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Thyrea porphyrella, a new species of the Lichinaceae from inselbergs in tropical South America
Lichenologist 33(3): 211-214 (2001) doi:10.1006/lich.2000.0317, available online at http://www.idealibrary.com on IDEJ^L
Thyrea porphyrella, a new species of the Lichtnaceae from inselbergs in tropical South America M. SCHULTZ, B. BUDEL and S. POREMBSKI Abstract: The new species Thyrea porphyrella Schultz, Budel & Porembski, in the Lichinaceae, is described from tropical South America. It grows on granite inselbergs in SE Brazil and in southern Guyana. Diagnostic characters are the very thin, superficially smooth but conspicuously uneven to folded thallus lobes of reddish-brown to crimson colour. £ 2001 The British Lichen Society
gelatinous, ecorticate, heteromerous thalli with a loose or compact central strand. Their type of ascoma development is unique to approximately half of the Lichinaceae. Ascogonia are formed beneath pycnidia, and the pycnidia are then transformed into apothecia (so-called pycnoascocarps, cf. Henssen 1963). Table 1 gives an overview of the species currently treated under Thyrea s. str. and their distribution. The species Thyrea of the Lichinaceae such as Gonohymenia, arenae A. Massal. ex Forssell, T. borzii Lempholemma, Lichinella, Paulia and Beltr. and T. camaromorpha (A. Massal.) Peccania (Henssen 1968, 1979, 1986;A. Massal. are dubious. Thyrea latissima Henssen & Jorgensen 1990; Moreno & Asahina was transferred to Anema by Egea 1992a). The genus Digitothyrea was Yoshimura (1968) due to the similar shape described by Moreno & Egea (19926) of the hyphae. However, in our opinion, the for those species of Thyrea s. lat. having a original placement by Asahina (1958) based ± fruticose growth form with elongated, on the heteromerous thallus structure better tongue-shaped lobes and a type of ascoma reflects the generic limits (at least for the ontogeny starting with ascogonia formed in moment) and hence should be maintained. a tangle of generative hyphae. Currently, In this paper we describe a new species of Thyrea s. str. contains approximately 12 Thyrea from two granite inselbergs in species that are quite large and foliose or tropical South America. These are the first small squamulose lichens predominantly records of Thyrea s.str. from the Southern colonizing calcareous rock. All of them have Hemisphere.
Introduction The genus Thyrea A. Massal. originally included various squamulose, foliose or small fruticose cyanolichens in the family Lichinaceae. The genus as circumscribed by Zahlbruckner (1925) also included species representing discordant elements that are not closely related. Subsequently, some of them have been transferred to other genera
M. Schultz and B. Budel: FB Biologie, Abt. Allgemeine Botanik, Universitat Kaiserslautern, Postfach 3049, D-67653 Kaiserslautern, Germany. S. Porembski: FB Biologie, Abt. Allgemeine und Spezielle Botanik, Institut fur Biodiversitatsforschung, Universitat Rostock, Wismarsche Str. 8, D-18051 Rostock, Germany. 0024-2829/01/030211+04 S35.00/0
Material and Methods Cryotome sections (16-20 um thick) were stained with lactophenol cotton blue. Anatomical measurements were made using semi-permanent mounts. The iodine reaction was tested after pretreatment with KOH. © 2001 The British Lichen Society
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TABLE 1. Species of Thyrea, occurrence and substratum
Species asahinae (Yoshim.) Yoshim. confusa Henssen deusta (Tuck.) Zahlbr. girardi (Dur. & Mont.) Bagl. & Carestia indica D. D. Awasthi & S. R. Singh latissima Asahina leptophylla (Tuck.) Zahlbr. mongolica H. Magn. nummularioides (Nyl.) A. Massal. pachyphytta (Mull. Arg.) Henssen plectopsora A. Massal.
plicatissima (Nyl.) Zahlbr.
Distribution
Substratum
Japan, E China Northern Hemisphere, Socotra Isl. Cuba Mediterranean region India Japan Cuba Mongolia Algeria Mediterranean region, Sonoran Desert Mediterranean region, Socotra Isl. Mediterranean region, Socotra Isl.
Siliceous rock Calcareous rock Limestone Calcareous rock Siliceous rock Calcareous rock
Micrographs were taken with a Zeiss (Jena) Axioskop compound microscope. Macrographs were taken with a Nikon F3 camera with a Zeiss Luminar 40 mm lens mounted on bellows.
Calcareous Calcareous Calcareous Calcareous Calcareous
rock rock rock rock rock
and folded, lobes rotundate (Figs 1 & 2), 1-5 mm wide; loosely attached to the substratum by bundles of rhizohyphae. Thallus structure heteromerous, ecorticate (Fig. 5); central strand thin, 25(-75) (im, composed of a few periclinally arranged hyphae, Results and Discussion without photobiont cells. Thallus periphery Thyrea porphyrella M. Schultz, Biidel with reticulate anatomy; fungal cells elon& Porembski sp. nov. gated, angular, 5-7-5 x 1-5 um. Photobiont a Thallus foliaceus, rubro-fuscus, porphyreus. Lobis unicellular cyanobacterium {Chroococcales), rotundatis, planis, inaequalibus vel plicatis; lobi ad 3-5 um across without sheath, 5-7-5 (im 5 mm lati. Thallus heteromerus; hyphis in centro thallo across including sheath; gelatinous sheath periclinis, ad marginem reticulatis. Cyanobacteria unicellularis, gelatina ad marginem lobi rubra; in centro thin, reddish (KOH + dirty violaceous) and thalli hyalina. Apotheciis raris, parvis; discus niger, with concentric granules at the thallus planus. Margo thallinus indistinctus. Hymenium hyali- periphery, hyaline in the thallus centre. num, in iodo caerulescens; excipulum proprium tenue. Apothecia rare, small, up to 0-35 mm Asci octospori, cylindracei, in iodo non tincti, leptoderdiam., up to 10 laminally on thallus lobes, mati. Ascosporae unicellulares, hyalinae, ellipsoideae vel globosae, parvae, 7-5-10(-12-5) x 7-5-10 urn. Pyc- immersed to semi-immersed; thalline marnidia immersa, globosa, ostiolum nigrescens; conidia gin flat and indistinct, up to 0-05 mm wide; ellipsoidea, 3 x 1 jxm. apothecial disc fiat, black, sometimes with Typus: Brazil, Est. Espirito Santo, near Pancas, colonies of free-living cyanobacteria. EpiFazenda Palmeiros, on exposed granite outcrop, c. 150 m alt, 19TS, 40°5'W, 11 April 1996, 5. Porem- hymenium tinged dirty olive, up to 10 um bski (2914), G. Martinelli & L. Leitman (RB 321989— high, KOH — . Hymenium (Fig. 4) hyaline, holotypus; B—isotypus). Guyana, Shea rock, on granite 85-100 um high, IKI+ blue. Subhymenium inselberg, 2°50'N, 59°9'W, 1997, F. Scheplitz (herb. hyaline, 10-25 (am high, IKI + blue. ParaPorembski, herb. Biidel, B, BM, M—paratypi). physes (Fig. 4) straight, sparsely branched and anastomosing, apically often furcate, (Figs 1-6) apical cell thickened, 3-5 urn wide. Exciple Thallus reddish brown to crimson, gelati- (Fig. 4) thin, 10-15 um wide, hyaline or nous when wet, foliose, polyphyllous, irregu- slightly brownish, of thin, gelatinous, interlarly branched and incised (Figs 1 & 3). woven hyphae. Asci (Fig. 4) cylindrical to Lobes ± flat at thallus margins (Figs 1 & narrowly clavate, 60-65 x 5-12-5 (im, ascus 2), irregularly ascending centrally (Fig. 3). wall thin throughout, IKI — . Ascospores 8, Lobe surface smooth or uneven, plicate simple, hyaline, globose to broadly ellipsoid,
FIGS 1-6. Thyrea porphyrella. 1, foliose thallus (holotype); 2 & 3, foliose thallus, lobes smooth, uneven to plicate, pycnidia indicated by arrow {Scheplitz 1997); 4, apothecium, old, thick-walled ascospores, paraphyses, excipulum (holotype); 5, thallus anatomy with thin, compact central strand {Scheplitz 1997); 6, ascogones beneath pycnidium, elongated conidiophores indicated by arrow (holotype). Scales: 1-3 = 3 mm, 4-6 = 25 |im.
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7-5-10(-12-5) x 7-5-10 um, spore wall distinctly thickened with age, up to 2-5 um wide (Fig. 4). Pycnidia immersed, globose, colourless, c. 0-1 mm wide, ostiolum dark coloured. Conidia simple, hyaline, ellipsoid or bacilliform, 3 x 1 urn, produced terminally on simple conidiophores. Ascomata develop from ascogonia formed beneadi pycnidia (Fig. 6), elongated conidiophores functioning as 'primary' paraphyses (Fig. 6) in juvenile apothecia (terminology according to Henssen 1963). Etymology. The species name is a diminutive indicating similarities in thallus shape and colour with the red alga Porphyra.
Vol. 33
America has a squamulose thallus and paraplectenchymatous anatomy of vertically arranged cells. Thyrea porphyrella was reported from Guyana by Schultz et al. (2000) under the name Thyrea sp. In spite of the large number of thalli present in the Guyanan collection, no apothecia were seen. However, juvenile stages in ascoma development with ascogonia beneath pycnidia were found. This observation justified its inclusion in Thyrea s.str., but the authors refrained from describing a new species at that time. We thank G. Martinelli and M. Leitman (both Rio de Janeiro), M. Jansen-Jacobs (Utrecht) and F. Scheplitz (Bonn) for collecting material and for field guidance.
Ecology and Distribution. Thyrea porphyrella
grows on granite inselbergs in exposed situations. To date, it is known from two inselbergs, one in SE Brazil and the other in southern Guyana. However, Thyrea porphyrella may occur in similar sites across tropical and subtropical regions of South America. The species is accompanied mainly by freeliving cyanobacteria (species of Gloeocapsa, Scytonema and other genera) but also by cyanobacterial lichens, Peltula auriculata,
REFERENCES
Asahina, Y. (1958) Lichenologische Notizen (§ 139). Journal of Japanese Botany 33: 67-69. Henssen, A. (1963) Eine Revision der Flechtenfamilien Lichinaceae und Ephebaceae. Symbolae Botanicae Upsalienses 18: 1-123. Henssen, A. (1968) Thyrea radiata, eine LempholemmaArt mit Hormocystangien. Berichte der Deutschen Botanischen Gesellschaft 81: 176-182. Henssen, A. (1979)[1980] Problematik der Gattungsbegrenzung bei den Lichinaceen. Berichte der Deutschen Botanischen Gesellschaft 92: 483-506. Henssen, A. (1986) The genus Paulia (Lichinaceae). Phylliscum vermiformis, Pterygiopsis guyanensis Lichenologist 18: 201-229. Henssen, A. & Jorgensen, P. M. (1990) New com(Schultz et al. 2000). binations and synonyms in the Lichinaceae. Lichenologist 22: 137-147. Notes. Other species of Thyrea have a Moreno, P. P. & Egea, J. M. (1992a) El genero greyish or blackish thallus colour and lack Lichinella Nyl. en el sureste de Espana y norte de Africa. Cryptogamie, Bryologie-Lichenologie 13: the distinct reddish tinge of T. porphyrella 237-259. that is caused by pigments in the gelatinous Moreno, P. P. & Egea, J. M. (19926) Digitothyrea, a sheaths of the photobiont cells. Thyrea plicanew genus in the family Lichinaceae. Lichenologist tissima (Nyl.) Zahlbr. has similarly thin thal24: 215-228. lus lobes but differs in the distinctly Moreno, P. P. & Egea, J. M. (1992c) Estudios sobre el complejo Anema-Thyrea-Peccania en el sureste de granulose surface and a preference for calla Peninsula Iberica. Acta Botanica Barcinonensia careous substrata (Moreno & Egea 1992c). 41: 1-66. Lichinella hondoana (Zahlbr.) Moreno & Schultz, M., Porembski, S. & Biidel, B. (2000) Egea, and L. cribellifera (Nyl.) Moreno & Diversity of rock-inhabiting cyanobacterial lichens: Egea, are similar in external appearance but studies on granite inselbergs along the Orinoco and in Guyana. Plant Biology 2: 482-495. have thallinocarpous ascomata and polysporous asci with smaller spores (<7-5 um Yoshimura, I. (1968) Japanese species of Anema. Journal of Japanese Botany 43: 354-358. long). Phylliscidium monophyllum (Kremp.) Zahlbruckner, A. (1925) Catalogus Lichenum Forssell from similar localities in South Universalis. Vol. 3. Leipzig: Borntraeger.
Accepted for publication 7 December 2000
Thyrea porphyrella, a new species of the Lichtnaceae from inselbergs in tropical South America M. SCHULTZ, B. BUDEL and S. POREMBSKI Abstract: The new species Thyrea porphyrella Schultz, Budel & Porembski, in the Lichinaceae, is described from tropical South America. It grows on granite inselbergs in SE Brazil and in southern Guyana. Diagnostic characters are the very thin, superficially smooth but conspicuously uneven to folded thallus lobes of reddish-brown to crimson colour. £ 2001 The British Lichen Society
gelatinous, ecorticate, heteromerous thalli with a loose or compact central strand. Their type of ascoma development is unique to approximately half of the Lichinaceae. Ascogonia are formed beneath pycnidia, and the pycnidia are then transformed into apothecia (so-called pycnoascocarps, cf. Henssen 1963). Table 1 gives an overview of the species currently treated under Thyrea s. str. and their distribution. The species Thyrea of the Lichinaceae such as Gonohymenia, arenae A. Massal. ex Forssell, T. borzii Lempholemma, Lichinella, Paulia and Beltr. and T. camaromorpha (A. Massal.) Peccania (Henssen 1968, 1979, 1986;A. Massal. are dubious. Thyrea latissima Henssen & Jorgensen 1990; Moreno & Asahina was transferred to Anema by Egea 1992a). The genus Digitothyrea was Yoshimura (1968) due to the similar shape described by Moreno & Egea (19926) of the hyphae. However, in our opinion, the for those species of Thyrea s. lat. having a original placement by Asahina (1958) based ± fruticose growth form with elongated, on the heteromerous thallus structure better tongue-shaped lobes and a type of ascoma reflects the generic limits (at least for the ontogeny starting with ascogonia formed in moment) and hence should be maintained. a tangle of generative hyphae. Currently, In this paper we describe a new species of Thyrea s. str. contains approximately 12 Thyrea from two granite inselbergs in species that are quite large and foliose or tropical South America. These are the first small squamulose lichens predominantly records of Thyrea s.str. from the Southern colonizing calcareous rock. All of them have Hemisphere.
Introduction The genus Thyrea A. Massal. originally included various squamulose, foliose or small fruticose cyanolichens in the family Lichinaceae. The genus as circumscribed by Zahlbruckner (1925) also included species representing discordant elements that are not closely related. Subsequently, some of them have been transferred to other genera
M. Schultz and B. Budel: FB Biologie, Abt. Allgemeine Botanik, Universitat Kaiserslautern, Postfach 3049, D-67653 Kaiserslautern, Germany. S. Porembski: FB Biologie, Abt. Allgemeine und Spezielle Botanik, Institut fur Biodiversitatsforschung, Universitat Rostock, Wismarsche Str. 8, D-18051 Rostock, Germany. 0024-2829/01/030211+04 S35.00/0
Material and Methods Cryotome sections (16-20 um thick) were stained with lactophenol cotton blue. Anatomical measurements were made using semi-permanent mounts. The iodine reaction was tested after pretreatment with KOH. © 2001 The British Lichen Society
212
THE LICHENOLOGIST
Vol. 33
TABLE 1. Species of Thyrea, occurrence and substratum
Species asahinae (Yoshim.) Yoshim. confusa Henssen deusta (Tuck.) Zahlbr. girardi (Dur. & Mont.) Bagl. & Carestia indica D. D. Awasthi & S. R. Singh latissima Asahina leptophylla (Tuck.) Zahlbr. mongolica H. Magn. nummularioides (Nyl.) A. Massal. pachyphytta (Mull. Arg.) Henssen plectopsora A. Massal.
plicatissima (Nyl.) Zahlbr.
Distribution
Substratum
Japan, E China Northern Hemisphere, Socotra Isl. Cuba Mediterranean region India Japan Cuba Mongolia Algeria Mediterranean region, Sonoran Desert Mediterranean region, Socotra Isl. Mediterranean region, Socotra Isl.
Siliceous rock Calcareous rock Limestone Calcareous rock Siliceous rock Calcareous rock
Micrographs were taken with a Zeiss (Jena) Axioskop compound microscope. Macrographs were taken with a Nikon F3 camera with a Zeiss Luminar 40 mm lens mounted on bellows.
Calcareous Calcareous Calcareous Calcareous Calcareous
rock rock rock rock rock
and folded, lobes rotundate (Figs 1 & 2), 1-5 mm wide; loosely attached to the substratum by bundles of rhizohyphae. Thallus structure heteromerous, ecorticate (Fig. 5); central strand thin, 25(-75) (im, composed of a few periclinally arranged hyphae, Results and Discussion without photobiont cells. Thallus periphery Thyrea porphyrella M. Schultz, Biidel with reticulate anatomy; fungal cells elon& Porembski sp. nov. gated, angular, 5-7-5 x 1-5 um. Photobiont a Thallus foliaceus, rubro-fuscus, porphyreus. Lobis unicellular cyanobacterium {Chroococcales), rotundatis, planis, inaequalibus vel plicatis; lobi ad 3-5 um across without sheath, 5-7-5 (im 5 mm lati. Thallus heteromerus; hyphis in centro thallo across including sheath; gelatinous sheath periclinis, ad marginem reticulatis. Cyanobacteria unicellularis, gelatina ad marginem lobi rubra; in centro thin, reddish (KOH + dirty violaceous) and thalli hyalina. Apotheciis raris, parvis; discus niger, with concentric granules at the thallus planus. Margo thallinus indistinctus. Hymenium hyali- periphery, hyaline in the thallus centre. num, in iodo caerulescens; excipulum proprium tenue. Apothecia rare, small, up to 0-35 mm Asci octospori, cylindracei, in iodo non tincti, leptoderdiam., up to 10 laminally on thallus lobes, mati. Ascosporae unicellulares, hyalinae, ellipsoideae vel globosae, parvae, 7-5-10(-12-5) x 7-5-10 urn. Pyc- immersed to semi-immersed; thalline marnidia immersa, globosa, ostiolum nigrescens; conidia gin flat and indistinct, up to 0-05 mm wide; ellipsoidea, 3 x 1 jxm. apothecial disc fiat, black, sometimes with Typus: Brazil, Est. Espirito Santo, near Pancas, colonies of free-living cyanobacteria. EpiFazenda Palmeiros, on exposed granite outcrop, c. 150 m alt, 19TS, 40°5'W, 11 April 1996, 5. Porem- hymenium tinged dirty olive, up to 10 um bski (2914), G. Martinelli & L. Leitman (RB 321989— high, KOH — . Hymenium (Fig. 4) hyaline, holotypus; B—isotypus). Guyana, Shea rock, on granite 85-100 um high, IKI+ blue. Subhymenium inselberg, 2°50'N, 59°9'W, 1997, F. Scheplitz (herb. hyaline, 10-25 (am high, IKI + blue. ParaPorembski, herb. Biidel, B, BM, M—paratypi). physes (Fig. 4) straight, sparsely branched and anastomosing, apically often furcate, (Figs 1-6) apical cell thickened, 3-5 urn wide. Exciple Thallus reddish brown to crimson, gelati- (Fig. 4) thin, 10-15 um wide, hyaline or nous when wet, foliose, polyphyllous, irregu- slightly brownish, of thin, gelatinous, interlarly branched and incised (Figs 1 & 3). woven hyphae. Asci (Fig. 4) cylindrical to Lobes ± flat at thallus margins (Figs 1 & narrowly clavate, 60-65 x 5-12-5 (im, ascus 2), irregularly ascending centrally (Fig. 3). wall thin throughout, IKI — . Ascospores 8, Lobe surface smooth or uneven, plicate simple, hyaline, globose to broadly ellipsoid,
FIGS 1-6. Thyrea porphyrella. 1, foliose thallus (holotype); 2 & 3, foliose thallus, lobes smooth, uneven to plicate, pycnidia indicated by arrow {Scheplitz 1997); 4, apothecium, old, thick-walled ascospores, paraphyses, excipulum (holotype); 5, thallus anatomy with thin, compact central strand {Scheplitz 1997); 6, ascogones beneath pycnidium, elongated conidiophores indicated by arrow (holotype). Scales: 1-3 = 3 mm, 4-6 = 25 |im.
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THE LICHENOLOGIST
7-5-10(-12-5) x 7-5-10 um, spore wall distinctly thickened with age, up to 2-5 um wide (Fig. 4). Pycnidia immersed, globose, colourless, c. 0-1 mm wide, ostiolum dark coloured. Conidia simple, hyaline, ellipsoid or bacilliform, 3 x 1 urn, produced terminally on simple conidiophores. Ascomata develop from ascogonia formed beneadi pycnidia (Fig. 6), elongated conidiophores functioning as 'primary' paraphyses (Fig. 6) in juvenile apothecia (terminology according to Henssen 1963). Etymology. The species name is a diminutive indicating similarities in thallus shape and colour with the red alga Porphyra.
Vol. 33
America has a squamulose thallus and paraplectenchymatous anatomy of vertically arranged cells. Thyrea porphyrella was reported from Guyana by Schultz et al. (2000) under the name Thyrea sp. In spite of the large number of thalli present in the Guyanan collection, no apothecia were seen. However, juvenile stages in ascoma development with ascogonia beneath pycnidia were found. This observation justified its inclusion in Thyrea s.str., but the authors refrained from describing a new species at that time. We thank G. Martinelli and M. Leitman (both Rio de Janeiro), M. Jansen-Jacobs (Utrecht) and F. Scheplitz (Bonn) for collecting material and for field guidance.
Ecology and Distribution. Thyrea porphyrella
grows on granite inselbergs in exposed situations. To date, it is known from two inselbergs, one in SE Brazil and the other in southern Guyana. However, Thyrea porphyrella may occur in similar sites across tropical and subtropical regions of South America. The species is accompanied mainly by freeliving cyanobacteria (species of Gloeocapsa, Scytonema and other genera) but also by cyanobacterial lichens, Peltula auriculata,
REFERENCES
Asahina, Y. (1958) Lichenologische Notizen (§ 139). Journal of Japanese Botany 33: 67-69. Henssen, A. (1963) Eine Revision der Flechtenfamilien Lichinaceae und Ephebaceae. Symbolae Botanicae Upsalienses 18: 1-123. Henssen, A. (1968) Thyrea radiata, eine LempholemmaArt mit Hormocystangien. Berichte der Deutschen Botanischen Gesellschaft 81: 176-182. Henssen, A. (1979)[1980] Problematik der Gattungsbegrenzung bei den Lichinaceen. Berichte der Deutschen Botanischen Gesellschaft 92: 483-506. Henssen, A. (1986) The genus Paulia (Lichinaceae). Phylliscum vermiformis, Pterygiopsis guyanensis Lichenologist 18: 201-229. Henssen, A. & Jorgensen, P. M. (1990) New com(Schultz et al. 2000). binations and synonyms in the Lichinaceae. Lichenologist 22: 137-147. Notes. Other species of Thyrea have a Moreno, P. P. & Egea, J. M. (1992a) El genero greyish or blackish thallus colour and lack Lichinella Nyl. en el sureste de Espana y norte de Africa. Cryptogamie, Bryologie-Lichenologie 13: the distinct reddish tinge of T. porphyrella 237-259. that is caused by pigments in the gelatinous Moreno, P. P. & Egea, J. M. (19926) Digitothyrea, a sheaths of the photobiont cells. Thyrea plicanew genus in the family Lichinaceae. Lichenologist tissima (Nyl.) Zahlbr. has similarly thin thal24: 215-228. lus lobes but differs in the distinctly Moreno, P. P. & Egea, J. M. (1992c) Estudios sobre el complejo Anema-Thyrea-Peccania en el sureste de granulose surface and a preference for calla Peninsula Iberica. Acta Botanica Barcinonensia careous substrata (Moreno & Egea 1992c). 41: 1-66. Lichinella hondoana (Zahlbr.) Moreno & Schultz, M., Porembski, S. & Biidel, B. (2000) Egea, and L. cribellifera (Nyl.) Moreno & Diversity of rock-inhabiting cyanobacterial lichens: Egea, are similar in external appearance but studies on granite inselbergs along the Orinoco and in Guyana. Plant Biology 2: 482-495. have thallinocarpous ascomata and polysporous asci with smaller spores (<7-5 um Yoshimura, I. (1968) Japanese species of Anema. Journal of Japanese Botany 43: 354-358. long). Phylliscidium monophyllum (Kremp.) Zahlbruckner, A. (1925) Catalogus Lichenum Forssell from similar localities in South Universalis. Vol. 3. Leipzig: Borntraeger.
Accepted for publication 7 December 2000