Lichenologist 26(3): 223-282 (1994) STUDIES ON THE LICHEN GENUS STICTA (SCHREBER) ACH.: I. SOUTHERN SOUTH AMERICAN SPECIES D. J. GALLOWAY* Abstract: Twelve species of Sticta are recorded from southern South America (Argentina and Chile including Juan Fernandez) south of latitude 30°S, viz., S. ainoae, 8. caulescens, S. fuliginosa, S. gaudichaldia, S. hypochra, 8. hypopsila, S. Fineariloba, S. limbara, S. longipes, S, sublimbara and S. weigelii, and S. santessonii D. Galloway is newly described. A key is provided, and details of anatomy, morphelogy, distribution, ecology and taxonomy are discussed. Introduction Species of Sticta (Schreber) Ach. are large, conspicuous, dorsi-ventral, foliose lichens characterized by a + tomentose lower surface with cyphellae as con- stantly occurring structures. The presence of cyphellac (Fig. 1) distinguishes Sticta from the superficially similar genera with which it has often been confused, viz. Lobaria (Schreber) Hoffm. and Pseudocyphellaria Vainio. Although the structure of ascospores and conidia are similar in all three genera (Lindsay 1859, 1869; Hue 1901; Vobis & Hawksworth 1981; Galloway 19886), the presence of cyphellae in Sticra correlates with an absence of acetone-soluble secondary metabolites and contrasts markedly with the situation in Lobaria and Pseudocyphellaria. The latter has pseudocyphellae (Fig. 1) on the lower surface and a richly diverse chemistry (Galloway 19884, 19915), whereas Lobaria has neither cyphellae nor pscudocyphellae on the lower surface, and a less diverse chemistry (Yoshimura 1971; James & Henssen 1976; Culberson er al. 1977; Galloway 1981). On chemical grounds, the diversity of acetone-soluble secondary compounds in Lobaria and Pseudocyphellaria suggests a closer affinity of these two genera to each other than either shows with Szicta, and for this reason, and correlating morphological considerations, species of Sticta should be placed in the family Stictaceae Zahlbr., with Lobaria and Pseudocyphellaria accommodated in the family Lobariaceae Chev., an arrange- ment earlier suggested by James & Henssen (1976) and Galloway (1985a, 1988b, 19915). Although a number of lichen collections were made by eighteenth century navigators and their naturalists from southern South America (Galloway 19924) only Archibald Menzies (1754-1842) collected specimens of Sticta. Employed as a naval surgeon during Captain Colnett’s Prince of Wales voyage of 1787-1789 (Galloway & Groves 1987), his February 1787 collections of S. caulescens, S. fuliginosa and S. gaudichaldia from an anchorage in Staten Island are the only known 18th century collections from cool temperate South America. *Department of Botany, The Natural History Museum, Cromwell Road, London SW7 5BD,UK. 0024-2829/94/030223 + 60 $08.00/0 © 1994 The British Lichen SocietyTHE LICHENOLOGIST Vol, 26 224 Cyrnentar. Fic. 1. Structure of pseudocyphella (1) and cyphella (2) from Schneider (1987 fig. 2) Inthe 19th century, British and French scientific expeditions to the Southern Hemisphere during the first five decades of the century (Galloway 19856) produced numerous published reports on their discov among which were eries names of South American species of Sticra. Sticta gaudichaldia was the first to be described (Delise 1825) from collections made in the Falkland Islands by Gaudichaud-Beaupré. Later the same year Fries (1825) published S. malovina and S. magellanica from S. gaudichaldia) from the Falkland Islands, Tierra del Fuego. The type (UPS) of the latter is a tiny fertile fragment that is ¢ presently impossible to ascribe to any known South American species with1994 Southern South American Sticta—Galloway 225, certainty, It is most closely similar to S, ambavillaria from East Africa and the Indian Ocean Islands, and the marginal arrangement and appearance of the fruits given in the illustration of this species in Swinscow & Krog (1988: fig. 152) is strongly reminiscent of the apothecia seen in the small fragment of S. magellanica, a fact earlier reported by Miiller Argoviensis (1887). In my opinion S. magellanica should be synonymized with S. ambavillaria (Bory) Ach., since it seems highly likely that Fries mixed material in his herbarium and that the provenance given of S. magellanica is in error. In a preliminary account of lichens collected by Carlo Bertero from Juan. Fernandez, Montagne (1835) recorded a green, stalked species, which he called S. filicina, noting that it had characteristics both of the New Zealand species S. filix and of the non-stalked Caribbean species S. damaecornis. For the stalked Juan Fernandez material he later described (Montagne 1852) two varieties of S. filicina, viz.; vars lineariloba and hypopsila, which are recognized in this account ‘as two separate species. Joseph Hooker in his account of lichens from the Falkland Islands and Tierra del Fuego discusses S$. filicina from Hermite Island in the context of a stalked, isidiate, cyanobacterial taxon, now referred to as S. hypockra. This erroneous interpretation of S. filicina was subsequently widely used in the literature relating to South American Sricta. As has been shown earlier (Galloway 1985a) S. filicina is itself a synonym of the endemic New Zealand species S. filix (Sw.)Nyl. Astalked, cyanobacterial species, 8. caulescens, was described by De Notaris (1851) from collections made near Valparaiso (a second new species, S. biatora De Not., is a synonym of S. caulescens), and Montagne (1854) recorded S. fuliginosa from southern South America for the first time. Crombie (1876) described S. filicina f. latissima (= broad-lobed S. caulescens)and S. fuliginosa f. Jfirmior (= S. hypochra) from Chilean collections made by R. O. Cunningham. ‘A further cyanobacterial, stalked species from Valdivia was named S. longpipes by Miiller Argoviensis (1882). At the close of the nineteenth century the following species of Sticta were known from southern South America: S. hypopsila and S. lineariloba from Juan Fernandez, an undescribed green algal species from Chile was incorrectly named S. damaecornis (see Galloway & Pickering 1990), and the remainder were the cyanobacterial species S. caulescens, S. filicina, S. fuliginosa, S. gaudichaldia, and S, longipes (Delise 1825; Fries 1825; Montagne 1835, 1852, 1854; Hooker & Taylor 1844; Hooker 1847; De Notaris 1851; Nylander 1855a,6, 1858, 1860, 1868, 1888; Crombie 1876, 1877, 1883; Krempelhuber 1876; Miiller Argoviensis 1882, 1883, 1887, 1888, 1889a,b; Jatta 1890; Hue 1890; Stizenberger 1895; Malme 1899). In the present century only very few new taxa in Sticta from southern South America were introduced. Vainio (1903) described S. hypochra for the cyanobacterial, stalked, isidiate species widely recorded earlier as S. filicina, and Jatta (1906) described S. quercizans var. glaucovirens for the same species. Additional new names were introduced by Rasiinen (1937, 1939), viz. Stictina caulescens f. pallidoglauca, Stictina caulescens f. brunnea (=S. caulescens), Stictina filicinetla vat. ochraceofusca (=S. hypochra). Galloway & Pickering (1990) described S. ainoae for the South American taxon widely and incorrectly reported as S. damaecornis.226 THE LICHENOLOGIST Vol. 26 Prior to the present revision, taxa recorded from southern South America this century included: S. ainoae, S. caulescens, S. fuliginosa, S. gaudichaldia, S. hypochra, 8. hypopsila, S. lineariloba, S. longipas and S. weigeliit (Hue 1901; Vainio 1903; Jatta 1906; Darbishire 1912; ZahIbruckner 1917, 1924, 1925, 1926, 1933; Cengia Sambo 1926a,b; Du Rietz 1926; Rasiinen 1932, 1936, 1937, 1939; Grassi 1950; Santesson 1954; Skottsberg 1956; Lamb 1959; Dodge 19653 Follmann 1966; Redon 1972, 1974, 1976; Galloway & Pickering 1990). The present study is based on a revision of material from South America south of latitude 30°S from both Argentina and Chile, including the Falkland Islands and Juan Fernandez. Materials and Methods ‘Typeand other material, about 475 collections, were studied from the following herbaria: BA, BM, BR, CONG, E, FH, G, H,H-NYL, KR, LINN-SM, M,NY, PC, 8, SGO, TNS, TUR, UPS, US, W. Scanning electron microscopy was performed on air-dried material coated with gold-palladium ‘on aluminium stubs using a Hitachi S-800 microscope. Morphology and Anatomy Caulescent taxa: In Sticta, a number of species are distinctly stalked, being attached to the substratum by a holdfast supporting the stalk from which the thallus lobes expand. The development of a holdfast and stalk is of considerable interest in Sticta since it is found in the macrolichens only in this genus. It is possible that stalk formation is an adaptation whereby the photosynthetic tissues of the lichen are held above competing bryophytes on tree trunks with a heavy epiphyte burden, and above litter on the forest floor when they are attached to stones and soil. In such low-light situations the presence of a stalk may be advantageous for what is basically a foliose lichen in exposing the photosynthetic tissues to the optimum amount of light. It is interesting, however, that other foliose genera (Nephroma, Pseudocyphellaria) capable of growing in very low-light habitats, do not show any tendency towards stalk formation, which means that the adaptation is not, or not only, @ response to low levels of light. Five species (40% of the total) in southern South America are stalked, viz.: S. caulescens, S. hypochra, S. hypopsila, S. lineariloba and S. longipes. Of these, two have green photobionts (S. hypopsila, S. lineariloba) and are endemic to Juan Fernandez, whereas the remainder have cyanobacterial photobionts, and are endemic in cool temperate forest habitats on the mainland of southern South America, not being recorded from Juan Fernandez. Sticta santessonii develops froma short, evanescent stalk in terricolous habitats but this attaching structure appears to be quickly lost as growth of the thallus proceeds. Rarely, specimens of S. ainoae may have a short stalk, especially when forming a photosymbiodeme with Dendriscocaulon (Fig. 2). This has so far been observed only twice. The South American temperate lichen flora has a higher percentage of stalked species of Sticta than does New Zealand, which has a similar number of species overall. In the New Zealand lichen flora only three species of Sticta are stalked (259%), and all of these have green photobionts (Galloway 19852).1994 Southern South American Sticta—Galloway 227 FIG. 2. Photosymbiodeme of Stictaainoae and Dendriscocaulon (S-R. Santesson 7604). Scale =5 mm. Upper cortex: In Sticta, as in Pseudocyphellaria (Galloway 19880: 19), the upper cortex in all species is a paraplectenchyma (pseudoparenchyma) of anti- clinically arranged, round to irregular, + isodiametric, + angularly thickened cells, 25-45 wm diam., with the outermost cells rather smaller and + com- pressed, and with somewhat thicker walls than the larger more loosely arranged cells adjacent to the photobiont layer (Fig. 3). Cell wall thickness is in the range 1-5-3-5 um, with lumina round to irregular. Scanning electron micrographs of the upper surface show the outlines of cells from the underlying tissue with a continuous cortical covering (epicortex), or else the outer cortical covering has small, scattered rather irregular pores. For detailed discussion of pored and228 THE LICHENOLOGIST Vol. 26 Fig. 3. Sticta limbata upper cortex and photobiont iayer ( x 1500). continuous epicortices see Hale (1981, 1987) and Angelsea er al. (1982). Species of Sticta from southern South America with a + continuous outer cortical layer include: S. hypopsila, S. limbata, 8. lineariloba, S. santessonii and S. weigelii (Fig. 4A-D). Taxa with pores in the outer cortical layer are: S. ainoae, S. caulescens, S. fuliginosa, S. gaudichaldia, S. hypochra, S. longipes and S. sublimbata (Fig. 5A-D). Lower cortex: This is similar in structure to the upper cortex but generally thinner; the outermost layer of cells is sometimes pigmented red-brown, giving rise to tomental hairs (Fig. 6). Isidia: These are terete in cross-section and are small outgrowths of the thallus with an outer cortex and internal photobiont and medullary layers and may be simple, + finger-like or only sparingly branched, as in S. fuliginosa and S. santessonii (Fig. 7A-B), to coralloid-branched, as in S. hypochra and S. wiegelii (Fig. 7C). Isidia of S. longipes are rather granular with a thin cortex and are readily eroded (Fig. 7D), appearing pseudosorediate. Isidia in S. fuliginosa (Fig. 8), S. hypochra (Fig. 9) and S. longipes (Fig. 10) collected from shaded and very humid sites may become distinctly lobulate or flattened-phyllidiate toFic. 4, Continuous upper cortex: A, Sticta hypopsilas B, S. imbara, C, 8. lineariloba; D, S. weigelit £1000). coralloid and Dendriscocaulon-like. The minute granular-coralloid isidia in S. sublimbata soon erode to form granular soredia. Soredia: Soredia are present in only two species: S. limbata and S. sublimbata (Fig. 11A-B). In S. limbata soralia are present as laminal and marginal erose patches, the soredia not being associated with the breakdown of isidia, whereas in S. sublimbata the soralia are primarily linear and marginal, the soredia arising from the breakdown of granular-coralloid isidia. Photosymbiodemes: In contrast to South American species of Pseudocyphellaria (Galloway 19926) only two photosymbiodemes were found in all of the collec- tions of South American species of Sticta examined, viz., young, atypically stalked specimens of S. ainoae from Laguna San Rafael, which were associated230 THE LICHENOLOGIST Vol. 26 Fic, 5, Pored upper cortex: A, Sticta ainoae; B, S. fuliginosa; C, S. hypochra; D, S. sublimbata (1000). with Dendriscocaulon [Galloway 9050 (BM)] and a similar photosymbiodeme from dense Valdivian rainforest at Enco [Santesson 7604 (S): Fig. 12]. As in other Dendriscocaulon-Sticta photosymbiodemes (James & Henssen 1976), the leaflets of the green S. ainoae were growing out of the basal Dendriscocaulon. Both collections are from shaded habitats of constant high humidity. In con- trast to the situation reported for S. filix-Dendriscocaulon from New Zealand (James & Henssen 1976), where the dominant expression of the lichen fungus is as the cyanobacterial Dendriscocaulon bearing secondary leaflets of the green S. filix, the South American photosymbiodemes seen have the green S. atnoae as the dominant expression of the lichen fungus. It seems, however, to be a rare phenomenon in South American populations of Sticta. Photobiont versatility in species of Szicta was first discussed at length by James & Henssen (1976), who pointed out that species with a green primary1994 Southern South American Szicta—Galloway 231 FIG. 6. Sticta libata lower cortex (x 1000). photobiont also have smaller amounts of cyanobacterial photobionts present in internal cephalodia. The physiological advantages of photobiont versa- tility with respect to the genera Sticta and Pseudocyphellaria in Southern Hemisphere temperate rainforest ecosystems is reviewed by Green & Lange (1991). Cyphellae: These structures are a characteristic feature of the genus and have well-defined margins and a cellular membrane lining the floor of the cyphella separating the medullary tissue from the exterior (Fig. 1). In southern South American species two major kinds of cyphellae are found: (i) with thick, prominent, raised margins projecting above the tomentum of the lower surface and bordering a rather narrow, deep pore, as seen in S. ainoae, S. caulescens, S. hypopsila, S. hypochra, S. lineariloba, S. longipes and S. santessonii (Fig. 12A-D)} (ii) with thin, rather narrow margins often not much raised above the tomentum, bordering a rather shallow often very broad pore as in S. fuliginosa, S. gaudichaldia, S. limbata, S. sublimbata and S. weigelii (Fig. 13A-D). Tomentum: These are strands of thick-walled, septate, generally unbranched, colourless to red-brown hyphae that grow outwards from the outermost cells of232 THE LICHENOLOGIST Vol. 26 Fi. 7. Isidia: A, Sticta santessonit ( x 2000); B, S. fuliginosa ( x 400}; C, S. Aypochra (x80); D, S. longipes (x 125). the lower cortex (usually in fascicles o£2-12 or more) and thatanchor the thallus tothe substratum (Fig. 14A). Development of omentum may also be important in the increased storage capacity of water, which is known to correlate positively with rates of nitrogen fixation (Snelgar & Green 1981; Green & Lange 1991). Some species, such as S. sublimbata, have tomental hairs, which may be branched at the apices (Fig. 14B), and S. weigelii consistently has laminal protuberances or knobs on the tomental hairs (Fig. 14C), as shown by Harris (1987). Sticta and nutrient cycling Of the 12 species of Sicta recorded here from southern South America, nine (75%) have cyanobacteria as the primary photobiont and the remaining three1994 Southern South American Sticta—Galloway 233 F1. 8. Lobulate-phyllidiate isidia developing at margins of Sticta fuliginosa(S-R. Santesson1 15). Scale=I cm. species have cyanobacteria also present in internal cephalodia. Cyanobacterial photobionts allow the species to photosynthesize in low-light habitats and many of the species of Sticra found in cool temperate forests in South America are shade-adapted. The caulescent habit, seen in S. caulescens, S. hypochra, S. hypopsila, S. lineariloba and S. longipes allows these taxa to compete effec- tively with more rapidly growing bryophytes on the ground and in understorey habitats, since the stalk raises the lichen frond up to or above the level of the bryophyte mat. The biomass of species of Pseudocyphellaria and Sticta in cool temperate forest habitats in southern South Ametica is impressive (Rundell 1980; Galloway 19914,, 1992a) as is also the potential for nitrogen fixation by lichen cyanobacterial photobionts in these habitats (Galloway 19922,)). This has already been noted for analogous habitats in New Zealand forests (Green etal, 1980; Galloway 1985a, 19884,b, 1991a,5; Green & Lange 1991). Species of Sticta are known to produce quantities of amines (which impart a fishy smell to a number of taxa, especially those with cyanobacterial photobionts) and a recent report (Greenfield 1992) shows the New Zealand, green caulescent species S. latifrons to have 5%, of its dry weight as detectable nitrogen. It now seems likely that species of Sticta have an important role in the nitrogen budgets of cool temperate South American forests, through fixing234 THE LICHENOLOGIST Vol. 26 Fig. 9. Phyllidiate to Dendriscocaulon-like development of isidia in Sticta hypochra (S-R. Santesson 7605). Scale= 1 em. atmospheric nitrogen and either leaking it to the ground during rainfall, or eventually releasing soluble or combined nitrogen after decomposition in the litter (Guzman et al. 1990; Greenfield 1993). Phytogeographical notes Southern South American species of Sticta show three differentbiogeographical elements or affinities. (1) Endemic element: this is the dominant element in the Sticta-flora of the region, comprising 50% of the species: S. ainoae, S. caulescens, S. gaudichaldia, S. hypochra, S. longipes and S. santessonii. Sticta gaudichaldia, a grassland species, is predominanuy Fuegian, occurring also on the Falklands, whereas the remaining taxa occur mainly as epiphytes in rainforest of mainland South America, being best developed south of latitude 40°S. It may be noted here that all of the endemic taxa, with the exception of S. ainoae, have cyanobacterial photobionts, which allows them to colonize low-light habitats. This gives them a special importance in temperate rainforest where they are significant contributors to the nitrogen budget of the forest ecosystem. Associating1994 Southern South American Sticta—Galloway 235 FIG. 10. Phyllidiate to Dendriscocaulon-like marginal isidia in Sticta longipes(S-R. Santesson 7681). Scale in mm. lichens including taxa from the following genera: Cladina, Cladonia, Degetia, Menegazzia, Metus, Nephroma, Pannoparmelia, Protousnea, Pseudocyphellaria and Psoroma are discussed in Galloway (1992). (2) Juan Fernandez element; two species or 17% of the Sticta-flora are found only in Juan Fernandez, viz. S. hypopsila, S. lineariloba adding to the list of lichens endemic to the region (Galloway 1991a). (3) Cosmopolitan element: four species or 33% of the Sticta-flora are cosmopolitan, viz. S. fuliginosa, S. limbata, S. sublimbata, S. weigelii, although the latter two are often more common in tropical regions than they are in temperate areas.Fic. 11. Soredia: A, Sticta limbata; B, S. sublimbata ( x 1000), { iB Fic. 12. Narrow-pored eypheliae: A, Sticta ainoae (x 150); B, S. caulescens {x 60); C, 8. hypopsila (150); D, 8, longipes (50).1994 Southern South American Sticta—Galloway 237 Fic. 13. Broad-pored cyphellae: A, Sticta fuliginosa (x 100); B, S. gaudichaldia (x 150); C, S. Fimbata ( x 100); D, Sticta weigelit ( x 150). Iris noteworthy that no species with austral affinities are found in the Sticta flora of South America. In cool temperate areas, Sticra has a higher incidence of regional endemism than does Pseudocyphellaria and this appears to hold in tropical areas as well, although in the tropics as a general rule many more species of Sticra are found than are species of Pseudocyphellaria (Galloway & Arvidsson 1990). Although there are many more species of Pseudocyphellaria (53 taxa) in temperate South America than there are of Sticra (12 taxa), of these only 38% are endemics whereas half of the Sticta species in southern South America are endemic to the mainland and if we include Juan Fernandez endemics this total rises to 67%. In New Zealand for comparison, of the 12 species of Sticra presently known 50% are endemic (Galloway 19852), whereas 52% of the species of Pseudocyphellaria are endemic (Galloway 19886).238 ‘THE LICHENOLOGIST Vol. 26 Fi. 14. Tomental hairs: A, Sticea limbata (x 250); B, S. sublimbata (x 3000); C, S. weigelii (3000). ‘The genus Sticta (Schreber) Ach. (1803) Thallus foliose, heteromerous, dorsiventral, lobate, spreading or sometimes stalked with a fruticose, erect, monophyllous to polyphylious frond, some- times arising from a holdfast or appearing + unattached, + loosely attached, 2-10(-20) cm diam., saxicolous, terricolous, muscicolous or corticolous. Lobes dichotomously to irregularly branching, rounded to imbricate to variously incised, often lacerate-notched, with or without isidia, phyllidia or soredia,1994 Southern South American Sticta—Galloway 239 FiG. 15. Sticta ainoae (SGO-holotype). Scale=2 em. tough, coriaceous to fragile and papery, thin to thick. Upper surface smooth, wrinkled or obscurely ridged, sometimes shallowly faveolate or pitt punctate-impressed, shining or matt, often maculate, or with or without i phyllidia, or soredia, without pscudocyphellae. Medulla white. Photobiont green (trebouxioid) or cyanobacterial (Nostoc); in species with a green primary photobiont, internal cephalodia containing cyanobacteria are also regularly present, Lower surface pale or dark, glabrous or tomentose, occasionally with anchoring tufts or rhizines at margins and centrally, cyphellae always present. Cyphellae round to irregular, shallow to deeply excavate, margins smooth,240 THE LICHENOLOGIST Vol. 26 swollen and with a small pore (thelotremoid), or margins sharply defined, raised and with a wide pore, basal membrane white to brownish yellow. Internal cephalodia generally present, visible on upper surface as small hemispherical swellings and more obviously on the lower surface as often prominent, globose structures. Ascomata apothecioid, often rather sparsely developed, hemiangiocarpic, laminal and marginal, sessile or subpedicellate, margins entire or crenate, disc matt or shining, epruinose, proper exciple + well-developed, smooth or verrucose-scabrid, without photobiont cells. Theciwm I+ blue, hamathecium of simple, septate, filiform paraphyses, generally 2m thick, swollen (up to 55m) and sometimes pigmented at apices, pigment usually external to paraphyses. Epithectum pale or pigmented, straw-yellow to red-brown, minutely granular or a + clear gel external to tips of paraphyses. Hypothecium opaque, of densely interwoven hyphae, pale to dark yellow-brown ot red- brown. Asci shorter than paraphyses, cylindrical to clavate. Ascospores 8 per ascus, colourless or brown, fusiform-cllipsoid to broadly ellipsoid, apices rounded or pointed, 1-3 (rarely 5 or 7)-septate (see De Notaris 1851; Lindsay 1869). Conidiomata pycnidial, immersed, punctate to subglobose (Lobaria-type), as in Pseudocyphellaria (see Lindsay 1859, 1869; Vobis & Hawksworth 1981; Galloway 1988b, 19924). Condidiogenous cells colourless, + cuboid, phialidic, on simple to slightly branched conidiophores bearing conidia laterally and terminally, Conidia colourless, short, bracillariform to slightly lageniform, Key to South American species of Sticta 1 Photobiontgreen .. 2... Photobiont cyanobacterial . 2(1) ‘Thallus distinctly stalked, lobes thin, papery, rather fragile ‘Thallus not stalked, lobes thick, coriaceous... ss. 1. S.ainoae 3(2) Lower surface glabrous, yellow or whitish... . . 6. S. hypopsila Lower surface minutely pubescent-velvety, pale yellow-brown to red- brown... Lecce eee ee 7. §.lineariloba 42) Thallus distinctly stalked. . Thallus not stalked... . . 5(4) Isidiate Without isidia, lobe margins entire . 6(5) Isidia terete to irregularly coralloid, marginal and laminal, dark red-brown; lobes + plane, broadly rounded, thick, + coriaceous; lower surface pubescent-velvety; cyphellae not thelotremoid nn 5. S. hypochra Isidia granular, eroded-pseudosorediate, mat |, grey; lobes narrow, + canaliculate, thin, papery; lower surface glabrous, shining, costate; cyphellae thelotremoid, margins swollen and shining 9. S. Jongipes 7(4) Thallus with isidiaorsoredia .. . 2... ee ee Thallus without isidia or soredia Ss. gaudichaldia 8(7) Isidiate 9 Sorediate eee 2. S. caulescens1994 Southern South American Sticta—Galloway 241 9(8) Lower surface brown to dark brown, + densely and evenly velvery- tomentose; lobes not monophyllous; cyphellae small, 0-1-1'5(-3) mm diam ». 10 Lower surface whitish or pale; tomentum white, thinly scattered; lobes broadly rounded, + monophyllous; cyphellae large, 0-5— 10(9) Isidia + densely coralloid; lobes plane, rounded, margins dissected 11 Isidia minute, granular or styliform; lobes + inrolled at margins, elongate-laciniate, margins entire... 2.2... 10. S. santessonii 11(10) Isidia soon eroding and becoming sorediate. . . . 11. S. sublimbata Isidia never becoming sorediate . . . 1... 12. S.weigelii 12(8) Lobes + cochleate or monophyllous; soralia marginal and laminal, in + erose patches, soredia not derived from isidia . . . .8. S.limbata Lobes irregularly spreading, not monophyllous; soralia linear, eroding lower surface of margins, rarely laminal, soredia derived from minute, granular to + coralloidisidia.......... 11. S.sublimbata The Species 1, Sticta ainoae D. Galloway & J. Pickering Bibliotheca Lichenologica 38: 92 (1990); type: Chile, X Region, Valdivia, Las Palmas, University Forest Farm, 10km northeast of Valdivia, 73°10'W, 39°45'S, 10-100 m, on forest trees in plantation, 28 November 1986, B. 7. Coppins, D. J. Galloway, G. Guzman & P. W. James 8200 (SGO!—holotype; BM!, CONC!, HI, UPS!—isorypes). (Fig. 15) ‘This species is characterized by: the + dichotomously branched to some- what entangled lobes with a green photobiont; a mainly smooth upper surface, which is not faveolate or punctate-impressed; + prominent thelotremoid cyphellae on the lower surface with broad, smoothly rounded margins and a relatively narrow pore. Apothecia are occasionally developed and are marginal or laminal, 1-2-5 mm diam., shallowly concave or plane to subconvex, sessile, the disc shining, epruinose, pale orange-yellow to red-brown. The ascospores are broadly fusiform, colourless to pale brown, !~3-septate and 18-28 x 4-5— Tym. The morphology, anatomy and variation of this Valdivian rainforest species are described in detail in Galloway & Pickering (1990). In the earlier literature this species was incorrectly reportedas S. damaecornis (eg. in Crombie 1876; Nylander 1888; Malme 1899; Rasinen 1932; Zahlbruckner 1933; Grassi 1950; Lamb 1959; Follmann 1966; Redon 1972, 1974). Distribution: (Fig. 16). Between latitudes 38°S and 47°S, close to and mainly west of the Andean Cordillera. Endemic to southern South America. Habitat ecology: Sticta ainoae occurs on forest trees and shrubs in humid, rather shaded habitats in Valdivian rainforest where it is a conspicuous epiphyte amongst mosses and large species of Pseudocyphellaria such as P. berberina, P. flavicans and P. nitida in particular. In a recent collection from bark of Nothofagus in coastal forest at Laguna San Rafael in southern Chile (XIeemueretenretinerrecenneermeenieeron Fig. 17. Sticra caulescens (BM-D. J. Galloway 10439). Scale in mm,1994 Southern South American Sticta—Galloway 243 Region) a short-stalked form of S. ainoae was associated with clumps of fruticose Dendriscacauion {Galloway 9050 (BM)], similar to associations found in New Zealand with the green species of Sticta, S. filix and S. latifrons (James & Henssen 1976). ‘Specimens examined: 20 (see also Galloway & Pickering 1990). 2. Sticta caulescens De Not. Osserv. Sticta: 12 (1851).—Stictina caulescens (De Not.) Rasinen, Annis Bor. Soc. zo0l.-bor. fenn. Vanamo 2(1): 43 (1932); type: Chile, presso Valparaiso, dalla collezione del Rev. De Negri ‘@RO—holotype, not seen; UPS!—isotype). Sticta biatora De Not., Ossery. Sticta: 13 (1851); type: Chile, Valparaiso, legit Puccio ex Rev. De ‘Negri (RO— holotype, not seen [Santesson (UPS) notes that the holotype label has * Sticta biatora, E Chile a Rev De Negri. No 3 9/vi 847°}; UPS!—isorype) Stictina filicina f. latissima Nyl. in Crombie, J. Linn. Soc. Bot, 15: 229 (1876); type: Chile, ‘Tuesday Bay, 23 December 1868, R. O. Cunningham (BM!—holotype). Stictina caulescens . pallidoglauca Risiinen, Revta Univ. Santiago 22: 202 (1937); type: Chile, Panguipulli, 230m, 19 August 1928, R. P. A, Hollermayer (H!—holotype). Stictina caulescens f. brunnea Rasinen, Revia Univ. Santiago 22: 203 (1937); type: Chile, Mal Paro (Corral), sobre rocas maritimas, 23 March 1935, Fuga Gunckel (H!—lectotype selected here). (Fig. 17) Morphology: Thailus + palmate from a basal holdfast, stalked below, irregularly spreading above, 2-8(-12) cm diam., and 2-6(-10) cm tall, firmly attached by a stout, + woody, rooted holdfast 2~4(-6) mm wide, supporting a short, terete stalk, 0-5-1-5(-3) cm tall and 2-5mm thick. Lobes irregularly branching from a central stalk, + monophyllous and somewhat reniform to polyphyllous, narrow (0:5 cm) to broad 1-2-5(-3) cm, broadly rounded to + clongate-laciniate, apices notched, incised or + bluntly furcate, sometimes rather ragged and irregularly fenestrate. Margins entire, slightly thickened and with a definite ridge below, apices often with short, soft, silky projecting white hairs (x 10 lens). Upper surface malachite green to dark slate-blue to blue-black when moist, pale olivaceous or greyish often suffused red-brown when dry, smooth, matt, undulate, occasionally shallowly pitted or punctate-impressed in parts, thin and papery to rigid, + coriaceous, maculate, lacking isidia, phyllidia, or soredia. Maculae distinct, white, common, forming a delicate reticulum, or as isolated spots giving a marbled appearance to upper surface when moist (x10 lens). Medulla white. Photobiont cyanobacterial. Lower surface whitish to pale yellow-buff in marginal parts, to brown or grey-brown centrally and on stalk, irregularly wrinkled, pitted or subbullate to subfaveo- late, + striate near stalk, evenly and finely arachnoid-1omentose from margins to centre, tomentum short (rarely long and entangled), pale, silky, white to greyish. Cyphellae frequent, scattered, punctate, minute 0-1 mm diam., or less, margins very thin, sharply defined, not or only very slightly raised above tomentum, basal membrane white. Apothecia common, scattered, solitary, + sessile, constricted at base, with a marked concavity on lower surface at point of attachment, mostly towards lobe apices, marginal and laminal, (0-5-)1-3(—4) mm diam., round to irregular; disc, concave at first then plane to subconvex, shining or matt, epruinose, pale to244 THE LICHENOLOGIST Vol. 26 Sis coh an teva fol ateen, fp Cate Cork Phe Apa ued wee. Fic. 18. Sticta caulescens (BM-R. O. Cunningham; holotype of Stictina filicina f. latissima) Scale inmm. dark orange to red-brown; exciple persistent, pale pinkish or whitish to brown, scabrid-areolate to smooth, prominent and often obscuring disc in young fruits, becoming thin with age. Pycnidia occasional to common, rather inapparent, slightly swollen, apical pore shallowly punctiform, 0-2 mm diam., concolorous with thallus. Anatomy: Thallus 100-220(-280) wm thick. Upper cortex 22-32 um thick, colourless, cells + isodiametric, round to irregular, 3-9 tm diam., Medulla 35-160 pum thick, of colourless, loosely interwoven hyphae up to 3um thick. Photobiont layer 28-42(-55) um thick, rather irregular, photobiont Nostoc in clusters, cells up to 6 um diam. Lower cortex 14-22 um thick, outermost layer of cells pale straw-yellow, remainder colourless, cells + isodiametric, round to irregular, 3-9j1m diam. Tomental hairs rather sparse, simple, septate, colourless, up to 6 um diam., and up to 100 ym Jong. Apothecia: Exciple without photobiont cells, 90-135 um thick, colourless, of radiating parallel rows of round to irregular thick-walled cells 3-9 um diam.1994 Southern South American Sticta—Galloway 245 Hypothecium 30-45 um thick, opaque, pale straw-yellow, unchanged in K. Thecium 100-120 wm tall, colourless; epithecium 8-12 um thick, very dilute straw-yellow gel without granules, unchanged or slightly darkening in K, colour external to tips of paraphyses, Asci clavate 70-76 x 14-18 am. Ascospores broadly ellipsoid with pointed apices, colourless, 1-3-septate 28-36(-40) x 5-5-8:5 pm. Distinguishing features: Sticta caulescens is characterized by a distinctive palmate, caulescent thallus attached to the substratum by a firm, woody, rooted holdfast, the fronds arising from a short, terete stalk. It has broadly rounded lobes with entire margins, a cyanobacterial primary photobiont, and lacks cither isidia or soredia. It is distinguished from S. hypochra by its lack of isidia; from S. longipes, which has a narrower, longer stalk, narrower lobes with marginal isidia, and a glabrous lower surface with thelotremoid cyphellae; and from S. hypopsila and S. lineariloba, both of which have a green primary photobiont. Variation: Sticta caulescens shows considerable variation in the size and branching of lobes from + monophyllous to + polyphyllous and from narrow (0-5 cm) to broad (1-3cm), The taxon Stictina filicina f. latissima (Crombie 1876) is an example of the broad-lobed condition (Fig. 18). Distribution: (Fig. 19). Between latitudes 38°S and 54°S, mainly close to the Andean cordillera in areas of high rainfall. Endemic to southern South America. Habitat ecology : Sticta caulescens grows on the trunks and stems of rainforest trees and shrubs in very humid and often deeply shaded habitats where it is commonly found at or near the bases of large forest trees densely shaded by understorey shrubs, where low light levels effectively inhibit growth of many other lichens. It also grows on rocks close to rapids or waterfalls where + continual spray keeps thalli moist and able to photosynthesize effectively in low-light conditions. Together with S. Aypochra and Nephroma plumbeum (all cyanobacterial lichens) it appears able to grow successfully in some of the most densely shaded habitats known for lichens in southern South America. Ithas an altitudinal range from sea level to 1200 m. Selected specimens examined: Argentina: Prov, Rio Negro: Lago Frias, Perez Moreau 4548 (BA), Prov. Chubut: Lago Menendez, Kiihnemann 4762 (BA). Prov. Neuguen: Cotrentoso, A. Burkout 6427 (BA 1525). Staten Island: New Ycar’s Harbour, A. Menzies (BM)—Chile: IX Region: Parque Nacional Conguillio, 17. Quithor 834 (BM). X Region: Lago Rigihue, Enco, 1940, R. ‘Santesson 7688 S); S slope of Volcan Osorno, H. Kashitadani 17380, 17446 (TNS); Puerto Montt, 1940, R. Santesson 2278 (S); Chiloé Chepu, E. J. Godley 369 (BM, CHR). XV Region: Puerto Aysén, 1940, R. Santesson 4332, 4371 (S); Laguna San Rafael, D. 7. Galloway 9559 (BM). XII Region: Tierra del Fuego, Isla Clarence, J. Hyténen 2916 (H). (Total seen: 80]. 3, Sticta fuliginosa (Hoffm.) Ach, Meth. Lich: 280 (1803)—Lobaria fuliginosa Hoftm., Deutsch. Fl. 2: 109 (1796)—Parmelia sylvatica var. fuliginosa (Hofim.) Hepp. Lich. Fl. Witrzburg: 26 (1824)—Parmelia fuliginosa246 THE LICHENOLOGIST Vol. 26 Fig, 19. Distribution of Sticta caulescens in southern South America. (Hofim.) Schaerer, Lich. Helv. spic. sect. 10: 4383 (1840)—Sticta sylvatica var. fuliginosa (Hofim.) Hepp, Abbild. Beschr, Spor.: 371 (1875).—Stictina fuliginosa (Hoflm.) Nyl., Syn. meth. ich. 1(2): 347 (1860); type: Wales, Cader Idris, August 1726 (Dillenius 1742 Tab. XXVI, Fig. 100A—lectotype; OXF 100A—typotype: see Laundon 1984: 218-219). (Fig. 20) Morphology: Thallus + monophyllous, often deeply indented-incised and appearing polyphyllous, 2-12(-20) cm diam., rather loosely attached by a + central irregular umbilicus, easily broken. Lobes broadly rounded to some- what ragged-lacerate (0-5-)1-5(-8) cm wide, folded to + imbricate centrally. Margins broadly rounded, occasionally shallowly indented or incised, thinly ridged below and often rolled under, commonly densely isidiate. Upper surface livid slate-grey, dark glaucous-grey to blue-black, suffused red-brown in parts when moist, pale glaucous-grey suffused brownish when dry, shallowly to deeply wrinkled or subfaveolate, ridges sharply defined, matt to somewhat scabrid-areolate, rather papery and thin when well-developed, flabby when moist, brittle when dry, isidiate, maculate, rarely phyllidiate, soredia absent. Isidiaminute, styliform, brown-black, shining, granular-coralloid, very delicate and friable, in clusters at margins and on thalline ridges, often coalescing into a1994 Santhern South American Sticta—Galloway 247 Fic. 20. Sticta fuliginosa (BM-B. J. Coppins, D. J. Galloway, G. Guzman & P, W. James 8205). Ob.248 THE LICHENOLOGIST ‘Vol. 26 FIG, 21. Distribution of Sticta fuliginosa in southern South America. dark, sooty, diffract crust, rarely developing into small, flattened phyllidia or into small secondarily attached lobes or forming distinctly stalked lobate struc~ tures or entirely terete, richly branched Dendriscocaulon-like growths 2-5 mm tall, in habitats of continuous high humidity (Fig. 8). Maculae minute, white, delicately marbling upper surface, best seen when wet (x 10 lens). Medulla white. Phorobiont cyanobacterial. Lower surface pale creamish-white to buff, rarely brownish centrally, conspicuously wrinkled-ridged to + faveolate, densely tomentose from margins to centre, tomentum woolly, white, glisten~ ing, rarely with a broad to narrow, naked, shining white, marginal zone. Cyphellae common, flat, crateriform, 0-5-3(-5)mm diam., very large and prominent centrally, minute towards margins, basal membrane white, margins sharply defined, narrow, slightly projecting above tomentum. Apothecia very rare (seen only in Valdivian material), scattered, laminal on reticulate ridges, or marginal, + sessile, insertion of fruit forming a prominent cavity on lower surface, rounded, 1-3(—4) mm diam.; disc plane to subconvex, matt, epruinose, pale to dark red-brown; exciple very thin, pale or darker than disc, persistent, occasionally with minute, glistening white hairs.1994 Southern South American Sticta—Galloway 249 Anatomy: Thallus 135-225(-330) um thick. Upper cortex 14-40 um thick, outermost layer roughened-irregular separating into clusters, of pale yellow- brown to colourless, + isodiametric, round to irregular cells 3-8-5 ym diam. Medulla 55-180 um thick, colourless, hyphae loosely interwoven, up to 3-5 jm diam. Photobiont layer very irregular, 40-70 um thick, + continuous near cortext; thinly developed and scattered near medulla, photobiont Nostoc in clusters, yellow-green above, blue-green near medulla, cells 3-9 um diam. Lower cortex 28-42 um thick, colourless, cells + isodiametric, rounded 3-6 um. Tomental hairs simple, in clusters, colourless, septate, up to 6 um diam., and 50-170 uum long. Apothecia: Exciple 45-115 um thick, yellow-brown in outer parts, colourless internally, of thick-walled round to irregular cells, 5-5-17 jum diam., splitting off in ragged clumps, occasionally with small, septate hairs. Hypothecium 30-45 um thick, opaque, yellow-brown, unchanged in K. Thecium 80-112 um tall, colourless to pale straw-yellow; epithecium 11-20 )um thick, yellow- brown, not granular, paler in K. Asci (60-)70-82 x 14-18 um. Ascospores ellipsoid, apices pointed, colourless, 1-3-septate, contents granular-vacuolate, (25-)28~31(-33) x 5-5-8'5 um. Distinguishing features: Sticta fuliginosa has a cyanobacterial photobiont, broadly rounded, + monophyllous lobes, delicate, granular coralloid isidia on margins and upper surface of lobes, and a pale creamish white of buff-pink lower surface with pale tomentum and large flat, conspicuous cyphellae. It is distinguished from S. santessonii and S. weigelii, both of which have dark- brown lower surfaces and different lobe morphology, and from S. gaudichaldia, which lacks isidia and has a different morphology and ecology. Sticra limbata has a somewhat similar thallus morphology but the upper surface is smooth and at times + shining, and it has conspicuous erose-soralia on margins and upper surface and sometimes also on the lower surface including the cyphellae. Variation: Sticta fuliginosa varies in thallus size according to habitat conditions, and also in the degree of development of apothecia, fertile speci- mens being the exception rather than the rule, although large, fertile specimens are common in favourable sites for lichen growth such as in humid rainforest at Choshuenco (Hildebrand-Vogel et al. 1990), an exceptional area for lichen biomass and biodiversity; large, fertile specimens are common. Over much of its range worldwide, S. fuliginosa tends to produce only sterile thalli. Fertile thalli from British and Macaronesian populations are much smaller on average than fertile specimens from southern South America (P. W. James pers. comm.). South American specimens are similar in all respects to material seen from other parts of the world, although in habitats of high humidity and especially in the dense shrub thickets near Rio Enco, specimens exhibit wide variation in the development of isidia and the formation of Dendriscocaulon-like struc- tures (Fig. 8), which are not known from any other populations of the species elsewhere. Distribution: (Fig. 21). From latitude 30°S (Fray Jorge National Park) to 55°S in southern South America. Cosmopolitan in temperate regions.250 THE LICHENOLOGIST Vol. 26 < halia DL. Suelo Gad Fic. 22. Sticta gaudichaldia (PC-LENORMAND-holotype). Scale in mm. Habitat ecology: Sticta fuliginosa is an epiphyte of bark and twigs of trees and shrubs and on rotten stumps in forest and scrub communities and oc cagionally also colonizes rocks in rather drier habitats. It occurs in humid scrub communities, along roadsides in Valdivian rainforest and in alpine scrub, where it is most commonly collected. It is known from the following phorophytes: Aextoxicon punctatum, Baccharis spp., Berberis buxifolia, Drimys winteri, Fuchsia magellanica, Gevuina avellana, Nothofagus dombeyi, N. obligua, N. pwnitio and Rhaphithamnus cyanocarpus. Sticta fuliginosa associates with the following lichens: Degelia gayana, Dictyonema glabratum, Erioderma leylandii, Hypotrachyna sinuosa, Leioderma pycnophorum, Nephroma antarcticum, Normandina pulchella, Pseudocyphellaria berberina, P. coerulescens, P. coriifolia, P. crocata, P. kirsuza, P. matiota, P. piloselloides, P. pluvialis, P. scabrosa, Poroma leprotomum and P. sphinctrinum. Ut bas an altitudinal range from sea Jevel to 2200 m. Selected specimens: Argentina: Prov. de Salta: Depto Candelaria, Puesta la Queira, S. Venturi (BA 3784). Staten Island: New Year's Harbour, A. Mensies (LINN-SM 1705.19, US).—Chile: IP Region: Parque Nacional Fray Jorge, H. Schindler & J. Redon 3612, 5616 (KR). IX Region: Parque Nacional Conguillio, W. Quilhor 832 (BM). X Region: Lago Rifiihue, Enco, R. Santesson 3885 (Ex Herb. Regnell, Exs. No 433] (BM); Chiloé, R. 0. Cunningham (BM). XI Region: Laguna San Rafacl, D. 7. Galloway 9306 (BM). X7I Region: Lago del Toro (L. Maravilla), Estancia Rio1994 Southern South American Sticta—Galloway 251 Fig. 23. Distribution of Sticta gaxdichaldia in southern South America. Payne, R. Santesson 6389 (S); Punta Arenas, R. O. Cunningham (BM); Canal Whiteside, Puerto Yartou, R. Santesson 6806 (S); Lago Roca, R. Santesson 1077 (S). {Total seen: 80]. 4. Sticta gaudichaldia Delise Men, Soc. tinn. Normandie 2:80, Tab. VII. fig. 23(1825).—Sticta gaudichaldiana (sic.] Persoon in Gaudichaud, Voy. Uranie Bot.: 200 (1827)—Sticta filicina var. gaudichaudii (Delise) Mont, Annis Sci. nat. Bor. séx. 3, 18: 308 (1852) —Sticta freycinerié var. gaudichaudii (Delisc) Lindsay, Trans. r. Soe, Edin. 22: 197 (1859).—Stictina gaudichaudit (Delise) Nyl., Syn. mech, lich. U2): 345 (1860).— Stictina quercizans var. gaudichaudil (Delise) Nyl., Flora, Jena 48: 297 (1865); type: [Falkland Islands} Iles Malouines. Mr Gaudichaud Naturaliste de "éxpedition Freycinet, 1820. Regu le ler Janvier 19822 (PC-LENORMAND!—holotype). Sticta malovina Fr., Syst, Orb, Veg, 1:283 (1825); type: Iles Malovines. Gaudich.n. 117 (UPS!— holotype; G!—isotype). ‘Sticta negeri Vainio, Hedwigia 38: 187 (1899}—Stictina negeri (Vainio) Datbishire, Wiss. Erecbn, Schwed. Sizdpolarexped, 1901-1903 4(11): 55 (1912)s type: [Chile] Inter muscos rupium prope Port Tamar in Freto Magellanico, Neger 304 (TUR 10546!—holotype). (Fig. 22) Morphology: Thallus irregularly spreading, 2-8(-15)cm diam., loosely attached at base, margins and apices + ascending, straggling through low252 THE LICHENOLOGIST Vol. 26 vegetation or at base of small shrubs in grassland. Lobes + elongate, + ascend- ing to horizontally straggling, undivided, subcanaliculate to plane, apices shallowly notched or incised, 1-3 cm wide. Margins entire, distinctly swollen, thickened and + darkened above, slightly ridged below sinuous and + inrolled when dry. Upper surface livid slate blue-grey to olivaceous-glaucous or dark blue-black, suffused reddish or brownish when moist, pale fawnish grey to red-brown or dark brown when dry; shallowly undulate, wrinkled in parts or obscurely ridged to irregularly and shallowly pitted, matt to glossy; rather thin and papery, brittle when dry; conspicuously maculate, lacking isidia, phyllidia, or soredia. Maculae common, white to pale buff or red-brown, in delicate, minutely reticulate patterns giving upper surface a marbled appearance when moist (x10 lens). Photobiont cyanobacterial. Lower surface wrinkled to indistinctly shallowly ridged to + subundulate and smooth, pale buff at margins darkening centrally to + uniformly dark brown to black, + glabrous in a narrow marginal zone to entirely regularly short-tomentose, tomentum velvety, even, pale whitish to buff to dark brown or black. Cyphellae rather widely scattered, more common at margins, rounded to subirregular 0-1-0-3(-0-8) mm diam., deeply excavate, basal membrane white, minutely granular; margins prominent even when young, swollen, raised, shining, projecting above tomentum. Apothecia rare to occasional, commonly absent, laminal, rarely submarginal, solitary, (0-5-)0-8-2(-3) mm diam., rounded to subirregular, margins thin, regular, smooth and shining, entire to minutely crenulate, dark brownish to + concolorous with thallus, commonly excluded by disc in mature fruits; disc plane to distinctly convex, pale orange-brown to red-brown, markedly paler than exciple, matt, epruinose; subpedicellate, pedicel short, less than half width of disc. Pycnidia scattered, frequent to occasional, marginal and laminal, up to 0-5mm diam., swollen, raised-papillate, ostiole red-brown when wet, dark brown to black when dry. Anatomy : Thallus 165-280 wm thick. Upper cortex 20-28 um thick, outermost 12-14 ym pale yellow-brown to red-brown, remainder colourless; cells + iso~ diametric, 8-10 um diam. Medulla 75-145 ym thick, of colourless, densely interwoven to somewhat loosely textured hyphae, up to 3 ym diam, Photobiont layer 55-90 um thick, rather irregular, photobiont Nostoc in clusters, cells 6-9 um diam. Lower cortex 14-20 um thick, pale to dark red-brown, cells + isodiametric, round to irregular, 6-8 um diam. Tomental hairs simple, septate, clustered, colourless, up to 6 jim diam. and up to 90 um long. Apothecia: Exciple 110-280 wm thick, colourless to pale straw-yellow, of radiating, + isodiametric, round to irregular cells, 512 um diam. Hypothecitun 55-90 um thick, opaque, pale yellow-brown, unchanged in K. Thecium 85~ 110 um tall, colourless to pale straw-yellow; epithecium 8-15 ym thick, pale yellow-brown, paler in K. Asci 80-85 x 14-18 um. Ascospores ellipsoid, apices pointed, colourless, 1-3-septate, 33-42 x 6-9 um. Distinguishing features: Sticta gaudichaldia is characterized by a cyanobacterial photobiont, broadly rounded to irregular, rather papery lobes (brittle when1994 Southern South American Sticta—Galloway 253 dry) with entire margins, and without isidia or soredia. It is similar to S. santessonii but lacks the isidia of that species. Both species have similar ecologies. Variation: Sticta gaudichaldia shows rather little morphological variation and as a rule thalli are sterile. It is still rather poorly collected. Distribution: (Fig. 23). From Valdivia (Puyehue National Park) southwards to Tierra del Fuego, Staten Island and the Falkland Islands, Most common in ‘Tierra del Fuego. Endemic to southern South America. Habitat ecology: Sticta gaudichaldia is a terricolous Fuegian species best developed in Magellanic tundra (see Pisano 1983; Tretiach et a/. 1989) in Patagonia, Tierra del Fuego and the Falkland Islands, and above the treeline (rarely) in Valdivia. On the shores of Laguna San Rafael (latitude 46°S) in damp sand colonized by introduced grasses, S. gaudichaldia grows vigorously together with Leptogium britanmicum, L. menziesii, Peltigera membranacea and P. scabrosa, forming very large colonies and undoubtedly contributing appreciable amounts of fixed nitrogen from their cyanobacterial photobionts. The growth of L. menziesii in this habitat is extremely rapid with the lichen actively overgrowing annual grass stems (Galloway 1992a). This habitat is noteworthy for the occurrence of the bipolar taxa L. britannicum and P. scabrosa, the former not recorded until now from outside Great Britain and a few sites in western Europe (Jorgensen & James 1983; Purvis et al, 1992). Specimens exantined: Argentina: Tierra de! Fuego. Sierra Sorondo above Las Cotorras, 20 km ENE of Ushuaia, R. Santesson 626 (S); Isla Grande, Peninsula Mitre, Bahia Valentin, S. Stenroas 2784 (A); Staten Island: New Year's Harbour, A. Mensies (BM, E, LINN-SM 1705.13 pr.p}.— Chile: X Region: Parque Nacional Puyehue, Antillanca, B. J. Coppins, D. J. Galloway, G. Guzman & P. W. James 8194, 8197, 8201 (BM); § slope Volca Osorno, H. Kashitvadani 17481 (TNS). XI Region: Laguna San Rafael, D. 7. Galloway 9118, 9119, 9168 (BM). XIT Region: Punta Arenas, W. Lechler 1348 (BM); Sholl Bay, Hassler Expedition 1872(BM); Tierra del Fuego, Isla Clarence, HH. Roivainen 1844 (H); Isla Clarence, Southern Peninsula, J. Hyodner 2849 (H); Ibid., S. Stenroos 2586 (H); Hermite Island, J. D. Hooker (1. 910 210-65)—East Falklands: Berkeley Sound, Port Louis, 1. M. Lamb 2645 (BM); Port Stanley, slopes of Mr William, 7. M. Lamb 1008, 2920(BM). 5, Sticta hypochra Vainio Res. Voy. S. ¥. Belgica Bot.: 29, Pl. IIT, figs 24-26 (1903); type: Argentina, Staten Island, Golf St Jean, G. Racovitza 185 pr.p (BR!—lectorype selected here). Stictina fuliginasa f. firmior Crombie, 7. Linn. Soc. Bot. 15: 229 (1876); type: Chile. Sandy Point (Punta Arenas), 16 March 1867, R. O. Cunninghan (BM!—lectotype selected here; E!—isotype).. Stictina quercizans var. glaucovirens Jatta, Malpighia 20: 6 (1906); type: Chile. Temuco, ad frunegs muscosos in locis umbriosis forestae antarcticae, G. 1, Scou-Ellior 165 (E!—holotype [as “1857 Stictina filicinella ver. ochraceofusca Risinen, An. Soc. cient, argent. 128: 143 (1939); type: Azgentina, Parque Nacional Nahuel Hwapi, 800 m, Donat (?H—not seen). [Sticea hypochra was also recorded erroneously under the following names: Stictina filicina var. ‘marginifera sensu Ny1. (1888); S. marginifera vat. coralloides sensu Mull. Arg.; S. orbicularis sensu Miill. Arg; S. filicina sensu Malme]. (Fig. 24)254 THE LICHENOLOGIST Vol. 26 sAMOULEUUENEEEEYVOBEGHeSUUOHeEyerasasseget. Fig. 24, Sticta hypochra (BM-D. J. Galloway 9060). Scale in mm. Fi. 25. Distribution of Sticta hypochra in southern South America1994 Southern South American Sticta—Galloway 255 Morphology: Thallus + palmate from a basal holdfast, 2-6(-8) cm diam., and 1-4(6) cm tall, firmly attached by a rooted, short, terete stalk 2-5 mm wide, 6-10(-20) mm tall. Lobes arising from basal stalk, polyphyllous, rarely monophyllous, 5-15(~40)mm wide, broadly rounded, + seniform, rarely laciniate-lacerate. Margins irregular, notched or incised or ragged, at maturity down-rolled and with a conspicuous ridge below, regularly and densely isidiate. Upper surface livid slate-blue, blue-black. Rarcly malachite-green, often suffused red-brown to dark-brown when moist, olivaceous-glaucous to red-brown to brown-black when dry, plane, undulate to shallowly punctate- impressed, smooth, matt, thin and papery to thick and coriaceous, pliable when wet, rather rigid and brittle when dry, isidiate, maculate, soredia absent. Isidia common, crowded-congested along margins or in scattered, laminal clumps, or ina + continuous diffract crust, terete, styliform at first becoming + coralloid- complex, 0-1-0-5 mm tall, often densely clustered, sometimes becoming + Dendriscocaulon-like, or forming smal] secondary lobules. Maculae minute, white, reticulate or as scattered spots (x 10 lens). Medulla white. Photobiont cyanobacterial. Lower surface whitish to pale yellow-buff to chestnut brown or red-brown, irregularly wrinkled, + bullate to subfaveolate in parts, evenly velvety tomentose from margins to centre, tomentum short, even, whitish to buff or brown. Cyphellae scattered, prominent, (0-1-)0-5-1-5(-2) mm wide, crateriform, margins sharply defined, not or only slightly raised above tomentum. Apothecia rather rare, often absent, scattered towards lobe margins, rounded 0-2-2(-4) mm diam., sessile to subpedicellate, insertion of pedicel visible as a distinct cavity on the lower surface; disc plane to subconvex, waxy, epruinose, yellow-orange to orange-brown; exciple prominent, persistent, smooth to crenate or + scabrid, paler than disc, + translucent when wet. Anatomy: Thallus 140-225(-300) ym thick. Upper cortex 22-28 um thick, pale straw-yellow, cells thick-walled + isodiametric, round to irregular up to 94m diam. Medulla 70-190 um thick, densely interwoven, colourless to pale straw-yellow, hyphae up to 3:5 jum diam. Photobiont layer 25-55 yum thick, photobiont ? Nostoc, in clusters, cells up to 6 um diam. Lower cortex 22-28 um thick, outermost zone dark yellow-brown, internally pale straw-yellow, cells round to irregular, 2-5-8 im diam. Tomental hairs pale yellow-brown, simple, septate, up to 8-5 um diam., and up to 80 um long. Apothecia: Exciple 55-170 um thick, colourless, of radiating, parallel, thick-walled, round to irregular cells, 6-22 um diam. Hypovhecium 90-110 um thick, pale yellow-brown, unchanged in K. Thecium 110-130 um tall, colour less; epithecium 8-14um thick, pale red-brown, unchanged in K. Asci 80-90 x 14-18 um. Ascospores colourless, ellipsoid, apices pointed, 1-3-septate, 30-40 x 5-5-8:5 pm. Distinguishing features: Sticta hypochra is a distinctive, caulescent species attached to the substatum by a broadly rooted holdfast. It has a cyanobacterial photobiont, + polyphyllous rather ragged lobes arising from a short support- ing stalk, marginal and laminal terete to coralloid isidia that rarely become Dendriscocaulon-like (Fig. 9), a velvety buff tomentum on the lower surface,256 THE LICHENOLOGIST Vol. 26 and crateriform cyphellae, It is distinct from S. longipes, which has a markedly longer and narrower stalk, narrower, + strap-like lobes, minutely granular marginal isidia, a glabrous lower surface and pin-prick-like, thelowremoid cyphellae. Sticta hypochra is the isidiate counterpart of S. caulescens. Variation: Sticta hypochra shows variation in both size and degree of robustness of lobes and also in number and position of isidia. It is most commonly collected without apothecia, but in habitats of + continuous high humidity it will produce apothecia in well-developed specimens. In sites of continuous high humidity, such as near Rio Enco in Valdivia, specimens of S. hypochra (together with S, fuliginosa and S. longipes) show a distinctive development of isidia into flattened, stalked lobules and Dendriscocaulon-like structures, a development not commonly seen elsewhere in the range of this species. Distribution: (Fig. 25). From latitude 38°S to 55°S. Endemic to southern South America. Habitat ecology: Sticta hypochra occurs as an epiphyte of trees and understorey shrubs in rainforest and will also grow among moss on rotten tree trunks, and twigs of shrubs, on stones and rock sand soil close to streams and watercourses, and on the ground in relatively dry, sparsely vegetated, deforested rocky areas. As with S. catélescens itis capable of colonizing substrata in very low-light situations providing humidity is high. Ir has an altitudinal range from sea level to 1200 m. Selected specimens: Argentina: Prov. Neuguen: Brazo Blest, Arroyo Bravo, V. Kull 2627, 2628 (Ba). Prov. Chubiat: Lago Menendez, 4. Castellanos 2474 (UPS). Lago Argentino, Fiotdo Mayo, PAV. James (BM). Tierra Del Fuego. Isla Grande, Bahia Brown, S. Stenroos 2378 (H); Ushuaia, R. Santesson 439 (Sy; Canal Beagle, Lapataia, R. Santesson 1143 (S.—Chile: LX Region: Parque Nacional Conguillio, G. Guzman 957 (BM). X Region: Valdivia, Krause (BM); Lago Rinihue, Cerro Tralcan, R. Santesson 3483S); Enco, 2. Santesson 7605 (S); Chaiten, W”. Quithot 2281 (BM). XI Region: Puerto Aysen, R. Santesson 4202 (8); Laguna San Rafael, D. J. Galloway 9754 (BM). XII Region: Rio Rubens, c. 50 km SE of Natales, R. Santesson 5667 (8); Canal Beagle, Yendegaia, R. Santesson 1506 (S); Ista Riesco, Mina Elena, R. Saniesson 2043 (BM, UPS); Isla Hoste, Peninsula Dumas, Canasaca, 29 February 1940, R. Santesson 7414 (S). [Total seen: 125]. 6. Sticta hypopsila (Mont.) Crombie J. Linn, Soc. Bot. 16: 223 (1877).—Sticta filicina var. lineariloba f. hypopsila Mont., Annls Sci. naz. Bot. sér. 3, 18: 308 (1852).—Sticta damaecornis var. hypopsila (Mont,) Nyl., Aunis Sei.nat, Bot. ser. 4, 11: 213 (1859).—Sticta lineariloba var. hypopsita (Mont.) Nyl.s Sym, meth. lich. (2): 355 (1860)5 type: Juan Fernandez, sine collectoribus lecta in litt. ad. el. J. D. Hooker (BM!—lectotype selected here; PC-MONTAGNE!— isotype). [BM material chosen as lectotype is labelled by Montagne Sticta hypopsila Montag, Fl. Chil. MS. Je vous ai promis un bel échantillon, le voila. Montagne] Sticra laciniata var. denudata sensu Zahir. (1924) non Nyl. (1860). (Fig. 26) Morphology: Thallus irregularly spreading from a basal holdfast, (2-)5— 12(-20) cm diam., firmly attached by a short, stout, terete holdfast 2-3 mm