests a possible route of migration to the northern Hemisphere.
REFERENCES
EKMAN S. & J43RGENSEN EM., in ed. - - Molecular studies in the lichen family Pannariaceae, JORGENSEN P.M., 1978 - - The lichen family Pannariaceae in Europe. Opera botanica 45, 123p. J43RGENSEN EM., 1994 - - Studies in the lichen family Parmariaceae VI: The taxonomy and phytogeography of Pannaria Del. s.lat. Journal of the Hattori Botanical Laboratory 76: 197-206. JORGENSEN EM., 2000a - - Survey of the lichen family Pannariaceae in the American Continent, North of Mexico. Bryologist 103:670-704 JORGENSEN P.M., 2000b - Studies in the lichen family Pannariaceae IX. A revision of Pannaria subg. Chryopannaria. Nova Hedwigia 69:405-414 OVSTEDAL D.O. & LEWIS SMITH R.I., 1993 - - A new Pannaria species from the Antarctic. Cryptogamie, Bryologie et Lichenologie 14(4): 337-340.
67
© 2001 Adac/l~litions scientifiques et m6dicales Elsevier SAS. Tous droits r6serv6s S0181158401010594/FLA
Studies in the fichen family Pannariaceae X. The lichen genus Protopannaria in the subantarctic islands. Per Magnus JORGENSEN* Dept. of Botany, University of Bergen, All~gt.41, N-5007 Bergen, Norway
Abstract - - The lichen genus Protopannaria is defined as different from Pannaria, being a bipolar taxon with three (of four) species in the subantarctic region: P. austro-orcadensis(13vst.) comb. nov. in the western part and P. alcicornis P.M. JCrg. & R.S. Poulsen sp. nov. as well as P. azorellae P.M. J~rg. & R.S. Poulsen sp. nov. in the eastern one (Kerguelen), showing the greatest diversity there. © 2001 Adac / l~ditions scientifiques et m6dicales Elsevier SAS lichen taxonomy / Pannariaceae / phytogeography / subantarctic region
R ~ s u m 6 - Le genre Protopannaria est accept6 et defini comme distinct de Pannaria. C'est un genre bipolaire dont trois (des quatre) esp~ces sont de la r6gion subantarctique: P. austro-orcadensis (13vst.) comb. nov. dans les zones occidentales; P. alcicornis P.M. JCrg. & R.S. Poulsen, sp. nov. et P. azoreUaeP.M. JCrg. & R.S. Poulsen, sp. nov., dans les zones orientales (Kerguelen), oh se trouve la plus grande diversit6. © 2001 Adac / Editions scientifiques et m6dicales Elsevier SAS
taxonomy des lichens / Pannariaceae / phytog~ography / r~gion sub-antarctique
INTRODUCTION W h e n JCrgensen (1994) remodelled the genus Pannaria, he discussed the problematic position o f Pannaria pezizoides (G.H. Weber) Trevis., and confirmed a placement in subg. Protopannaria Gyeln., but also repeated an earlier idea (JCrgensen, 1978) that it m a y belong to Psoroma. Molecular studies (Ekman & J~rgensen, 2000) clearly put this species outside both genera, and as a consequence this subgenus was accepted as a genus (JCrgensen, 2000a), Protopannaria (Gyeln.) P.M. JCrg. & Ekm. The genera can be distinguished as shown in Tab. 1.
* Correspondence and reprints.
68
M. JCrgensen
Tab. 1. Comparison of characters between PannarialProtopannaria.
Thallus Chemistry Apothecia Hymenium Asci Distribution
Pannaria
Protopannaria
Foliose to squamulose Pannarin and related subst. Squamulose margin I + clear blue in vicinityof asci Without amyloid apical structure Pantropical to warm temperate
Squamulose to crustose No substances Granular margin Totally I + blackish blue Without arnyloid apical structure Bipolar, arctic/alpine
When revising material from the subantarctic islands additional species where found. They are described below.
T H E SUBANTARCTIC SPECIES These species can be distinguished in the following way: la Thallus mostly crustose-granular, richly fertile, in dying Azorella-cushions ............................................................................................. P. azorellae lb Thallus small-squamulose,sparingly fertile, on mosses or soil ................................. 2 2a Squamules coralloid, dark, matt brown, western .................. P. austro-orcadensis 2b Squamules like small antlers of moose, shining brown, eastern ...... P alcicornis 1. Protopannaria austro-orcadensis (Ovst.) P.M. Jc~rg., comb. nov. Pannaria austro-orcadensis Ovst., Crypt., Bryol. Lich6nol.14(4):337 (1993). Type: SOUTH ORKNEY ISLANDS, Signy Isl., Moraine Valley, Jan.1967, R.I.L. Smith 985 (AAS, holotypel). (Fig.l) Thallus squamulose to coralloid, dark brown, forming circular patches up to 5 cm on bryophytes. No lichen acids found. Apothecia rare, to 2.5 mm diam, with dark brown disc and nodular thaUine margin. Hymenium I + blackish blue with asci without amyloid apical apparatus. Spores simple, colourless, broadly ovate 16--18 x 12-13 larn Notes: As pointed out by Ovstedal & Smith (1993: 339) this species is closely related to P. pezizoides with the same ascus structure, so it must be referred to Protopannaria. It differs, however, clearly from P. pezizoides in the darker brown, coralloid thallus which has a lower cortex, and the smaller, smooth spores. It is closer related to P. alcicornis which has a different thallus with more elongated, branched squamules, which looks like small moose antlers, with a different anatomy (thicker upper cortex, no lower one) and different (narrower, rugulose) spores. Habitat and distribution: This is a species of moist moss-tufts on weakly alkaline, mineral soil in solifluction streams (Ovstedal & Lewis Smith, 1993: 328-329). It is often associated with Pannaria caespitosa P.M. J¢rg. and occasionally with Placopsis contortuplicata Lamb. P austro-orcadensis is as yet oialy known from the South Orkney and South Shetland islands, and may prove to be endemic to this region of subantarctic islands. Other material examined: SOUTH SHETLAND, Signy Isl., Robin Peak Plateau, alt.180 m, 20.1.85, R.I. Lewis Smith (AAS).
The lichen genus Protopannaria
69
Fig. 1. a. Protopannaria austro-orcadensis from Signy Isl., showing the coralloid thallus; b. Protopannaria alcicornis, detail of type, showing the characteristic, antler-like squamules.
70
M. Jergensen
Fig. 2. Pannaria azorellae, detail of type specimen, showing the numerous, small apothecia.
2. Protopannaria alcicornis EM. Jerg. & R.S. Poulsen, sp. nov. Protopannariae austro-orcadensis similis sed thallo squamis alcicornis et apotheciis magnis. Typus: KERGUELEN, Peninsule Courbet, just below the summit of Mt.Crozier, alt. 950 m, 18.12 1998 R.S. Poulsen 301 (C, holotype) Thallus squamulose, shining brown, effuse to 5 cm; individual squamules elongated, incised, often with isidioid branchlets, rather like antlers of moose (Alces alces); upper cortex thick to 50 grn, of irregular structure, no lower cortex. No chemical reactions, and without lichen substances. Apothecia irregularly expanding, often with sinuose thalline margin, to 2 mm broad. Hymenium to 100 ~rn, I + dark blue; asci without apical amyloid structures; spores, coloudess, pyriform with apical thickening of perispore, 15-17 x 9 larn. Notes: P. alcicornis is a most characteristic species by the very unusual squamules which in parts appear coralloid, like P. austro-orcadensis which has a darker brown, dull thallus of different anatomy, and smaller, more regular apothecia. Habitat and distribution: This species appears to be confined to naked soil or moss tufts, occasionally onto basaltic rocks in moist places, such as snow beds, at high altitudes in Kerguelen, and gives the impression of being a local endemic. Additional specimens examined: KERGUELEN, La Mortadelle, SE of the summit of La Mortadelle, upper S-exposed side of mountain, sloping scree, on damp gravelly soil, alt. 650 m, R.S. Poulsen 712 (C). 3. Protopannaria azorellae P.M. Jerg. & R. S. Poulsen, sp. nov. Protopannariae pezizoidis similis, sed thaUo atrissimo, granuloso, fertilissimo et apotheciis et sporis minutis.
The lichen genus Protopannaria
71
Fig. 3. Distribution of the genus Protopannaria in the Antarctic region.
Typus: KERGUELEN, Ile Foch, SE side of Baie Phillips, 2.5 km from W end of bay, 0.5 km from the coast, alt. 30 m, R.S. Poulsen 976 (C, holotype). Thallus crustose of granulose, dark brown squamules growing effusely over the substrate, to several centimetres diam. No chemical reactions or substances found. Apothecia plentiful, up to 1 m m diam., with distinct granular thalline margin. Hymenium up to 100 pm, I + dark blue; asci without amyloid apical structures; ascospores simple, colourless, ellipsoid, rugulose, 12-14 x 5-7 jam. Notes: P. azorellae is superficially rather like P. pezizoides, but has a darker, more granular thallus covered in numerous small apothecia with distinctly smaller spores which at most are mgulose, not regularly warted. Habitat and distribution: As yet only known from Kerguelen and there in a very specialized habitat, inside dying tufts of Azorella selago Hook. ill. in exposed coastal habitats, finding protection in these cushions. It remains to be seen whether it is confined to this habitat, and follows the circumantarctic distribution of its host. One collection, however, is on stems in carpets of Acaena magellanica, indicating a less species specific dependance, but one to hosts protecting it from great climatic changes.
72
M. JOrgensen
Other specimens examined: KERGUELEN, Ile Foch, Phillips, ca 2 km from W end of Bay, on dying Azorella Acaena magellanica, 19.2.1999 R.S. Poulsen 968 (C); He Mayes, corner, 100 m from the sea, on moribund Azorella selago, alt. Pouisen 202 (C).
along SE side of Baie selago and stems of Golfe du Morbihan, SE 10 m, 17.12. 1998 R.S.
CONCLUSIVE REMARKS The discovery of three further species in the genus Protopannaria in the subantarctic region is of considerable taxonomic and phytogeographic interest. It proves that this genus is not monospecific and that it most possibly originated in the cold parts of the Southern Hemisphere, far away from the tropical region in which the genus Pannaria s. str. has its main distribution (JCrgensen, 1994). Since P. pezizoides still has not been discovered in the Antarctic region, its origin remains obscure. It is, however, most closely related to the other species, particularly P. azorellae from Kerguelen which is an ancient island where the genus has its greatest species-diversity (50 %, Fig.3), just as in Pannaria subgen. Chryopannaria P.M. J~rg.(J¢rgensen, 2000b). The presence of P. pezizoides in the East African mountains (J~rgensen, 1978), suggests a possible route of migration to the northern Hemisphere.
REFERENCES
EKMAN S. & J43RGENSEN EM., in ed. - - Molecular studies in the lichen family Pannariaceae, JORGENSEN P.M., 1978 - - The lichen family Pannariaceae in Europe. Opera botanica 45, 123p. J43RGENSEN EM., 1994 - - Studies in the lichen family Parmariaceae VI: The taxonomy and phytogeography of Pannaria Del. s.lat. Journal of the Hattori Botanical Laboratory 76: 197-206. JORGENSEN EM., 2000a - - Survey of the lichen family Pannariaceae in the American Continent, North of Mexico. Bryologist 103:670-704 JORGENSEN P.M., 2000b - Studies in the lichen family Pannariaceae IX. A revision of Pannaria subg. Chryopannaria. Nova Hedwigia 69:405-414 OVSTEDAL D.O. & LEWIS SMITH R.I., 1993 - - A new Pannaria species from the Antarctic. Cryptogamie, Bryologie et Lichenologie 14(4): 337-340.