Distribution and habitat ecology of the sorediate species of Menegazzia ( Parmeliaceae , lichenized Ascomycota ) in Chile Distribución y ecología de las especies sorediosas de Menegazzia ( Parmeliaceae , Ascomycota liquenizado ) en Chile

The taxonomy and ecology of the sorediate species of Menegazzia from the southernmost regions of Chile and Argentina and the South Atlantic Islands was recently published, only with sporadic reports from the more northern regions. In the present work the distribution patterns and habitat ecology of the sorediate species are discussed, with emphasis on the area north of 48o S. Eleven species are treated. Menegazzia subpertusa , n epiphyte of sclerophyll scrubs, is recorded from South America for the first time (Chile and Argentina). Menegazzia neozelandica h s a disjunct distribution in Chile, with occurrences in Fray Jorge (Fourth Region of Chile) and on Islas Juan Fernández, and along the coast south of latitude 38o S. Menegazzia kawesqarica and M. tenuis are most common in the southernmost part of Chile, but are also found at high altitudes at lower latitudes. Additional treated species are M. chrysogaster , M. fumarprotocetrarica, M. globulifera, M. magellanica, M. norsorediata, M. sanguinascens and M. wandae. Several of the sorediate species are early colonisers of newly developed substrates. They show variable occurrences along light and humidity gradients. Distribution maps and a revised key are presented.


INTRODUCTION
The genus Menegazzia A. Massal.constitutes a prominent part of the lichen mycobiota in the forests of southern South America (Santesson 1942).The checklist of Chilean lichens lists 14 species (Galloway & Quilhot 1999).Subsequently, the sorediate species from the southernmost regions of Chile and Argentina were revised by Bjerke & Elvebakk (2001), including descriptions of two new species.An additional new species known between latitudes 38º30' and 46º40' S Distribution and habitat ecology of the sorediate species of Menegazzia (Parmeliaceae, lichenized Ascomycota) in Chile Menegazzia were obtained, and the new results are presented here.Special emphasis is given to the distribution of species in the regions not treated by Bjerke & Elvebakk (2001), viz.all regions north of the Magellanic region (Duodécima Región de Magallanes y de la Antártica Chilena).Some reports on the sorediate species of Menegazzia from regions Four to Eleven exist (e.g., Räsänen 1932, Redon et al. 1975, Quilhot et al. 1975, and literature reports cited in detail in results and discussion).These reports are revised here.Distribution maps and a revised key to the sorediate species are presented.

MATERIAL AND METHODS
Field studies and collecting trips were performed in all regions of Chile, including Islas Juan Fernández.No material of Menegazzia was found north of latitude 30º S. Laboratory studies were mainly done on material collected by the present authors, especially on the numerous specimens collected by the third author during the last 30 years and deposited in the Herbarium at Universidad de Valparaíso (UV).Most specimens collected by the first and second authors are housed in TROM.Additional material, including specimens of extra-American species for comparisons and many type specimens, were seen in or obtained from the following herbaria: B, BG, BM, GZU, H, HIP, HO, MSC, O, S and UPS.Specimens examined by Bjerke & Elvebakk (2001) are included on the maps here, and in the total number of examined specimens, but not referred to in any more detail.For Quilhot's specimens, the collection numbers are equivalent to the herbarium's accession numbers.The brief descriptions of all treated species are based on descriptions and illustrations in Bjerke (2001) and Bjerke & Elvebakk (2001), and new data.Other sources for morphological traits are cited when available.Thin-layer chromatography of acetone extracts was performed using standardised procedures (Culberson 1972, White & James 1985).Only commonly-occurring secondary substances are mentioned below.For less common substances, see Bjerke & Elvebakk (2001).Nomenclature follows Galloway & Quilhot (1999) for lichens, except for Coccotrema coccophorum (Schmitt et al. 2001), and Marticorena The distribution of the sorediate species of Menegazzia is related to the bioclimatic zones and subzones in order to understand more about the species' ecological requirements.
Menegazzia chrysogaster is a South American endemic species that has convex white or yellowish soralia, wide, slightly convex and partly glossy lobes, and a characteristic ochre-yellow upper side of the internal cavity.The upper surface is greyish-green with small maculae, and often with slightly pruinose lobe tips.The margins of the lobes become black, especially towards the centre.
Variation: Menegazzia chrysogaster has a more variable morphology than first thought, thalli from more light-exposed microhabitats tend to be more greyish than the more common pale green thalli found in shadier habitats.The crater-like depressions found in one specimen from Argentina (Bjerke & Elvebakk 2001) were not seen in specimens from Chile.The degree of maculation, pruinosity and blackening of margins is also variable.Lobes vary from contiguous towards apices and imbricate to contiguous with open spaces between lobes.Two collections from Isla Desolación made by P. Dusén in 1896 and published as M. sanguinascens by Santesson (1942) were discussed in context with M. chrysogaster by Bjerke & Elvebakk (2001).More detailed morphological and chemical studies revealed that these collections include thalli of M. chrysogaster, M. norsorediata and M. sanguinascens.
Distribution: In Chile from latitude 36º50' S (Termas de Chillán, VIII Region of Chile) to 53º22' S (Península Brunswick, XII Region of Chile) (Fig. 1).It is also known from Isla de los Estados in southern Argentina (Bjerke & Elvebakk 2001).Most known localities are from forests surrounding the Andean cordillera, but a few localities closer to the Pacific Ocean are known, e.g., along Cordillera de La Costa.
Habitat ecology: Menegazzia chrysogaster is an epiphyte of trees and shrubs in low-to medium-light situations in  Adler, Mycotaxon 59: 369 (1996).
Menegazzia fumarprotocetrarica is a South American endemic species that has soralia formed on 1-2 mm tall pustules, a greenish, glossy upper surface, and small perforations with flat or slightly elevated rims.Its major medullary constituent is fumarprotocetraric acid, which makes it chemically very distinct from other South American representatives.It is discussed in detail and illustrated in Adler & Calvelo (1996) and in Bjerke & Elvebakk (2001).
Variation: Menegazzia fumarprotocetrarica shows some variation in thallus size and morphology of soralia.The widest lobes are found in the specimens from the south (to 2.0 mm wide), whereas lobes from shadier habitats in the north are not wider than 1.1 mm wide.The southern populations are also duller and slightly more pruinose than in the north.The form and size of the sorediate pustules appears to depend upon the age of the soralia.In old pustules, few soredia are remaining, and an open duct to the cavity is evident, whereas younger pustules have welldefined convex, richly sorediate apices.
Menegazzia globulifera is a pan-austral species having vesicular, helmet-shaped soralia, a yellow-green shining upper surface and usnic acid in the upper cortex.It is discussed in detail in Santesson (1942), Galloway (1985) and James & Galloway (1992).
Variation: Menegazzia globulifera has a rather constant morphology apart from shape of soralia and thallus colour.Older soralia tend to erode, with only the basal part of the vesicle remaining, whereas young, immature soralia are more or less convex.In shaded habitats, M. globulifera partly looses its yellow-green colour.Only trace amounts of usnic acid are detected in specimens from these habitats.
Habitat ecology: Menegazzia globulifera is an epiphyte of trees and shrubs in light-exposed, relatively dry vegetation.It is occasionally also saxicolous.It is known from the following phorophytes: Nothofagus pumilio, N. dombeyi, N. betuloides, N. antarctica, N. alpina, Araucaria araucana, Chiliotrichum diffusum, Berberis linearifolia, B. empetrifolia, B. ilicifolia and Embothrium coccineum.In regions Eight to Ten, it is a high-altitude species with altitudinal range from ca. 500 to 1,800 m, only with scattered occurrences at lower altitudes.It is probably more Menegazzia kawesqarica is a southern South American endemic species characterised by a greyish, patchily blackened and corrugate upper surface, by large, convex, partly confluent soralia, and by wide, convex, prominently maculate lobes.
Variation: the northern populations of M. kawesqarica differ slightly from the southern populations by having less corrugate and less pruinose lobes, and by having soralia that are not so confluent.Some specimens of M. kawesqarica produce apothecia without producing mature asci and ascospores (Bjerke & Elvebakk 2001).One large apothecium was found in a specimen from the Eleventh Region of Chile (W.Quilhot 3005), but asci were not found.
Distribution: in Chile from latitude 37º49' S (Nahuelbuta, Eleventh Region of Chile) to 53º22' S (Península Brunswick, Twelfth Region of Chile) (Fig. 4).It is still not recorded from Argentina.These are the first records from Chile north of latitude 47º S.
Habitat ecology: in the Ninth Region of Chile it is an epiphyte of large trees in high-altitude Menegazzia magellanica is endemic to southernmost South America.It has a greyish-green, smooth upper surface, convex, slightly pruinose and maculate lobes, and maniciform soralia with a n o p e n d u c t t o t h e c a v i t y .M e n e g a z z i a magellanica occasionally produces apothecia with two-spored asci.Bjerke & Elvebakk (2001, pp. 382-383)  gions Eight to Eleven identified as M. magellanica were seen.These were referable to M. wandae and M. neozelandica, in particular.For distribution map, morphological variation, habitat ecology and list of examined specimens, see Bjerke & Elvebakk (2001).It is, however, slightly less variable than stated by Bjerke & Elvebakk (2001), because some problematic specimens assumed to belong to M. magellanica are here redetermined to M. neozelandica after comparison with more specimens of the latter species (see below).
Variation: Menegazzia neozelandica has a rather constant morphology apart from the soralia, which occasionally are not associated with perforations, especially in small thalli.The soralia that do not surround perforations are subcapitate, and resemble the soralia of several other species.In particular, specimens from near the limits of its Chilean distribution range and specimens encircling tiny twigs deviate by having many soralia not associated with perforations.A specimen (A.Elvebakk 98: 255) identified as M. magellanica by Bjerke & Elvebakk (2001) is here redetermined to M. neozelandica because of elevated rims, a shining upper surface, and a few soralia surrounding perforations, which were previously considered true maniciform soralia.
Distribution: It has a disjunct distribution in Chile (Fig. 5).It is known from latitude 30º30' S (Fray Jorge, Fourth Region).It is so far not known from the area between Fray Jorge and Lago Lleulleu (Eighth Region).It has a more or less continuous distribution from latitude 38º20' S to latitude 52º40' S (Isla Riesco, Twelfth Region).It is also present on Islas Juan Fernández.It is much more common in Chile than previously expected (see Bjerke & Elvebakk 2001).These are the first records from Islas Juan Fernández and from the mainland north of latitude 45º30' S, and it is the only species of Menegazzia known from the marginal and relict forests of Fray Jorge and from the isolated archipelago of Juan Fernández.It is also known from a few localities in Argentina (Calvelo & Adler 1994), from Tasmania (James & Galloway 1992), and from New Zealand (Galloway 1985).
Habitat ecology: Menegazzia neozelandica is an epiphyte of trees and shrubs in Valdivian rainforests in moderate to dense shade, often growing intermixed with mosses, Hymenophyllum ferns, and other lichens.When growing on trunks of tall trees it is often more attached to other epiphytes than directly to the bark.Its occurrence on erect Hymenophyllum shoots is very striking, and it is a first-successional species, producing numerous small thalli; and locally as the most common pioneer species in such habitats.When attached directly to bark, it prefers branches of smooth-barked species such as Amomyrtus luma, A. meli, Laurelia phillipiana, Aextoxicon punctatum, Tepualia stipularis, Gaultheria mucronata, Embothrium coccineum, Drimys winteri, Berberis spp., Kageneckia oblonga (Fray Jorge), Baccharis sp., Ovidia pillo-pillo and Rhaphithamnus spinosus.It is also found on Nothofagus, e.g., N. nitida, N. dombeyi, N. betuloides, N. antarctica and N. alpina, and on planted Populus along roadsides on Chiloé.One single specimen was found on a rock at the forest floor.It associates particularly with shade-loving lichens with cyanobacterial photobionts, such as Degelia gayana, Leioderma pycnophorum, Pseudocyphellaria bartlettii, P. intricata, Menegazzia norsorediata Adler & Calvelo, Mycotaxon 59: 368 (1996).
Type: Argentina, Province of Tierra del Fuego, 10 km from the intersection of Moat River and the route from Moat to Ushuaia, November 1993, S. Calvelo & M. Adler (BAFC 37732,holotype).
Menegazzia norsorediata is a southern South American endemic species.It has greyish-green shining lobes that are generally epruinose and slightly maculate, evenly distributed perforations with flat or elevated rims, and convex, wellseparated soralia with orange patches.It is chemically distinct from other South American species.
Variation: Menegazzia norsorediata shows some minor morphological variation.Some thalli produce several sympodial, toe-like branches, but generally, these are absent.Large thalli growing on trunks often have radiating lobes with perforations lined on parallel lobes, whereas on twigs, the lobes are more complexly divided.The size and colour of soralia vary with age and habitat.Young soralia are small and with a greenish hue in fresh conditions.Apothecia are rare.
Habitat ecology: Menegazzia norsorediata is a common epiphyte of trees and shrubs in both Valdivian rainforest and high-altitude mixed deciduous-coniferous forest, in microhabitats with variable light conditions.In Petrohué (Tenth Region), it is common both on open twigs of isolated trees of Nothofagus pumilio and N. dombeyi at the forest edge, and on myrtaceous shrubs in the understorey vegetation of well-developed N. dombeyi forests permitting little direct solar radiation to reach the lower strata.It is among the first macrolichens to colonise the bark of young shrubs and trees.Other known phorophytes are N. alpina, N. nitida, N. betuloides, N. antarctica, Podocarpus alpinus, planted Pinus, Drimys winteri, Embothrium coccineum, Euchryphia cordifolia, Gevuina avellana, Weinmannia trichosperma, Berberis ilicifolia and Maytenus magellanica.It has a wide altitudinal range from sea level on Chiloé and in Laguna San Rafael to 1,400 m, and probably higher in and around Conguillío.Since it has a wide ecological amplitude, it associates with many different species.However, it prefers smooth bark, and is only rarely found in close association with broad-lobed species of Sticta and Pseudocyphellaria that prefer more roughened bark.
Parmelia sanguinascens Räsänen, Annales Botanici Societatis Zoologica Botanica Fennicae Vanamo 2, 1: 18 ( 1932 This characteristic austral species has broad, shining and emaculate lobes, few to numerous toe-like lateral branches, and neat, convex soralia that are UV+ white.It is often prominently green when wet and fresh, and has small, gaping perforations with slightly elevated margins.It is discussed in detail and illustrated in Santesson (1942), in Calvelo & Adler (1994) and in Bjerke & Elvebakk (2001).Santesson (1942) had a wide interpretation of this species, including in his list of examined specimens material that has proven to be morphologically and chemically different from M. sanguinascens s. str.
Variation: As stated by Bjerke & Elvebakk (2001), M. sanguinascens is a variable species in terms of its lobe size and morphology, colour of the upper surface, and number and position of the soralia.These features vary with microclimatic conditions.This being said, material from regions Nine to Eleven has a quite uniform morphology and colour, and is easily distinguished from the other sorediate species of Menegazzia.Thus, most variability is seen in the southernmost populations.
Habitat ecology: Menegazzia sanguinascens is a prominent epiphyte of trees in moderately to deeply shaded forests of moderate to high humid Menegazzia subpertusa P. James & D. J. Galloway, New Zealand Journal of Botany 21: 195 (1983).
T y p e : N e w Z e a l a n d , R a n g i t i k e i G o r g e , Wellington, 17 June 1980, J. K. Bartlett (BM, holotype).
This widespread Australasian species is here reported for the first time from America.Detailed descriptions are given in Galloway (1985) and James & Galloway (1992).It is illustrated in James & Galloway (1992) and in Kantvilas & Jarman (1999).
V a r i a t i o n a n d d i s t i n g u i s h i n g f e a t u r e s : Menegazzia subpertusa is characterised by a closely attached thallus with convex, irregularly spreading and often imbricate lobes, small perforations with flat rims, and laminal, convex soralia that are often in groups.Single soralia are small, generally not wider than 0.5 mm.The upper surface is minutely pruinose in parts, smooth, pale grey and often with slightly shining apices.Apothecia containing numerous two-spored asci are common.It is distinguished from M. kawesqarica, which has slightly convex, reticulately maculate, corrugate and dull lobes, and large convex soralia that are readily confluent and spread over the upper surface.Apothecia with mature asci are not known in M. kawesqarica.They also inhabit different habitats.Menegazzia neozelandica is distinct from M. subpertusa by the soralia that are surrounding perforations.However, as stated under M. neozelandica above, it occasionally has soralia that are convex and not associated with perforations.Furthermore, M. subpertusa sometimes has soralia with a central opening, resembling the soralia of M. neozelandica or M. magellanica.The specimen illustrated by James & Galloway (1992, p. 243) actually has one such soralium.In such cases, M. neozelandica is distinct by the glossier lobes, the elevated rims of perforations, the absence of pruina and maculae, and at least some soralia associated with perforations.The more typical convex soralia of M. subpertusa are illustrated in Kantvilas & Jarman (1999).Specimens of M. subpertusa were identified as M. sanguinascens by Santesson (1942), e.g., H. Roivainen s.n., 12.IV.1929.Menegazzia subpertusa is distinct from M. sanguinascens, which has a shiny upper surface, broad, greygreen lobes, and hypothamnolic and thamnolic acids as the major medullary constituents.
Distribution: in South America from latitude 36º50' S (Las Trancas, Eighth Region of Chile) to 54º49' S (Isla de los Estados, Argentina) (Fig. 8).The Argentinean localities reported here are not Habitat ecology: Menegazzia subpertusa is mainly found growing on sclerophyll bushes and scrubs in rather sunny and dry microhabitats surrounded by well-developed forests, e.g., along forest edges, river beds and bogs, and not so much on trees in typical forest vegetation.It prefers smooth bark, such as on young twigs, where it encircles the twigs, so that lobe apices from the same thallus meet and compete for space.It is known from the following phorophytes: Ugni molinae, Berberis spp., Amomyrtus spp., Nothofagus nitida, N.   Quilhot et al. (2002aQuilhot et al. ( , 2002b) ) reported the following sorediate species from Laguna San Rafael: M. chrysogaster, M. fumarprotocetrarica , M. globulifera , M. magellanica, M. neozelandica, M. norsorediata, M. sanguinascens, M. wandae.The presence of M. chrysogaster , M. magellanica and M. sanguinascens in Laguna San Rafael could not be confirmed here.

Ecological requirements
The affiliations of the sorediate species of Menegazzia to the temperate and antiboreal vegetation zones, as defined here, are outlined in Table 1.The temperate high-altitude mixed deciduous-coniferous forest and the antiboreal rainforest are the most species-rich vegetation zones, with eight affiliated species each.The high-altitude forest zone, which is located between 600 and 2,000 m from latitude 37º to 44º S, has several species in common with the antiboreal evergreen and deciduous forests in the southernmost part of Chile.Menegazzia tenuis has a remarkable disjunct distribution, with one known locality on N. dombeyi in a mixed A. araucana-N.dombeyi forest.Also M. kawesqarica is in Central southern Chile mostly restricted to higher altitudes within this vegetation zone.Several lichens from other genera that are common at low altitudes in the antiboreal region are confined to higher altitudes in the temperate region, e.g., Pseudocyphellaria freycinetii, P. lechleri, P. Presentación gráfica de las preferencias ecológicas de las especies sorediosas de Menegazzia en el parte austral de América del Sur.Se presentan las escalas en valores relativos.Nombres ciéntificos son abreviados (las tres primeras letras del epíteto).
ecology: Menegazzia fumarprotocetrarica is most common in fairly open, mixed Nothofagus-Araucaria forests along Cordillera de Nahuelbuta and the Andean Cordillera in the Ninth Region of Chile, where it grows on the hard bark of trunks and branches of Nothofagus alpina, N. dombeyi and Araucaria araucana.On the latter phorophyte, it finds a remarkable habitat between and underneath the needles.In the Ninth Region of Chile, it associates with Menegazzia globulifera, M. norsorediata, Hypogymnia s u b p h y s o d e s , P a r m e l i a p r o t o s u l c a t a , Pannoparmelia angustata, Protousnea spp.and Bryoria sp.The only known locality of M. fumarprotocetrarica in the Eleventh Region of Chile is from a shrubby vegetation dominated by
Fig. 11: Graphical depiction of the ecological preferences of the sorediate species of Menegazzia in southern South America.The scales show relative values.The species names are abbreviated (three first letters in species epithet).
Nothofagus forests from about 150 m in Nahuelbuta to 1,400 m close to the Argentinean border in the Ninth Region of Chile.It has an altitudinal range from sea level to 200 m in the southernmost part of its distribution range.It is most common and best-developed on the lower parts of the smooth-barked trunks of Nothofagus dombeyi, N. betuloides and N. pumilio.

TABLE 1
Vegetation zone affiliation of sorediate species of Menegazzia in Chile.See text for full names of vegetation zones Afiliación de zonas de vegetación para las especies sorediosas de Menegazzia en Chile.Ver el texto para nombres completos de las zonas de vegetación THE SOREDIATE SPECIES OF MENEGAZZIA IN CHILE species associated with Pseudocyphellaria guzmanii and P. compar in Valdivian rainforest (Galloway 1992b).This report of M. magellanica is probably referable to M. wandae.Menegazzia neozelandica was reported from the Eleventh Region, and considered vulnerable and insufficiently known by Quilhot et al. (1998).Based on literature reports (see above), Elix & McCarthy (1998) listed M. sanguinascens and M. terebrata from Islas Juan Fernández.In his treatment of important lichen habitats in Chile, Galloway (1998) reported that M. globulifera, M. magellanica and M. sanguinascens occur in Patagonian Nothofagus forest (regions Eight to Twelve of Chile).Bjerke (2001) described M. wandae and mapped six lo-M.kawesqarica, M. neozelandica and M. norsorediata from localities north of Twelfth Region.