Syntaxonomical review of sub-Antarctic orotemperate ...

International Journal of Geobotanical Research Vol. nº5. 2015. pp. 13-35

Syntaxonomical review of sub-Antarctic orotemperate forests (Adenocaulo chilensis-Nothofagetalia pumilionis) in the Valdivian biogeographic province Javier AMIGO(1) & Manuel A. RODRÍGUEZ-GUITIÁN (2) (1) Laboratorio de Botánica, Facultad de Farmacia, Universidad de Santiago de Compostela (USC). E-15782 Santiago de Compostela (Galicia, España). Phone: 34-881 814977. E-mail: [email protected] (2) Departamento de Producción Vexetal. Escola Politécnica Superior de Lugo-USC. 27002-Lugo (Galicia, España). E-mail: [email protected]

Abstract: Lenga forests (Nothofagus pumilio) are present all over the Andean mountain range throughout the whole of the Valdivian-Magallanic region; they attain their greatest floristic richness in the province of Valdivia. In half a century of phytosociological studies, some 15 associations of forests dominated by lenga have been described in the Chilean-Argentinian strip between latitudes 38º and 42º south. The scarcity of data to support the early published associations, combined with a number of inaccuracies in the identification of species of flora, have led to inconsistent interpretations and successive syntaxonomical proposals, resulting in an over-fragmentation of the units. We have gathered a large number of published relevés and compiled our own relevés in the main areas where the previous data were obtained (almost 600 in total) to propose a reordination and clarification of the syntaxonomy of these forests in the class Nothofagetea pumilionis-antarcticae, and, in the light of our findings, their separation from forestsin the class Wintero-Nothofagetea. We propose that the taxonomy of these forests, in the territory where they attain their greatest biodiversity should be simplified into only four associations, in addition to another two with a more markedly southern presence. Key Words: Araucaria araucana, Argentina, Chile, lenga, orophilous forest, forest flora. Abbreviations: NP = National Park; CNP = Conguillío NP; HNP = Huerquehue NP; LNP = Lanín NP; NHNP = Nahuel Huapi NP; PNP = Puyehue NP; TNP = Tolhuaca NP; VNP = Villarrica NP; VPRNP = Vicente Pérez Rosales NP; VP = Valdivian biogeographic province; MP = Magellanic biogeographic province. Only in the tables: Q-P = Quinchamalio-Pernettyetea Introduction Forestspresided by the deciduous species Nothofagus pumilio (Poepp. & Endl.) Krasser, known by its local name of lenga, are among the most characteristic in extratropical Latin America. They extend from latitude 35º 30’ to 55º south in the meridional areas of the continent and are present as the altitudinal limit of the forest on both the Chilean and Argentinian faces of the Andes throughout their entire extension. Their forest stands have been the object of numerous studies focused both on the dominant species and its characteristics as a forest species (DONOSO 1995; 2006), and on the forest dynamic (VEBLEN ET AL. 1996). From the geobotanical point of view, the formations of this species, along with its close relative Nothofagus antarctica (G. Forster) Oerst., have been used to justify the designation of sub-Antarctic

deciduous forests whose distribution delimits areas of biogeographical importance for several authors: SubAntarctic province (CABRERA & WILLINK 1971), SubAntarctic subregion (MORRONE 2001), South America Pacific Maritime (JOSSE ET AL. 2003) and ValdivianMagallanic region (RIVAS-MARTÍNEZ ET AL. 2011a). Studies have also been published on the variations in the different plant formations throughout this territory in correlation with the bioclimatic belts proposed according to the worldwide classification of RIVAS-MARTÍNEZ (online) and RIVAS-MARTÍNEZ ET AL. (2011b); the works of AMIGO & RAMÍREZ (1998), LUEBERT & PLISCOFF (2006) and AMIGO & RODRIGUEZ-GUITIÁN (2011) covers the widest geographic scope. In spite of the scarcity of precise climate data from meteorological stations in Andean elevations, we can assume the orotemperate

14 J. Amigo & M.A. Rodríguez-Guitián

character attributed to the lenga forests when they develop in pure stands, although at different latitudes in the VP they may be mixed with some other tree species such as Araucaria araucana, without relinquishing their orotemperate character (AMIGO & RAMÍREZ op. cit.; AMIGO ET AL. 2007). This orotemperate belt is located at an altitude of 1700 m at parallel 35º, but is already at sea level at latitude 52º south. With this vast range of latitude and altitude it is unsurprising that lenga should form part of different plant communities throughout its more than 2,000 km of territorial extension. When OBERDORFER (1960) established the basic syntaxonomical checklist for extratropical Chile, he proposed the class Nothofagetea pumilionisantarcticae to include these forestsand differentiate them from the rest of the broadleaved communities in the temperate, boreal and even Mediterranean territories belonging to the class Wintero-Nothofagetea. In his original description he defined three associations: one dominated by Nothofagus antarctica, another by Araucaria araucana and a third by lenga (Nothofagus pumilio); in the case of the last, he interpreted this single association as being present from latitude 36º to Tierra de Fuego. It is therefore only natural that several subsequent studies focusing on more restricted territories including lenga forests found it useful to define new associations for these formations; this is the case of the Argentinian lenga forests in the NHNP described by ESKUCHE (1973) as Macrachaenio gracilis-Nothofagetum pumilionis, the Chilean forests in the VPRNP described by VILLAGRÁN (1980) under the name of Senecio acanthifolii-Nothofagetum pumilionis, and the forests on the volcanoes between the Chilean regions of La Araucanía and the northern part of Los Lagos, which FREIBERG (1985) called Carici minutissimae-Nothofagetum pumilionis. Argentinian authors also proposed lenga communities in the territory of the MP(ROIG ET AL. 1985). The syntaxonomical synthesis published at the time by HILDEBRAND-VOGEL ET AL (1990) was therefore of great practical use, and proposed a distribution in orders, alliances and associations to classify all the known lenga forests, particularly those in the territory of the VP. The proposal of HILDEBRAND-VOGEL ET AL. (op. cit.) provided a model with a biogeographical basis by separating the class Nothofagetea pumilionis-antarcticae into two orders: Adenocaulo chilensis-Nothofagetalia pumilionis and Violo magellanicae-Nothofagetalia pumilionis, differentiated by the fact that the first is distributed in the VP, and the second in the MP. Given that the territory in the VP has a number of bioclimatic features that greatly favour a wider diversity of flora, and added to the fact that it has been the subject of more phytosociological studies, the work included six different associations of lenga forests for the VP and only two for the MP. However, several studies were subsequently made in both the Chilean and Argentinian part, in each case defining new associations of lenga forests and often reclassifying relevés or renaming whole previously published associations to adapt them to each new proposal. In nomenclatural terms there were arguments to support some of these changes, as the previous associations had not been typified or were published as “provisional

name”. This gave rise to a very complicated scenario, in which some 15 different names were proposed for associations distributed on the two Andean faces of the VP, as described in detail in AMIGO & RODRIGUEZ-GUITIÁN (2011). The primary aim of the present work is to resolve these conflicts on the diversity of associations described, and to establish how many and which ones should be recognised. Study area According to the proposals of RIVAS-MARTÍNEZ ET (2011a), the Valdivian-Magallanic region is part of the Austro-American Neotropical kingdom, and the VP is located within it; its delimitation extends from parallel 35º south, at the limit of the Mediterranean/Temperate bioclimate, until 46º south, with a degree of asymmetry in both latitudes depending on whether it is on the Pacific coast or in the Andean mountain range. However the main territory –from which the great majority of the published relevés are taken– can be established more precisely between the Chilean administrative regions of La Araucanía, Los Ríos and Los Lagos, and the Argentinian provinces of Neuquén and Río Negro, between latitudes 38º and 43º. This is also where we have focused our samplings. This Argentinian-Chilean area of territory meets the conditions of being relatively accessible and containing a substantial extension of well-preserved native forests; it marks the confluence of four large contiguous NPs, two in Chile (PNP and VPRNP) and another two in Argentina (LNP and NHNP), together representing a total of 15,000 km2 of well-conserved autochthonous vegetation (see Figure 1). In addition to these large NPs there are several others in the Chilean territory that also contain lenga forests and which have also been used as a data source. Our analysis excludes more northerly lenga forests between parallels 36º and 38º south in Chilean territory. Although some studies have included some of these lenga forests, citing part of their floristic content (PFANZELT ET AL. 2008), or have even cartographically delimited “vegetation belts” dominated by Nothofagus pumilio (LUEBERT & PLISCOFF 2006), they have in no case contributed phytosociological relevés that enable their comparison with the data from more southerly areas; we are therefore unable to venture any opinion on these forests. Nor have there been more than very scarce phytosociological studies in the southernmost part of the VP. We are aware of the publication of little more than one association of lenga forests in Chilean territory above latitude 45º south (HILDEBRAND-VOGEL ET AL. 1990), and another described in Argentinian territory at latitude 43º 50’ (ESKUCHE 2002); both associations are included in our analysis, although we do not have our own data on these more southerly territories. AL.

Methodology All the information compiled from the scientific literature and our own field samplings follows the Sigmatist phytosociological methodology (BRAUN-BLANQUET 1979, updated by GÉHU & RIVAS-MARTÍNEZ 1981); this uses floristic composition and quantitative presence indexes to establish the type of data for the comparative

Syntaxonomical review of sub-Antarctic orotemperate forests in the Valdivian biogeographic province

analysis of the results of the bibliography, and the rules for accepting or rejecting syntaxonomical names (WEBER ET AL. 2000). We have also personally covered and inventoried lenga forests throughout the most biodiverse and best-conserved areas of the VP in several campaigns carried out in the austral summers of 2006, 2007, 2008, 2009, 2010 and 2015.

15

Uncinia are particularly significant, although the inaccuracies in the older works are more understandable, as new species were discovered in both genera in the last decade of the 20th century (see WHEELER 1989, 1997a and 1997b). In compiling the comparative table (Table 1) we have attempted to show some cases in which the determination of a species was erroneous, either demonstrably or presumably. Examples are the citations of Uncinia negeri which are assumed owing to its confusion with U. lechleriana; in some cases the error has been proven, while in others it is only presumed. The quantitative information for the species in this second case is marked with the symbol “?”. Other examples of confusions are Carex lateriflora interpreted as Carex minutissima, and the presence of Ribes nitidissimum initially published as Ribes sp. in lenga forests in territories where our own data have demonstrated the exclusive presence of that particular species of Ribes. Cases have also occurred where the author was originally unable to distinguish between two very similar taxa; these are highlighted in Table 1, although applying a valuation based on our own criterion, as is the case of the pair Blechnum microphyllum/B. penna-marina jointly included by FINCKH (1996) in his tables, but for which we have been able to supply a little more information in our own recent studies (AMIGO & CASTRO 2015). The phytosociological assignment of characteristic species takes into account the criteria of previous authors, but particularly our own criteria; as highlighted previously (AMIGO ET AL. 2010), it is useful to assume some species of flora that can be considered shared between the forests of Nothofagetea pumilionis-antarcticae and those of Wintero-Nothofagetea. Results & Discussion

Figure 1. Map of the central-northern Valdivian biogeographical province (VP), showing the protected areas (NP) and sites (ranges, hills, volcanoes, lakes, passes) cited in the text.

One of the main problems affecting all the phytosociological studies in these forests is the difficulty in identifying some of the species of flora in the communities in which the lengas are present. A recent publication (AMIGO & CASTRO 2015) has highlighted some identification errors, inconsistencies and inaccuracies, some of which continue unresolved today; this point is important when comparing data on the floristic contents of lenga forests published over half a century ago, and we attempt to highlight it in our synthetic table (Table 1) in order to minimise possible errors. In our floristic samplings we have particularly taken into account the main taxonomic bibliography published on Argentinian-Chilean flora in the last two decades, and we use the Flora del Cono Sur (ZULOAGA ET AL. 2009) as a reference. One exception is the case of the species of the genus Ribes, where we adopt a Latin binomen that is merely assumed as a synonym in the work of ZULOAGA ET AL. (op. cit.); this genus was organised based on the proposals of SPARRE (1984), and advances have been made today thanks to the monograph of HECHENLEITNER (2007). The errors in the determinations of some cyperaceae in the genera Carex and

The synthetic table (Table 1) shows a large part of the published diversity of the communities dominated by lenga, organised in associations according to the authors’ proposals. The first columns (no.1-11) contain the northernmost communities included in the present study: the orophilous forests of Nothofagus pumilio and Araucaria araucana (forests of lenga-pehuén, to use their local name) that extend throughout the region of Araucanía in Chile and the province of Neuquén in Argentina. The following columns show other communities of lenga forests described in both countries in the areas where Araucaria is no longer present; the orotemperate forests are therefore the exclusive domain of lenga, except in specific cases in which they include Nothofagus betuloides. The communities included in Table 1 are not the only ones that have been described with a presence or even dominance of lenga. Relevés have been collected and studied on both sides of the mountain range in communities characterised by the coexistence of Nothofagus pumilio and part of its characteristic companion species, along with mainly evergreen trees and herbaceous species typical of the class Wintero-Nothofagetea with which the lenga forests come into contact as they descend from the supratemperate belt. These contact situations between two phytosociological classes have been interpreted in different ways by the various authors who

Column number 1 2 3 21 Number of relevès Araucanía-Neuquén territory Araucaria araucana 3 V Alstroemeria aurea · III Valeriana laxiflora · II Vicia nigricans · III Leucheria thermarum · · Anemone antucensis 1 II Olsynium scirpoideum (* = subsp. luridum) 1? III* Senecio pilquensis · II Calceolaria filicaulis · + Festuca scabriuscula (sub F. thermarum) 1 · Senecio argyreus · · Los Lagos ultraperhumid territory Blechnum penna-marina · · Gunnera magellanica · · Dysopsis glechomoides · · Nothofagus betuloides · · Festuca thermarum · · Acaena antarctica · · Acaena magellanica · · Senecio acanthifolius · · Ranunculus chilensis · · Oxalis magellanica · · Schizeilema ranunculus · · Differential shrub taxa Drimys andina · V Escallonia alpina · II Chusquea culeou + Ch.·montana · IV Ovidia andina (sub O. pillo-pillo) · + Chiliotrichum rosmarinifolium 2 + Near 44º or further South Senecio patagonicus · · Anthoxanthum redolens · · Leucheria cf.·magna · · Berberis ilicifolia · · Characteristic taxa of Nothofagetea pumilionis-antarcticae Nothofagus pumilio 3 V Maytenus disticha 2 V Adenocaulon chilense 3 V Codonorchis lessoni 3 III Berberis serrato-dentata 1 I Viola reichei 1 V V V II V V IV III V I II IV IV · · III · + + IV · I · · · · · · · · · · · · · · · · II · · · · V III V V I ·

V II III III · I III III III II · · · · · · · · · · · · · III · · II · · · · V V V II · V

V IV V III IV IV

· · · ·

III · V · ·

· · + · · · · · · · ·

4 16

3 7

5 14

· · · · · · · · · · ·

V IV V IV III IV

· · · ·

· · · ·

· I · · ·

· · · · · · · · · · ·

· · · ·

I I · I ·

· · · · · · · · · · ·

· · · · · · · · · · ·

· · · ·

V II II II · II

· · · ·

· · · ·

II + I · ·

· · + · · · · · · · ·

· · · ·

· · · · ·

· · I · · · · · · · ·

· · · · · · · · · · ·

· II III III III II · · III · · + · · · · · · · · · ·

· III III V IV · · · · · III · · · · · · · · · · ·

· · 1 3 2 · · · · · 1

· · · ·

V V V III V II

· · · ·

V V V + V ·

· · · ·

3 3 3 3 3 2

· · · ·

· · · 1 II + I · I I · 2 · · · · V II III 1

· · · · · · · · · · ·

· II IV V III + · · · · +

5 V V V + IV IV V 1 V V V · IV III II + III V V 1 V II III

· · · ·

· · · · ·

· · · · · · · · · · ·

I · + · IV · V II II · II I V · II III II · IV IV · 1 V III · ? IV · · · · · · · II + I · · · · · · ·

III I III · + III II I · ·

· · + · · · · · · · ·

V V V V V IV IV V I IV IV V II II IV IV I I + III · II III V

· · · ·

I I + · III · I III · ·

· · · · · · · · · · ·

IV III V IV II IV · · I + II · · II + III · · I · · · · · · · · · · · I · · III IV · · · · · · · · · · + I II V I · · · · · · II + · · · · + · · ·

II I II · II · · + · · ·

II · I II · I + · · · ·

· · · · · · I · · I · · + · · · · · · II* · · · · · · · · · · · · ·

· · · · · · · · · · ·

· · · · · · · · · · ·

· · · · · II · · · · ·

· · · · · I · · · · ·

· · · · · · · · · · ·

V IV V III IV IV

· I · ·

V V V IV V V

· I · ·

V V V II II V

· · · ·

V V IV III V II

· · · ·

· · · · · · I* · · · ·

· · · · · · · · · · ·

· · · · · + · · · · ·

V V V IV · V

· · · ·

V V III II r IV

· · · ·

IV V III · I IV

· · · ·

V V V III IV V

· · · ·

V V IV II III III

· · · ·

V V V IV V V

· · · ·

V V V IV II III

· · · ·

I IV · · ·

V IV I III I V

· · · ·

IV V + · ·

· · · · · · · · · · ·

· · · · · · · · · · I

II · · · · · · · · · ·

· · I · · · · · · · ·

III III III IV · · · · · · ·

· · · · · · · · · · ·

V III V III I V

· · · ·

V V V III V ·

V V V V V V

V V IV · I V

II III · III IV · · II · · · II

I · I · III III III IV · · · · · · · · · V V ·

III I I IV III V I III II IV III I · II III V V I · · · IV II III III II III I II V · · · · I · III IV · · · · · · · + II · + II · III II II · · · · II V · + + + · · · IV IV V V III · · · · I IV · II I · · · · · I I II + · · · · · I · II II

· · · · · · · · · · ·

V V V V V V V V V IV V · III III V IV V IV IV IV III IV II · II r V · I I · · III + IV IV V · · I III I · · · · · · · ·

· III I · · · · I · · ·

· · II · · + · · · · +

6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 22 13 8 15 13 8 1 11 19 10 23 16 3 19 20 22 13 11 40 7 13 8 8 7 14 24 12 16 5

Table 1: Main published relevés of Chilean and Argentinian lenga or lenga-pehuén forests in VP.


Berberis montana (sub B. buxifolia subsp. andina) · V III I III IV Perezia prenanthoides 1 V IV V III III Gavilea lutea 1 II V IV · II Rubus geoides 2 III II + + II Macrachaenium gracile 2 II II · + I Uncinia negeri (sub U. lechleriana?) · IV II II II III Senecio prenanthifolius 2 + I I II I Lycopodium magellanicum 1 II · I II I Carex lateriflora (sub C. minutissima) 2 II II · I + Perezia pedicularifolia 1 II III I · I Lagenophora harioti · II I · II + Ribes nitidissimum (sub Ribes sp.) · V V I · III Blechnum microphyllum (* = + B. penna-marina) 1 II III III? III? III* Trisetum cernuum · + I I I · Ranunculus peduncularis · + II · · · Arachnitis uniflora 1 · · IV II I Hypochaeris tenuifolia 1 + · · · · Nothofagus antarctica · · · · · + Nothofagetea pumilionis-antarcticae & Wintero-Nothofagetea shared taxa Myoschilos oblonga 1 V V V III IV Ribes magellanicum 1 II II + V III Osmorhiza chilensis · IV III V III III Polystichum plicatum 1 + I · III I Berberis microphylla · + II + II II Valeriana lapathifolia · I I · · · Embothrium coccineum · I · · · + Lagenophora hirsuta 2 · · · · · Berberis trigona · IV I · · + Uncinia scabriuscula (sub U. triquetra) · I · · · · Gaultheria phyllireifolia · · · · · · Myrceugenia chrysocarpa · II · · · · Uncinia tenuis 1 · · · · · Azara alpina · III I · · · Dioscorea brachybotrya · II · I + · Companion taxa Acaena ovalifolia · II I + V II Gaultheria poeppigi 1 I III · · II Poa obvallata (Q-P) · · · V · + Gaultheria pumila (Q-P) · + · · · I Bromus catharticus · · · III II · Misodendron punctulatum · · · · · I Ribes cucullatum · · · · · + Uncinia chilensis (sub U. brevicaulis?) · + I I I · Hypochaeris arenaria (incl.·aff. palustris) · + · IV · II Misodendron linearifolium · · · · · + Senecio chilensis (sub S. chionophilus)(Q-P) · · · · · + + + + II · II · · I · I

· · I I I I I · III IV V V · · · · II II · · II · · · · · III I · II · I II

IV IV II V V IV III I IV IV II V II* · · · II · IV II III I · · + I II · · + I · ·

IV II · · III · II · I · · · · · ·

IV II · I II · I · · · · · · · ·

V · IV II + III + II I III · IV II* · · II I + IV I III I IV · + · I · · + · · ·

II II · · · II · IV I III · II II* · · · · ·

V · I II · I · · · · · V I* · · · · ·

V I · · · II II · · II ·

II V V III IV · · · · · · II · · ·

IV · I · · I · · · · · · I* · · · · I

V V I + IV II V · · · · I II* V · + · ·

V V III II I V IV · III · + IV II? I II · · ·

V V II II II III III · · · · V II? II · · · ·

IV V V II I III II · III + · V III + II · · ·

· V IV · · III · · · I · IV · + · · · ·

2 2 1 · 2 1 · · 1 · 1 · 2? · · 1 · ·

· · · · · · · · · · ·

V III IV III II 2 · · II · II 1 + I II III V 1 · · · · · · · II III · III 1 + · · · · 3 · · II · III · · · · · · · I I · I + · + · · · · 3 · · · · · ·

· + III IV III IV 1 + V III · II · 2 + V V IV V V 3 · I + II II I 1 · · · I · II 1 2 III · · I · · · · · · · · · · II · · · · · · · · · · · · · · III · II · · · · · · · · · · · · · · · · + + · · · · · · · · · · · · · · · · · · ·

· 4 (+) 1 + · + · · · · · + +? · · · ·

III III II II V II · + I · II IV II? II · + · ·

IV III I III IV IV II · · I II IV IV · III · · ·

IV + II IV III II? + IV · + I · · · · · · ·

V · · V r · · III r IV · · · · r · · II

III · · IV II · · I · · · · · · · · · · III III II + · V · · · · + · · · ·

I II II II IV II? I II · · II · · II · · · · I III IV I · V · II · · I · · · ·

II III I II IV I? I III I · I · · II · · · ·

· II I III II II · · · · · · · · · · · · · · · · +? · · · · · · · I · ·

· I I V IV III II · · · · · · · · III · · + II III V III III · · I · · · I r II · · · · · · · · · · · ·

V · · V V · · II I IV · · · · · · · +

III III IV + V + IV III II IV II II II + + · III II · · · + IV V I + · · · · · · · · · · I · · · · + · · · · III? +? + · · · · · · · · · · · · · · · + IV

IV IV III II · II III I III IV III I + + II + I I + + II III III · + · I · · · · + · II + · I I II · · · · · · · + · · · · · · · · · · · · ·

V III III IV IV II · II I III III V III? + · · · · III · III · · IV · · · · · · · · ·

V V III V V I · III III III III V III · V · · · · II · · · V · II · · + · · · ·

II + · III V +? + III · III III · · · · · · +

· II II II IV III · I · · · II · · · I · · · · · · · · · · · · · · · · ·

· II IV I · V · · · · · · · · ·

V V · II III · II · · II · IV · · · · · ·

I + · + · · + I? · · ·

II III IV · · IV · II · · · · · · ·

II IV II IV IV +? I + II III II · · I + · · ·

+ V · · · V III · · IV ·

V · III · III · · · · · · · · · ·

· · IV III · · · · · · · · · · · + · ·

III II III IV · · · · II · V · II · · · · · III · · ·

V I I II V I · I II · II IV · · · · · · II · · · · · · · · · · ·

· · V · V · II II III V III · · · · · I · · · I II · · · · · · I · + · · · · ·

Syntaxonomical review of sub-Antarctic orotemperate forests in the Valdivian biogeographic province 17

· 2 · · · · · · · · · · · · · · 1 · · ·

· · · · · · · · · · · · · · · · II · · ·

· · · · · · · · I · · · · · · · I · · ·

· III · · · · · · · · · · · · · · · · · ·

· · · II · · · · · · · · · · · · · · · ·

+ · + I · I · + · · · · · · · · · · · ·

· · · · I · · · · · · · · · · · · · · ·

V · · · I I · + V II I IV V II V II · · · · · · · · · · · · · · · · · · · II · · · ·

· · · · · I · · · · · · · · · · · · · ·

· · · II · · · · · · · · · · · · · II · ·

· · · · · · · · · · · · · · · · · · · ·

· · · + · + · · · · · · · · · · · · · ·

· · · · · · · · · · · · · · · · · · · ·

· · · · · · + · · · · · · · · · · · · ·

· · · · · · · · · · · · · · · · · · · ·

· + · + + · + · · · · · · · · · · · · ·

· · · · · · · · · · · · · · · · · · 1 1

· · · · · · · · II · · · · · · · · · · ·

· · · · · · · · I · · · · · · · · · · ·

· · · · · · · · · · · II · · · · · · · ·

I · · · · · · · · · · · · · · + · · · ·

· II · · · · + · · II + III · · · · · · · ·

IV III · · II · III · · II II IV · · · · · · · ·

· · · · · · · · · · · II · · · · · · · ·

· · · · · · · · · · · · · · · · · · · ·

· · · · · · · · · · · · · · · I · · · ·

· · · · · · · · · · · · · · · II · · · ·

· · · · · · · · · · · I II I II · · · · ·

I · + · I · · · · · II I IV II · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · · I · · · · II I · · · II + · · · + I · · · + · · · I · II* · · · · · II · · · · III IV · · · III I ·

Sources of relevés (designed with the original names proposed by the authors). The syntaxa names between quotation marks are considered rejected names or misinterpretations of the original definitions: 1) Carici-Araucarietum:·OBERDORFER (1960):·Tb·38: 3 rel., number R5 excluded. 2) Carici lateriflorae-Araucarietum araucanae subass. drimydetosum andinae: 17 own relevés + 4 from FINCKX 1996). 3) Carici lateriflorae-Araucarietum araucanae subass. typicum: 7 own relevés. 4) “Nothofago-Araucarietum araucanae typicum”:·ESKUCHE (2002). 5) “Nothofago-Araucarietum araucanae chusqueetosum”: ESKUCHE (2002). 6) “Chusqueo-Nothofagetum pumilionis (Fragmentarische Ausbildungen)“: FINCKH (1996). 7) “Pernettyo-Nothofagetum pumilionis”: FINCKH (1996). 8) “EmpetrumAraucaria gessellschaft:“ FINCKH (1996). 9) “Festuco scabriusculae-Nothofagetum pumilionis“: FINCKH (1996). 10) “Macrachaenio-Nothofagetum pumilionis”:·FINCKH (1996). 11) “Nothofagetum pumilionis disturbed”:·FINCKH (1996). 12) Anemono-Nothofagetum typus relevè: OBERDORFER (1960). 13) Anemono-Nothofagetum pumilionis FINCKH (1996). 14) Anemono-Nothofagetum pumilionis typicum·ESKUCHE (2002). 15) Anemono-Nothofagetum pumilionis chiliotrichetosum rosmarinifolii: ESKUCHE (2002). 16) Anemono-Nothofagetum pumilionis (own Argentinian relevés). 17) “Senecioni linariifolii-Nothofagetum pumilionis”: ESKUCHE (2002). 18) Macrachaenio-Nothofagetum pumilionis: ESKUCHE (1973). 19) Macrachaenio-Nothofagetum pumilionis escallonietosum: ESKUCHE (2002): tb·4 (rel.·1-2 excluded). 20) Macrachaenio-Nothofagetum pumilionis typicum:·ESKUCHE (2002). 21) Macrachaenio-Nothofagetum pumilionis subass. ovidietosum andinae (own Chilean relevés). 22) “Drimydo andinae-Nothofagetum pumilionis” (typicum + var. Empetrum rubrum + var.·Nothofagus betuloides): FLORES-TORO & HILDEBRAND-VOGEL (2006). 23) “Carici minutissimae-Nothofagetum pumilionis Ausbildung mit Valeriana lapathifolia und Macrachaenium gracile”: FREIBERG (1985) sensu FINCKH (1996). 24) “Carici minutissimae-Nothofagetum pumilionis” Ausbildungen mit Embothrium coccineum·FREIBERG (1985) sensu FINCKH (1996). 25) “Chusqueo-Nothofagetum pumilionis nothofagetosum betuloidis” FREIBERG (1985) sensu FINCKH (1996). 26) “Valeriano lapathifoliaeNothofagetum pumilionis typicum”: FLORES-TORO & HILDEBRAND-VOGEL (2006). 27) “Valeriano lapathifoliae-Nothofagetum pumilionis gunneretosum magellanicae”: FLORES-TORO & HILDEBRANDVOGEL (2006). 28) Senecioni acanthifolii-Nothofagetum pumilionis VILLAGRÁN (1980). 29) Senecioni acanthifolii-Nothofagetum pumilionis subass. gunneretosum magellanicae (own Chilean relevés). 30) Senecioni acanthifolii-Nothofagetum pumilionis subass. gunneretosum magellanicae: FLORES-TORO & HILDEBRAND-VOGEL (2006). 31) Senecioni acanthifolii-Nothofagetum pumilionis (typicum + “var.·de Perezia pedicularifolia”): FLORES-TORO & HILDEBRAND-VOGEL (2006). 32) Chiliotricho rosmarinifolii-Nothofagetum pumilionis typicum: ESKUCHE (2002). 33) Chiliotricho rosmarinifoliiNothofagetum pumilionis violetosum reichei:·ESKUCHE (2002). 34) Berberido ilicifoliae-Nothofagetum pumilionis HILDEBRAND-VOGEL ET AL.·(1990).

The symbol “?” means that there is not absolute certainty that all individuals determined by each author correspond to the name given in this table. Shaded columns are those that contain the relevé typus of the associations.

Empetrum rubrum (Q-P) Sisyrinchium arenarium Poa alopecurus Elymus angulatus Quinchamalium chilense (Q-P) Sisyrinchium pearcei Anthoxanthum juncifolium (Q-P) Adesmia emarginata (Q-P) Senecio subpubescens Agrostis philippiana (Q-P) Senecio triodon (Q-P) Cortaderia egmontiana (Q-P) Caltha appendiculata Ourisia breviflora Marsippospermum philippii Hymenophyllum peltatum Cardamine aff.·cordata (* =?) Discaria chacaye Misodendron oblongifolium Misodendron quadrifolium



have studied them using the Sigmatist phytosociological methodology. Table 2 shows a compilation of 136 relevés and the different designations with which they were proposed by their authors. Some authors preferred to propose the communities in this contact as new associations, such as FINCKH (1996) in the VNP and VILLAGRÁN (1980) in the VPRNP; those who produced work in geographically more extensive territories have chosen to use the rank of subassociation or variant. In our opinion the second option is preferable: the interpretation of these contacts as communities of a lower rank than association and preferably as a subassociation or variant of the altitudinally lower association, as proposed by the authors of the communities in columns 1, 2 and 3 in Table 2. It is biogeographically more consistent to interpret a supratemperate forest presided by Nothofa-

19

gus dombeyi, which may be constituted by three different associations in TNP, VPRNP and NHNP, as a possible subassociation nothofagetosum pumilionis when –for reasons of its altitudinal position– it includes the presence of lenga, rather than the opposite alternative: an orotemperate lenga forests that is modified towards a subassociation nothofagetosum dombeyi on descending to the supratemperate belt and mixing with different forests of Wintero-Nothofagetea. We do not therefore consider the proposals for associations shown in columns 4 to 9 (Table 2) to be correct, and judge it more acceptable to propose a future syntaxonomical ordination that subordinates these communities to other supratemperate associations, a task which goes beyond the scope of the present work.

Table 2: Mixed forests of Nothofagus pumilio/other Nothofagus species proposed to be classified in class Nothofagetea pumilionis-antarcticae Column number 1 2 3 4 5 6 7 8 9 15 15 18 24 29 5 13 6 11 Number of relevés Characteristic taxa of Wintero-Nothofagetea Nothofagus dombeyi V V V V II V V IV II Gaultheria phyllireifolia IV II IV IV r III II IV I Berberis trigona III III II IV III II IV V · Azara lanceolata IV V IV I r IV II · I Desfontainia spinosa · III V IV + V III III I Maytenus magellanica IV I II II + III I · · Blechnum magellanicum I II III · · I · II II Asteranthera ovata · · III · · V I II II Berberis darwini I I · · · I · · · Nothofagus alpina V IV · · · · · · · Raukaua laetevirens III · · · · · · · + Philesia magellanica · · · · · · · II + Nothofagus betuloides · · · · · · · IV + Taxa of the Araucanía-Neuquén area Alstroemeria aurea IV I I + II · · · · Araucaria araucana r · I III II · · · · Valeriana laxiflora I I · r I · · · · Vicia nigricans I + · I II · · · · Taxa of the Northern Los Lagos area Blechnum penna-marina · · · · · · · III V Acaena magellanica · · · · · · · III + Gunnera magellanica · · · · · · · · II Senecio acanthifolius · · · · · · · · II Characteristic taxa of Nothofagetea pumilionis-antarcticae Nothofagus pumilio II V III V V V V V V Maytenus disticha IV V IV V V V V V V Adenocaulon chilense IV II I III IV II V IV IV Berberis serrato-dentata II I I I III II V IV V Viola reichei II II III II IV II V V IV Berberis montana I I I II V III V II III Codonorchis lessoni II I + · II · IV IV + Blechnum microphyllum (*+ B.·p-m) · +? I* I* II* III? IV? · · Macrachaenium gracile · · · r II II V IV IV Escallonia alpina I · · I II · · V III Perezia prenanthoides · + · I II · II · III Rubus geoides · · + · I · · III III Ribes nitidissimum · · + II IV · · · · Uncinia negeri · · + + II · · · · Lycopodium magellanicum · · · r I · · · + Lagenophora hariotii · · · · · · III · III Carex lateriflora · · · r I · · · · Nothofagetea pumilionis-antarcticae & Wintero-Nothofagetea shared taxa Drimys andina V V V V V V V V V Chusquea culeou + Ch. montana V V V V IV V V II III Osmorhiza chilensis III III III I II I IV III III Myrceugenia chrysocarpa V V V V IV V II · ·

20 J. Amigo & M.A. Rodríguez-Guitián Dysopsis glechomoides Valeriana lapathifolia Embothrium coccineum Polystichum plicatum Ribes magellanicum Lagenophora hirsuta Myoschilos oblonga Arachnitis uniflora Ovidia andina Companion taxa Acaena ovalifolia Gaultheria poeppigi Nertera granadensis Schinus patagonicus Adesmia emarginata Hypochaeris arenaria Gaultheria pumila Anthoxanthum juncifolium Ribes cucullatum Uncinia brevicaulis

I · II I III + III · ·

II III + II III + II + ·

I I I I · · · · ·

r · I + · · · r +

r r I I · · · r II

· III · · II · · · ·

+ V · + · · · · ·

II V III · IV V · · ·

II V · · · I · · ·

I · + + · · · · · ·

II I I I · · · · · ·

II I · · + · · · · ·

+ II · · · r r r · ·

II II · · + + I r r r

I I III · · · · · · ·

II II · · · · · · · ·

· II · · · · · · · ·

I III · · · · · · + +

Taxa present in only 1 relevè: Characteristic taxa of Wintero-Nothofagetea: Azara microphylla I in 1; Berberis microphylla I in 2; Blechnum mochaenum + in 1; Dioscorea brachybotrya I in 1; Elytropus chilensis II in 1; Fuchsia magellanica I in 1; Greigia landbeckii I in 9; Hydrocotyle poeppigi I en 1; Laureliopsis philippiana + in 1; Uncinia phleoides II in 1; Uncinia scabriuscula + in 2. Characteristic taxa of Nothofagetea pumilionis-antarcticae: Gavilea lutea + in 2; Hypochaeris tenuifolia + in 5; Leuceria thermarum + in 2; Nothofagus antarctica r in 5; Olsynium scirpoideum subsp. luridum + in 5; Perezia pedicularifolia + in 5. Companion taxa: Acaena antarctica + in 9; Cortaderia egmontiana r in 5; Desmaria mutabilis II in 1; Dioscorea auriculata II in 1; Empetrum rubrum r in 5; Gamochaeta spiciformis + in 9; Hymenophyllum peltatum II in 8; Hymenophyllum rugosum I in 8; Lycopodium paniculatum II in 8; Misodendron punctulatum I in 5; Mutisia spinosa I in 1; Myrceugenia ovata var.·nanophylla II in 1; Ovidia pillo-pillo III in 8; Perezia nutans I en 1; Ranunculus chilensis + in 9; Senecio chilensis r in 5; Senecio otites I in 1

Sources of relevès: 1.·Nothofagetum dombeyi-alpinae nothofagetosum pumilionis·Malleco Province, Araucanía Region (POLLMANN 2001). 2.·Nothofagetum dombeyi-alpinae subass. pumilietosum·Neuquén Province (ESKUCHE 1999). 3.·Gaultherio phyllireifoliae-Nothofagetum dombeyi·VNP, Araucanía Region·(FINCKH 1996). 4.·Gaultherio phyllireifoliae-Nothofagetum dombeyi araucarietosum VNP, Araucanía Region (FINCKH 1996). 5.·Chusqueo culeou-Nothofagetum pumilionis·VNP, Araucanía Region (FINCKH 1996): 30 (- 1) relevés by subtraction of rel. H-2 from HILDEBRAND-VOGEL ET AL. (1990). 6.·Myrceugenio-Nothofagetum dombeyi pumilietosum·Neuquén Province (ESKUCHE 1999). 7.·Macrachaenio-Nothofagetum pumilionis nothofagetosum dombeyi·NHNP, Río Negro Province (ESKUCHE 2002). 8.·Nothofagus dombeyi-Nothofagetum pumilionis·VPRNP, Los Lagos Region (VILLAGRÁN 1980). 9.·Valeriano lapathifoliae-Nothofagetum pumilionis (dombeyetosum + tipo de Chusquea),·several sites from Los Lagos Region (FLORES-TORO & HILDEBRAND-VOGEL 2006). Forests of lenga-pehuén As shown in the first column of Table 1, the orotemperate Chilean-Argentinian territories of Araucanía and Neuquén contain extensive forests of lenga in which the most visible element is the pehuén (Araucaria araucana), which always overshadows the lenga owing to its greater height. The community it forms in the most part is a mesophytic forest association on trumao-type soils derived from volcanic activity (BESOAÍN 1985). The association that may serve to explain and encompass the floristic contingent of lenga-pehuén forests is the one proposed by OBERDORFER (1960): Carici laterifloraeAraucarietum araucanae (the nomenclatural precision derived from the correct identity of Carex lateriflora is discussed in AMIGO & CASTRO 2015). The original publication was presented by OBERDORFER (op. cit., Table 38, page 138/139) and included only four relevés in one table, of which three were presence/absence lists of species taken from REICHE (1907); the only relevé contributed by Oberdorfer came from Quetrupillán in the centre of the VNP (OBERDORFER op.

cit., Table 38, rel. 230). Given that this relevé was the most abundant in species and the only one with quantitative abundance-dominance indices for each species, it was taken as the reference point by numerous subsequent authors for reinterpretations of this community. It was considered equivalent to the holotype for the association; when Oberdorfer published his work, there was no code of phytosociological nomenclature in existence, nor was there a requirement to designate the nomenclatural type. However, any proposed lectotype for this name must necessarily be based on that single complete relevé. The set expressed by this table is shown as column 1 in our Table 1. In the same work Oberdorfer presented a detailed study of the soil profile in the terrain where he collected his relevé no. 230, showing a soil with a depth of 120 cm, “of the same type as the brown soils in the beech woods of Central Europe”; it is important to recognise that although this relevé has a sunny aspect, it did not correspond to any position that was more xeric in soil terms nor particularly deviating from what can be interpreted as the position of a climactic forest.


In a study that intensively inventoried VNP, FINCKH (1996) recognised the site of Oberdorfer’s relevé and classified it as a place “with a particular accumulation of snow”; for this reason, and due to the presence of the hemicryptophytes typical of chionophilous lenga forests, he decided to interpret the Carici-Araucarietum as a “variant of Araucaria araucana”, and within a typical association of lenga forest: the Macrachaenio-Nothofagetum pumilionis described for the Argentinian territory by ESKUCHE (1973). A more drastic deviation, however, was the interpretation made subsequently by ESKUCHE (2002), who identified as “Carici-Araucarietum Oberdorfer 1960” an impoverished version found in Argentinian territory in the area around the Ruca-Choroi lake, whose composition he recognised as being “degraded by grazing”; he collected three relevés with a scant presence of the hemicryptophytes and geophytes typical of lenga forests, to which he added as differential species Carex setifolia, Carex aphylla, Lathyrus multiceps and Luzula racemosa, none of which were present in OBERDORFER’s original table (op.cit.). With this interpretation, he deviated the concept of “Carici-Araucarietum Oberdorfer 1960” and described a new association which he called Nothofago pumilionis-Araucarietum araucanae, for which he collected around 30 relevés in Argentinian territory in the province of Neuquén, in what CABRERA (1976) called the “District of El Pehuén”. These relevés are shown in columns 4 and 5 of Table 1. In our opinion the lenga-pehuén forests in territories in both Chile and Argentina should be interpreted as a single main association: Carici lateriflorae-Araucarietum araucanae. The name proposed by ESKUCHE (2002) is therefore superfluous as it refers to the same community. By way of comparison, Table 3 shows a sample of lenga-pehuén forests studied by us in mainly Chilean territory corresponding to the region of La Araucanía (Figure 2).

Figure 2. Araucaria araucana trees usually stand up over the lenga canopies in lenga-pehuén forests like this located at the foothills of Llaima Volcano; Conguillío NP (La Araucanía Region, CHI)

● Variability: orotemperate lenga-pehuén forests at a lower altitude As seen in Table 3, lenga-pehuén forests are recognisably differentiated into two subassociations, which both overlap at different altitudinal levels. Forests at higher altitudes which thus receive a greater quantity of snow and for

21

a longer time are characterised by a clearer understory containing few shrubby species of a modest height (Berberis montana, Ribes nitidissimum and Maytenus disticha are generally recognised as camephytes), thus giving far more opportunity for the development of the hemicryptophytes and geophytes typical of lenga forests. The indisputable dominance in the upper tree canopy of Araucaria araucana means that the community as a whole is somewhat more heliophilous than pure lenga forests(with which they share numerous species); evidence of this greater heliophilia is found in the presence of species in the understory that are well adapted to the upper edge formations in the forest, such as Festuca scabriuscula and Chiliotrichium rosmarinifolium. This is the community we identify as the typical association. At a lower altitude, but still in the orotemperate belt, lenga-pehuén forests may have an understory with a substantial shrubby biomass although comprising a reduced number of species. These are mainly Drimys andina, Berberis trigona, Myrceugenia chrysocarpa and Chusquea montana + Chusquea culeou, among others: we propose this community as Carici lateriflorae-Araucarietum araucanae subassociation drimydetosum andinae subassociatio nova hoc loco (holotypus rel. 4, Tb. 3). ESKUCHE (2002) also noted a phenomenon in Argentinian territory equivalent to that of this subassociation, and therefore proposed a subassociation chusqueetosum culeou for his Nothofago-Araucarietum nomen nudum, which he differentiated due to the abundant presence of Chusquea culeou in the understory. AMIGO & CASTRO (2015) have already discussed the difficulty of identifying the Chusquea that extend into lenga forests, and the advisability of closely following the delimitation of these species in forestson the Argentinian side; it is also possible that the lenga-pehuén forests inventoried by ESKUCHE (op.cit.) are participated by Ch. montana to a large degree –in common with those on the Chilean side–, and thus Carici lateriflorae-Araucarietum subass. drimydetosum andinae could itself be interpreted as being present in Argentinian territory. If on the other hand it were to be demonstrated that only Chusquea culeou predominates in these lower-altitude lengapehuén forests in the province of Neuquén –as opposed to the presumably majority Ch. montana in Chilean territory– there would be an argument for establishing a different subassociation (such as the proposed chusqueetosum culeou) to distinguish the forests on the eastern side of the mountain range. The relevés designated by ESKUCHE (op.cit.) to define his subass. chusqueetosum culeou are also lacking in some plants that are present in the subass. drimydetosum andinae in Chilean territory, such as Azara alpina, Senecio pilquensis and Olsynium scirpoideum subsp. luridum; in contrast, there is a presence of Leucheria thermarum which does not appear in any of our Chilean relevés. In summary, the consideration of a single subassociation for the less orophilous lenga-pehuén forests, or else two (one Chilean and one Argentinian) should be supported to a large extent by a detailed study to identify the species of Chusquea involved in both cases.

1 1445 10 SE 95 22-28 200 15

2 1490 10 SSE 95 12-20 400 24

1 1 + · 2 3 · · · · · · · · · ·

2 + 1 + + · · 1 · · · + · 1 · ·

3 1 1 + 2 3 · · · · · · · · · ·

3 1530 15 WSW 90 20-25 250 22

2 · 3 · · · ·

Nothofagus pumilio Maytenus disticha Viola reichei Adenocaulon chilense Perezia prenanthoides Rubus geoides

5 3 1 1 1 ·

Characteristic taxa of Order & Class

Drimys andina Berberis trigona Chusquea culeou + Ch. montana Azara alpina Dioscorea brachybotrya Myrceugenia chrysocarpa Berberis serrato-dentata

5 2 1 1 + ·

· + 3 · · · ·

5 2 · 2 2 ·

1 1 + 2 + · ·

Differential taxa of subass. drimydetosum andinae

Araucaria araucana Ribes nitidissimum Berberis montana Uncinia negeri Vicia nigricans Alstroemeria aurea Olsynium scirpoideum luridum Anemone antucensis Valeriana laxiflora Blechnum microphyllum Senecio pilquensis Carex lateriflora Calceolaria filicaulis Uncinia scabriuscula Chiliotrichum rosmarinifolium Festuca scabriuscula

Characteristic taxa of Association & Alliance

Column number Altitude (m.a.s.l.) Slope (º) Aspect Canopy cover (%) Heigh of tree layer (m) Plot area (m2) Number of taxa

5 2 + 1 1 ·

2 + 1 · + · ·

2 1 + · 2 4 · · · 1 + · + · · ·

4 1470 20 SE 100 15-20 200 20

5 1 1 1 · ·

2 1 · 2 1 · ·

2 2 · 1 + · · · 1 · + · · · · ·

5 1380 10 NW 95 15-18 300 22

5 3 2 2 1 1

4 · 3 · · · +

2 r + + · 1 1 1 + · · · · · · ·

6 1330 15 SSE 90 15-22 450 24

5 2 3 2 2 1

3 + + · · · 1

2 · + + · 2 1 + + + · · · · · ·

7 1400 10 NE 100 22-26 500 25

5 3 2 2 1 1

4 + · · · · +

1 1 1 1 · 2 + 2 · · · · · · · ·

8 1420 20 SE 100 20-22 500 24

5 2 2 2 1 +

2 + + · · · ·

1 1 + 1 · · 1 1 · · 1 · · · · ·

5 1 + 1 1 +

1 1 · 2 · 3 ·

2 1 1 1 + 1 · 1 1 · · + · · · ·

5 2 + 1 + ·

· + 5 1 · · ·

2 · + 1 1 2 · r + · · · · · · ·

5 3 1 2 1 ·

· · 4 · · · ·

2 + + + + 1 · 1 2 · · · 1 · · ·

5 2 2 3 3 ·

1 + · 2 + · ·

3 1 1 + 1 3 · · · 1 1 · · · + ·

5 3 1 1 1 +

3 · 4 + + 2 ·

2 + + · · · + · + 1 + · · · · ·

5 3 2 2 1 +

3 · 2 + · 2 ·

1 1 + · · + · · + 2 · · · · · ·

5 2 1 1 1 ·

+ + 3 r + 2 ·

2 1 · r + · · · · 1 1 · · · · ·

9 10 11 12 13 14 15 16 1430 1450 1590 1585 1610 1330 1410 1390 5 20 15 20 35 15 35 20 WSW S NNW SE WNW WSW S SE 90 95 90 85 100 85 90 90 20-25 16-18 14-25 20-30 12-22 15-25 15-22 15-20 300 300 600 400 400 500 200 300 21 31 22 23 21 24 20 22

5 2 2 2 2 ·

+ + 1 + + · ·

1 1 + · 1 3 · · · + 1 · · · · ·

17 1495 25 SE 100 20-26 300 23

4 2 2 1 · 2

1 + · · · · ·

2 + + 1 · · 1 · · · · 3 · · · ·

18 1320 5 140 17

5 2 3 1 1 1

1 + · · · · +

+ + + 1 · · 1 · · · · 2 · + · ·

19 1410 10 S 150 23

4 2 2 1 · 1

+ + · · · + ·

+ 1 + 1 · · 1 · · 1 · 3 · · · ·

20 1360 15 S 50 19

4 1 1 2 + ·

+ + · · · · ·

2 1 · 1 · · 1 · · · · 1 · 1 · ·

21 1360 5 S 140 23

5 4 + 2 1 ·

· · · · · · ·

2 1 2 · 1 · r 1 1 · · · + · · ·

22 1585 35 SSE 80 16-25 250 24

5 3 2 2 1 1

· + · · · · ·

1 2 + + · · 1 · · 1 (+) r + · · ·

23 1570 30 SSW 95 12-16 200 23

5 2 1 2 2 +

· · · · · · ·

+ 2 · + · · + · 1 · · 1 · · · ·

24 1590 45 SSW 90 10-18 150 20

2 4 1 2 · ·

· · · · · · ·

5 1 · · · · · · · 1 1 · 1 · 1 1

5 4 1 2 2 ·

· · · 1 · · ·

2 1 1 · 2 1 · · · · 1 · · · 1 ·

5 3 · 1 1 ·

· · · · · · ·

3 + 1 · 1 4 · · 1 · + · + · · +

5 2 + · · ·

· · · · · · ·

2 1 · · · · · · · 1 · · · · · ·

25 26 27 28 1660 1650 1615 1625 25 30 40 10 NNW W W NE 90 95 95 100 15-20 10-18 15-17 3-5 150 300 800 200 17 19 22 11

Table 3: Chilean-Argentinian lenga-pehuén-forests of Carici lateriflorae-Araucarietum araucanae subass. drimydetosum andinae (rel.1-21) and subass. araucarietosum araucanae (rel. 22-28).


r · · · · · · · · · ·

· r · · · 1 · · r · ·

· · 1 + · · · 2 · · ·

· · + · · · · · · · · · 1 · · · ·

· 1 · +

1 · · · · · + · · · ·

· · · · · ·

2 1 · · · · ·

2 1 · · · r

· 1 + · · · ·

· · + · · ·

3 1 · + · · ·

· · · · · ·

3 + · · · · · · · 2 · · ·

+ · · + 2 · · 1 · · · · ·

1 r · · · + · 1 r · · · ·

1 r · · · + ·

1 · · · · 1 · · · · ·

1 + · · · ·

1 + · · · · ·

+ r · · · 1 1 · · · ·

1 · · · · ·

· 1 r · · · ·

+ · · · · · · · r · ·

+ 1 + · · ·

1 + + · · · ·

+ + + · · + + · · r ·

· + · + + ·

+ + + · · · ·

· + · · · · · · · · ·

1 1 · · · ·

1 1 1 · · · ·

· r · · · · · · · · 1

· · · · · ·

1 · · · · · ·

· + · · · · · 1 · · ·

· · · · · ·

1 · + · · · ·

· · · + 1 · · · · · ·

· · · · · ·

1 · · · · · ·

· · r · 2 · · · · · ·

· · · · · ·

2 · · · 2 · ·

· · · 1 1 · 1 · · · ·

· · · · · ·

1 + · · · · ·

r · 1 · · + · + · · ·

· · · · · ·

+ · · · · · ·

· · · 1 1 · · · · · ·

· · · · · ·

· + · · + · +

1 · · + 1 2 · · · · ·

· · · · · ·

1 · · · · · ·

· · + 1 1 · · · · · ·

· · · · · ·

1 · · · · · +

+ · + 1 1 · + · · · ·

1 · 1 + 2 +

+ 1 + · · · ·

· 1 · · · · · · + · +

· · · · · ·

1 1 · · · · ·

r + · + 1 + · · · 1 ·

· + · · · ·

· · · · · · ·

1 · 1 1 1 · · · · 1 ·

· · 3 · · ·

1 · · + · · ·

· + · + · · · 1 · · ·

· · 1 · · ·

1 · · + · · ·

· 1 + · · · · · · · ·

· · · · · ·

2 1 1 · · + ·

· 1 · · · · · · · · ·

· · 2 · · ·

1 · · · · · ·

· + 1 + · · · · · · ·

1) 38º 38’ 17”// 71º 36’ 17”· 2) 38º 26’ 04”// 71º 27’ 34”· 3) 38º 38’ 05”// 71º 36’ 10”· 4) 38º 37’// 71º 36’· 5) 38º 41’ 55”// 71º 48’ 40”· 6) 39º 07’ 45”// 71º 42’ 18”· 7) 39º 07’ 36”// 71º 42’ 20”· 8) 39º 07’ 26”// 71º 42’ 19”· 9) 39º 07’ 40”// 71º 41’ 53”· 10) 38º 42’ 00”// 71º 48’ 38”· 11) 38º 25’ 51”// 71º 26’ 01”· 12) 38º 26’ 03”// 71º 26’ 14”· 13) 38º 37’ 46”// 71º 36’ 03”· 14) 38º 12’ 35”// 71º 43’ 56”· 15) 38º 12’ 35”// 71º 43’ 44”· 16) 38º 12’ 42”// 71º 43’ 56”· 17) 38º 38’ 09”// 71º 36’ 08”· 22) 38º 25’ 37”// 71º 27’ 50”· 23) 38º 42’ 10”// 71º 48’ 11”· 24) 38º 42’ 20”// 71º 48’ 07”· 25) 38º 37’ 20”// 71º 35’ 58”· 26) 38º 37’ 32”// 71º 36’ 05”· 27) 39º 00' 03''// 71º 22' 30''· 28) 38º 37’ 10”// 71º 35’ 50”.

Localities: all relevés from La Araucanía region (CHI) except column 27 from Neuquén province (ARG). From CNP, pathway to Sª Nevada (rel·1, 3, 4, 13, 17, 25, 26, 28). From CNP, Llaima Refuge area (rel·5, 10, 23, 24). From Alto Las Raíces-Lonquimay (rel 2, 11, 12, 22).·From TNP, Laguna Verde area (rel·14, 15, 16). From HNP, between Laguna Verde and Laguna Los Patos (rel 6, 7, 8, 9). From FINCKH (1996) in VNP, Table1/2, Group 10, rel·406, 533, 408, 535, as Macrachaenio-Nothofagetum, variante mit Araucaria araucana (rel·18, 19, 20, 21).

Taxa present in only 1 relevè: Characteristic taxa of Wintero-Nothofagetea: Azara lanceolata + in 14; Desfontainia spinosa + in 6. Companion taxa: Bromus araucanus 1 in 2; Baccharis gr· patagonica + in 5; Carex aphylla: + in 27; Diplolepis descolei: r in 27; Gaultheria pumila + in 21; Hieracium sp· + in 24; Hypochaeris aff· palustris + in 18; Hypochaeris tenuifolia + in 21; Lycopodium paniculatum 1 in 14; Nertera granadensis + in 21; Ourisia alpina 1 in 15; Poa tristigmatica: + in 27; Senecio subpubescens + in 26; Viola maculata: + in 27·

Acaena ovalifolia Cardamine aff. cordata Gaultheria poeppigi Uncinia chilensis Lathyrus subandinus Stellaria arvalis

Companion taxa

Myoschilos oblonga Osmorhiza chilensis Ribes magellanicum Berberis microphylla Embothrium coccineum Polystichum plicatum Ovidia andina

Nothofagetea pumilionis-antarcticae & Wintero-Nothofagetea shared taxa

Codonorchis lessoni Gavilea lutea Escallonia alpina Perezia pedicularifolia Macrachaenium gracile Lagenophora hariotii Lycopodium magellanicum Valeriana lapathifolia Senecio prenanthifolius Ranunculus peduncularis Trisetum cernuum



It is interesting to note this modification in the lengapehuén forests in the lower level of the orotemperate belt, as a similar phenomenon occurs in pure lenga forests as explained in the section below. As highlighted earlier, the forests of Carici lateriflorae-Araucarietum, when combined with other broadleaved trees such as Nothofagus dombeyi and understory plants of WinteroNothofagetea, are interpreted as indicators of the supratemperate bioclimatic belt, thus justifying their classification as an/other different association/s; this criterion can be applied to the discrete populations of lengapehuén existing in the part of the mountain range on the coast of Chile known as Nahuel Buta, whose phytosociological position should be considered different from that of Carici lateriflorae-Araucarietum. ● Variability: orotemperate lenga-pehuén forests at a higher altitude Another interesting modification can be seen in lenga-pehuén forests at their upper altitudinal limits, not so much because they attain such extremely rigorous altitudes, but because they contact and overlap with the high-Andean supra-forest communities dominated by nanocamephytes and hemicryptophytes adapted to the regular deposits of volcanic ash. This type of vegetation is ideally represented by the class Quinchamalio-Pernettyetea described by FREIBERG (1985) with relevés in the volcanic areas of Chile in the regions of La Araucanía and Los Lagos. The most characteristic species that mark this trend are Senecio chionophilus (along with their very similar S.chilensis), Quinchamalium chilense, Gaultheria pumila, Poa obvallata, Anthoxanthum juncifolium, Adesmia emarginata, Carex aphylla and Sisyrinchium pearcei. (Figure 3)

Figure 3. Orotemperate lenga-pehuén forests reaching 1.600 m asl along the Lonquimay route; Alto Las Raíces (La Araucanía Region, CHI).

In VNP, FINCKH (1996) collected 15 relevés from this type of forests which he gave the consideration of an independent association designated Festuco scabriusculae-Nothofagetum pumilionis; as can be seen in Table 1 (col.9), these forests contain these aforementioned differential species, and have an abundance of Festuca scabriuscula –which is a more heliophilous than nemoral species–, but still include a significant number of cha-

racteristic species of Nothofagetea pumilionis-antarcticae, in addition to the constant presence of the same dominant trees. We therefore concur with the criterion of ESKUCHE (2002: 18) in questioning the consideration of an independent association under the name “Festuco scabriusculae-Nothofagetum pumilionis” and propose for these formations the treatment of Carici laterifloraeAraucarietum subass. quinchamalietosum chilensis (Finckh 1996) subassociatio nova hoc loco (lectotypus FINCKH 1996: Tab. 1/ Teil 2/Gr. 14, rel. 318). As a result of volcanic activity, which causes the periodic deposit of ash over areas of varying extensions and promotes a dynamic of degradation of the forest stands and the advance of Quinchamalio-Pernettyetea, FINCKH (op. cit.) highlighted a “Community of Empetrum rubrum and Araucaria araucana” that would play a dynamic role in the recolonisation of trees on pioneering camephytic formations, and therefore be a precursor community of Carici-Araucarietum subass. quinchamalietosum chilensis. At least this was observed in the VNP and the eight relevés included in Table 1, col. 8. A similar phenomenon was noted by FREIBERG (1985) in volcanic areas in the region of Los Lagos and which also has significance for the variability of lenga forests in these territories. Pure lenga forests Throughout La Araucanía and Neuquén there are also stands of pure lenga forests that tend to prefer southern exposures and hollows with more intense and longer snow coverage, as opposed to the heliophilous formations of lenga-pehuén. OBERDORFER (1960) first proposed the name Anemono antucensis-Nothofagetum pumilionis for these pure lenga forests, although to define them he proposed a table that contained his own relevé and another seven from previous authors such as SKOTTSBERG (1916), REICHE (1907) and ROIVAINEN (1954); this sample of eight relevés covered an extremely broad area of territory from the Termas de Chillán to Punta Arenas (from parallel 36º to 53º south). With the syntaxonomical synthesis published by HILDEBRANDVOGEL ET AL. (1990) this association became easier to interpret, as it was placed in a suballiance with a more northerly and continental biogeographic significance – compared to other Chilean associations which were somewhat more southerly and hyperhumid– and clearly separated the lenga forests in the VP from those of the MP. The possible lectotypus for this association from among the original relevés, according to the interpretation of HILDEBRAND-VOGEL ET AL. (op. cit.), would also come from the VNP in the same territory in which Carici lateriflorae-Araucarietum was described: Oberdorfer, op. cit., Tb. 38, Auf. 231. This relevé, as the only candidate for the lectotype for the association, is highlighted in Table1 in col.12. Our current interpretation of the ass. Anemono antucensis-Nothofagetum pumilionis recognises a type of lenga forest without Araucaria, which maintains the presence of a series of characteristic species of lenga forests in the area of La Araucanía-Neuquén, such as those highlighted in the first block of columns in the Table 1. Anemone antucensis, Valeriana laxiflora, Vicia nigricans, Alstroemeria aurea and Leucheria thermarum


form the characteristic indicator set for this unit. There are testimonies on the composition of lenga forests of Anemono-Nothofagetum pumilionis both in the Chilean (in the VNP, FINCKH 1996) and the Argentinian part (in the LNP, ESKUCHE 2002), for which we also provide here a series of our own relevés from the LNP (Table 4). From the comparison shown in Table 1, we interpret that Anemono-Nothofagetum pumilionis and Carici lateriflorae-Araucarietum form the set of communities that can be defined by a certain continentality such as the one that HILDEBRAND-VOGEL ET AL. (op. cit.) described under the suballiance Vicio-Nothofagenion pumilionis. To this aspect of “continentality” we should also add another bioclimatic character, namely the sub-Mediterranean influence; both the Chilean lenga forests in La Araucanía and the Argentinian ones included in Anemono-Nothofagetum pumilionis –as they are in a temperate territory– are very near the boundary with the Mediterranean bioclimate, and this influx must be the one that marks the floristic differences between these lenga forests and those in Chilean territory in the regions of Los Ríos, and particularly Los Lagos (Figure 4).

Figure 4. Pure lenga-forest of Anemono antucensis-Nothofagetum pumilionis at Cerro Chapelco (NHNP, ARG)

● Argentinian lenga forests in the VP Although the lenga forests in the LNP and in a large part of the NHNP are identified with Anemono-Nothofagetum pumilionis, another type of pure lenga forest was described in the southern sector of the NHNP, and is worth mentioning due to the disappearance of a substantial proportion of the characteristic species of the previous association, and the notable abundance of other more chionophilous hemicryptophytes and geophytes. This situation was highlighted by ESKUCHE (1973) who described for it an association he called MacrachaenioNothofagetum pumilionis which was represented by only three relevés from the area around Mascardi lake (approx. 41º 15’ south). This association was subsequently accepted by HILDEBRAND-VOGEL ET AL. (1990) as a participant in the more continental suballiance (Vicio-Nothofagenion pumilionis) and its presence in Chile was confirmed in the VNP by FINCKH (1996). ESKUCHE (2002) subsequently published numerous relevés from more diverse points of the NHNP which he included in the same association, thus providing more information

25

on its floristic content, while incurring in certain nomenclatural inaccuracies that should be corrected following the ICPN, as will be discussed below. All these relevés are shown in Table1 (cols.18,19 and 20). In our interpretation, this Macrachaenio-Nothofagetum pumilionis is representative of Argentinian lenga forests with a greater oceanic influx throughout the NHNP and which can be identified in the latitudes to the south of Nahuel Huapi lake in the elevations nearest Chilean territory. Table 4 includes four of our own relevés that can be identified with MacrachaenioNothofagetum pumilionis on the Argentinian slopes of Cerro Tronador. It is worth noting in Table 4 that a more ombrophilous Macrachaenio-Nothofagetum can be detected at practically the same altitude near the border with Chile; and an Anemono-Nothofagetum within barely 20’ degrees of variation in longitude towards the east (comparing relevés 24-27 with group 16-23, in Table 4). Even in the Cerro Catedral itself where we identified this Anemono-Nothofagetum, ESKUCHE (2002) inventoried some lenga forests which showed a greater poverty of species in the alliance and the order, and which he highlighted due to the presence of Senecio argyreus, Senecio linariifolius, Bromus catharticus and Festuca pallescens. He gave it the category of new association with the name of Senecioni linariifolii-Nothofagetum pumilionis; its floristic content is shown in Table1 (col. 17). As we have personally studied lenga forests in this massif and have verified that this floristic composition is the exception rather than the norm, we consider these relevés of ESKUCHE (op. cit.) to be a “variant of Festuca pallescens” but of Anemono-Nothofagetum, whose ecological significance we interpret as a certain degradation and aperture of the tree canopy with the penetration of species from more heliophilous supraforest communities (such as F. pallescens) or from the shrubby edge (such as Senecio linariifolius), or even as a certain degree of anthropisation (the presence of Bromus catharticus). The overview of the Argentinian lenga forests in the VP can be completed with the association described by ESKUCHE (2002) in clearly more southerly latitudes: this author defined an ass. Chiliotricho rosmarinifoliiNothofagetum pumilionis in the tributaries of General Vintter lake (43º 50’ south) which is noteworthy more for the loss of characteristic species of Vicio-Nothofagion and Adenocaulo-Nothofagetalia than for the incorporation of his own set of typical nemoral species. However he attributes to it certain differential species such as Anthoxanthum redolens, Senecio patagonicus and Leucheria cf. magna which, together with the constant presence of Chiliotrichum rosmarinifolium, have been used by its author to highlight the characteristic floristic combination of this association. ● Chilean lenga forests in the VP There are also continuous formations of lenga forest on the western slope of the Andean mountain range, for which several associations have been proposed in essays by different authors. The lenga forest in the territory of La Araucanía can be identified as Anemono antucensisNothofagetum pumilionis, coinciding with the territorial area where it coexists with lenga-pehuén forests. However several studies on the territory have been published

· · 1 + · · · · + · · · ·

Nothofagus pumilio Maytenus disticha Adenocaulon chilense Berberis serrato-dentata Perezia prenanthoides Ribes nitidissimum Gavilea lutea Berberis montana Uncinia negeri Blechnum microphyllum Codonorchis lessoni Viola reichei Rubus geoides Senecio prenanthifolius Ranunculus peduncularis

5 2 2 · + 1 1 1 r + 1 1 1 · +

5 3 1 2 1 1 + 1 r · 1 · · · ·

· + 1 + · · + · · · · · ·

5 4 2 3 1 · + + 1 · 1 + · + +

+ · · · · · · 1 · · · · ·

5 2 4 2 1 1 1 1 2 4 + + 1 · ·

· · 1 · · · · · · · · · · 5 4 4 2 1 1 + 2 2 · · + + · ·

· r 1 · · · · · · · · · · 5 3 3 2 + 1 + 1 2 2 1 · 1 · ·

· · 1 · · + · · + · · · · 5 3 3 2 1 1 1 1 1 1 · · · r ·

1 2 · r · · · 1 · · · · ·

1 2 3 4 5 6 7 1650 1560 1510 1580 1570 1570 1570 15 30 10 10 20 10 45 WNW S NW NW W W SW 100 90 100 95 95 90 90 18-22 18-22 20-25 > 20 15-20 18-20 18-22 800 600 400 500 300 300 200 22 19 20 16 17 18 20

Characteristic taxa of Order & Class

Vicia nigricans Valeriana laxiflora Carex lateriflora Calceolaria filicaulis Leucheria thermarum Alstroemeria aurea Chiliotrichum rosmarinifolium Anemone antucensis Uncinia scabriuscula Senecio argyreus Escallonia alpina Lycopodium magellanicum Drimys andina

Differential taxa of associations

Column number Altitude (m.a.s.l.) Slope (º) Aspect Canopy cover (%) Heigh of tree layer (m) Plot area (m2) Number of taxa

5 3 3 3 1 1 1 2 2 · 1 · · · ·

· 1 2 · · · · · 1 · · · · 5 2 1 3 · + · 1 1 · · · · + +

1 · · · · 1 · · + · · · · 5 3 3 3 1 1 · 1 · · · · · + ·

2 2 · + r · + 1 · · · · · 5 3 3 3 1 1 1 2 2 1 · · 1 · ·

· r + · · + · · 1 · · · · 5 1 2 2 1 1 + + · 2 · · · 1 ·

4 · · · · · + · · · · · · 5 1 2 1 1 1 r + · 2 · + · 1 1

2 · · + 3 · · + · · · · · 5 1 1 2 1 · + 1 · 2 · + · · +

1 · · + 2 · · 1 · · · · ·

8 9 10 11 12 13 14 1560 1535 1560 1590 1490 1480 1470 10 10 45 15 35 50 35 SW WNW SW NW WSW S E 90 95 90 85 95 90 90 15-22 20-25 15-20 17-22 15-20 18-20 18-22 400 150 250 400 400 500 400 18 21 21 17 19 23 22

5 + 1 3 + r · + · 1 · 1 · + 1

·

4 · · · 3 · · · · · ·

15 1480 30 ENE 85 17-25 400 20

5 4 2 3 2 + + · + 2 · · · · ·

1 1 · · 1 · · · · · · · ·

16 1630 25 NNE 90 18-25 250 18

5 3 2 3 2 1 r · · 2 · · · · ·

2 2 · · 2 + + · · · · · ·

17 1605 30 NNE 95 18-22 400 19

5 1 4 2 3 1 1 · + · r · · · ·

· · 1 2 2 · · · · · r · ·

18 1585 35 SSE 90 20-25 500 15

5 3 3 1 + 1 1 · · 1 1 · · · ·

· · · + · · + · · · · · ·

19 1590 25 ENE 95 15-20 200 16

5 4 1 3 2 1 1 · · 1 · · · · ·

3 2 · · 1 1 · · · · · · ·

20 1550 25 NNE 85 15-22 250 18

5 2 3 4 4 1 1 · · · 1 · · · ·

· · 1 + + · · · · · · · ·

21 1525 30 SSE 95 22-25 400 13

5 3 + 2 1 · + · 1 1 · · · · ·

1 2 · · 1 2 · · · · · · ·

5 2 1 3 2 2 1 · · · + 1 + · ·

· · 1 + · · 2 · · · · · ·

5 2 3 2 1 2 1 + · 1 1 1 · · ·

· · · · · · + · · 1 + 1 ·

5 4 3 + 1 2 1 + r 1 1 · + · ·

· · · · · · + · · 1 1 2 ·

5 2 3 2 1 2 1 1 1 · 2 2 + · 1

· · · · · · · · · 1 + 2 ·

5 2 2 4 1 2 1 2 · 3 2 2 + · ·

· · · · · · · · · 1 1 · 1

22 23 24 25 26 27 1525 1555 1440 1455 1465 1475 30 35 20 5 10 15 NNE SE SSE WSW S E 95 90 95 95 95 95 15-25 15-20 20-25 18-22 18-22 15-20 250 300 350 300 250 200 17 21 22 20 23 18

Table 4: Argentinian lenga-forests of Anemono antucensis-Nothofagetum pumilionis (rel.1-23) and Macrachaenio-Nothofagetum pumilionis (rel. 24-27)


· · · ·

· · · ·

· · · ·

· · · ·

· · · ·

· · · ·

· + · + · · · ·

+ 1 + · ·

· · · 1 · r · ·

1 1 · · r

1 · r · r · · ·

+ · · 1 ·

· · · · · r · ·

+ · · · · · + · · · r · ·

+ r · · · + · · · · · · ·

+ 1 · · · · 1 · 1 · · · +

+ + · · ·

· · · ·

· 1 · · · + + ·

1 1 · · ·

· · · ·

4 · + · 2 · + ·

1 · + 1 ·

· · · r

1 1 · · · · + +

1 · · · ·

· · + ·

· · · · · · · ·

1 + · · ·

· · · ·

+ · · · · · · ·

+ + · · 3

1 · · ·

1 · 1 · 2 · · ·

1 · + + ·

+ 1 · ·

1 · · ·

1 · +

2 · 1 2 1

· + · ·

3 · 1 · 1 · · ·

1 · + 1 ·

·

· +

+ 1 1 · · · · ·

1 · + · ·

· · · ·

1 1 1 · · · · ·

1 · + + ·

· · · ·

· · · · · · · ·

3 + · · ·

· · · ·

· + · · · · · ·

1 1 · · ·

· · · ·

1 + 1 · · · · ·

1 1 · · ·

· · + ·

· · · · · · · ·

3 + · · ·

· · · ·

1 · 1 · · · · ·

1 · + 1 ·

· · · ·

+ + · · · · · ·

1 1 · · ·

+ · · 1

· · · + · · · ·

+ 2 · · ·

2 + · ·

· · · 1 · · · ·

· 1 · · ·

+ · · ·

+ · · 1 · · · ·

1 3 · · ·

· · · ·

· · · · · · · ·

2 1 + · ·

· · · ·

1) 40° 12’ 43”// 71° 17’ 46”· 2) 40° 12’ 20”// 71° 18’ 03”· 3) 40° 12’ 15”// 71° 18’ 15”· 4) 40° 12’ 36”// 71° 18’ 20”· 5) 40° 12’ 42”// 71° 18’ 28”· 6) 40° 12’ 47”// 71° 18’ 29”· 7) 40° 12’ 52” // 71° 18’ 35”· 8) 40° 13’ 01”// 71° 18’ 38”· 9) 40° 12’ 59”// 71° 18’ 41”· 10) 40° 12’ 52”// 71° 18’ 36”· 11) 40° 12’ 40”// 71° 18’ 26”· 12) 40° 45’ 13”// 71° 36’ 23”· 13) 40° 45’ 26”// 71° 36’ 21”· 14) 40° 45’ 26”// 71° 36’ 17”· 15) 40° 45’ 18”// 71° 36’ 13”· 16) 41° 10’ 44”// 71° 27’ 40”· 17) 41° 10’ 45”// 71° 27’ 34”· 18) 41° 10’ 50”// 71° 27’ 28”· 19) 41° 10’ 55”// 71° 27’ 33”· 20) 41° 10’ 58”// 71° 27’ 26”· 21) 41° 10’ 53”// 71° 27’ 23”· 22) 41° 10’ 44”// 71° 27’ 25”· 23) 41° 10’ 37”// 71° 27’ 28”· 24) 41° 11’ 42”// 71° 46’ 56”· 25) 41° 11’ 38”// 71° 46’ 59”· 26) 41° 11’ 33”// 71° 47’ 00”· 27) 41° 11’ 26”// 71° 47’ 05”.

Localities: Relevés 1-11: LNP, San Martín de los Andes, Cerro Chapelco; 12-15: NHNP, Villa La Angostura, Cerro Bayo; 16-23: NHNP, Bariloche, Cerro Catedral; 23-27: NHNP, Mascardi, Cerro Tronador; sector Almohadilla.

Taxa present in only 1 relev: Characteristic taxa of Nothofagetea pumilionis-antarcticae: Lagenophora hariotii + in 12; Olsynium scirpoideum subsp. luridum + in 24; Uncinia tenuis + in 3. Companion taxa: Berberis microphylla + in 19; Cortaderia pilosa + in 19; Galium hypocarpium + in 10; Gaultheria poeppigi + in 12; Geranium sessiliflorum r in 1; Gnaphalium sp. + in 16; Gramineae·1 in 26; Hieracium sp.·r in 24; Marsippospermum philippi + in 23; Poa pratensis 1 in 9; Poa sp. + in 26; Ranunculus aff. acaulis 1 in 23; Senecio subpubescens 1 in 25; Senecio triodon + in 26.

Acaena ovalifolia Poa obvallata Bromus coloratus Perezia foncki Stellaria arvalis Hypochaeris arenaria Rumex acetosella Senecio angustissimus

Companion species

Osmorhiza chilensis Myoschilos oblonga Polystichum plicatum Ribes magellanicum Chusquea grex culeou

Nothofagetea pumilionis-antarcticae & Winthero-Nothofagetea shared taxa

Macrachaenium gracile Valeriana lapathifolia Trisetum cernuum Perezia pedicularidifolia


Column number 1 1310 Altitude (m.a.s.l.) 20 Slope (º) SW Aspect 95 Canopy cover (%) 18-20 Heigh of tree layer (m) 250 Plot area (m2) 23 Number of taxa Characteristic taxa of Association & Alliance Drimys andina 2 Escallonia alpina 1 Valeriana lapathifolia · Ranunculus peduncularis + Ovidia andina 2 Chusquea montana · Uncinia scabriuscula 1 Gunnera magellanica · Characteristic taxa of Order & Class Nothofagus pumilio 5 Adenocaulon chilense 4 Viola reichei 3 Maytenus disticha 3 Macrachaenium gracile · Ribes nitidissimum 2 Berberis montana 1 Blechnum microphyllum 3 Uncinia negeri 1 Perezia prenanthoides · Rubus geoides 2 Codonorchis lessoni 2 Berberis serrato-dentata · Senecio prenanthifolius · Olsynium scirpoideum subsp. luridum 1 Lagenophora harioti + Perezia pedicularidifolia 1 Gavilea lutea · 3 2 · · 1 · + · 5 2 2 4 · 1 1 1 1 · + 1 · · · 1 + ·

5 · · + · · + ·

5 1 1 2 · · 1 2 1 · · + · + · · · ·

5 2 + 2 · 1 + 1 1 · + + · · · · r ·

5 · · · + · + · 5 3 2 4 1 1 1 2 + · · 1 · · 1 1 · 1

3 2 · r 1 · 1 · 5 + 1 2 2 2 1 2 1 · · · · 1 + · · ·

4 1 · + 1 · · 1 5 2 2 2 3 1 1 2 1 · + 1 · 1 1 · · ·

2 1 · + 1 · + · 5 1 2 + + · · · · + · · + · · · · ·

+ · 2 + + 4 · · 4 1 2 2 1 · · + · 1 · · · · · · · ·

2 · 2 + + 4 · · 5 1 + 1 2 · · · · 1 · · · · · · · ·

1 · 4 + · 2 · · 5 1 2 · 1 · · · · 1 · · · · · · · ·

+ · 4 · · 1 · · 5 1 2 2 2 · · · · 1 · · · · r · · ·

3 · 4 · + 3 · ·

2 3 4 5 6 7 8 9 10 11 12 1315 1275 1235 1320 1275 1200 1250 1280 1305 1365 1365 20 20 15 15 15 35 20 15 35 35 30 NW S WNW WNW NW SSE S S SSW SSE S 90 95 100 90 90 95 90 70 100 90 85 12-20 12-18 15-20 15-18 12-18 20-22 22-25 10-18 15-22 20-25 15-20 200 150 300 200 150 250 400 200 200 150 200 19 22 20 23 23 25 16 16 15 12 13

5 2 1 4 + 2 1 1 + · 1 · · · · + · ·

5 + · · · 2 · · 5 + 1 3 · 1 1 · + · 1 · · · · · r ·

5 · · · · · · · 5 + 1 4 · 2 1 + + · + · · · · · · ·

5 · · · + + · · 5 1 1 2 4 + 1 · · r + 1 + · · · · ·

2 1 4 · · · · 2 5 2 2 2 2 1 + · + · 1 1 · · · + · ·

3 1 · · · · · 2

4 1 1 4 1 + + 2 + + · · 2 1 · · · ·

4 1 2 1 · · · ·

13 14 15 16 17 18 1060 1150 1140 1190 1270 1285 15 20 20 20 30 25 SSW WSW WSW SE NNW NE 95 95 95 95 90 95 14-20 12-16 15-20 15-20 12-16 5-20 400 200 400 600 300 200 21 18 19 23 19 21

Table 5: Chilean lenga-forests of Macrachaenio gracilis-Nothofagetum pumilionis subass. ovidietosum andinae.

4 2 + 4 + + + 1 · r · · 2 + · · · ·

3 1 2 + · 2 · ·

19 1320 35 NNE 95 20-23 120 21

4 1 + 4 + + · 1 · r · · 2 + · · · r

4 1 3 · · · · ·

20 1305 30 NNE 90 20-22 150 17

5 1 1 3 2 · · 2 + · · · 1 · · · · 1

4 · 1 · · · · ·

21 1300 15 NNE 95 20-22 200 15

5 2 1 3 1 + 1 2 + 1 · · 1 1 + · · ·

3 2 3 + · · · ·

22 1260 30 ENE 95 20-25 200 21


· 2 · 1 1 · 1 · · · · + · · · r · · ·

· · + 1 · + · · r · · 1 · 1 · · · · ·

1 · · · · · 1 ·

· 2 · · · + · · · ·

·

1 · · · · · · ·

· 2 · · · + · · r ·

·

1 · · · · · 1 ·

· + · · · + · · · ·

·

1 · · · · · · ·

· + · + · · · · · ·

1

· · · · · · · ·

· · · + · · · · · ·

+

· + · · · · · ·

· + + · · · + · + ·

·

· 1 1 1 · · · +

· · · · · · 1 · · +

·

· 1 + · · · · r

· · + · · · + · · +

·

1 + · · · · · ·

2 + + 1 1 · · · · ·

·

1 + · · · · · ·

1 · · · 1 · · · · ·

·

1 · · · · · · ·

1 2 + · · + · · · ·

·

1 · · · · · · ·

1 + · + · · · · · ·

·

1 · · · · · · ·

1 · + · · · · · · ·

·

· 1 · · · · · ·

+ · + · · · · · · ·

·

1 · · · · · · ·

+ 1 · + · · · · · ·

·

1) 39º 56' 37'' // 72º 04' 21''· 2) 39º 56' 36'' // 72º 04' 17''· 3) 39º 56' 40'' // 72º 04' 27''· 4) 39º 56' 38'' // 72º 04' 36''· 5) 39º 57' 07'' // 72º 04' 04''· 6) 39º 57' 09'' // 72º 04' 09''· 7) 39º 56' 59'' // 72º 04' 21''· 8) 39º 47' 01'' // 71º 40' 50"· 9) 39º 47' 02'' // 71º 40' 53"· 10) 39º 46' 57'' // 71º 40' 56"· 11) 39º 46' 50'' // 71º 40' 58"· 12) 39º 46' 48'' // 71º 41' 01"· 13) 40º 46' 25'' // 72º 10' 16''· 14) 40º 46' 30'' // 72º 11' 54''· 15) 40º 46' 30'' // 72º 11' 58''· 16) 40º 43’ 31”// 71º 55’ 46”· 17) 40º 42’ 54”// 71º 56’ 43”· 18) 41º 04' 34''// 71º 50' 05''· 19) 41º 04' 35''// 71º 50' 09''· 20) 41º 04' 36''// 71º 50' 07''· 21) 41º 04' 34''// 71º 50' 08''· 22) 41º 04' 34''// 71º 50' 03''·

Localities: Relevés 1-7: Choshuenco Volcano, Los Ríos Region (CHI); 8-12: Carirriñe Pass (CHI-ARG border); 13-15: Cº Antillanca, Los Lagos Region (CHI); 16-17: Puyehue Pass (CHI-ARG border); 18-22: VPRNP, Cerro Riggi (CHI-ARG border).

Taxa present in only 1 relevè: Characteristic taxa of Nothofagetea pumilionis-antarcticae: Anemone antucensis r in 10; Calceolaria filicaulis + in 3; Carex lateriflora 1 in 7; Lycopodium magellanicum 1 in 21; Trisetum cernuum + in 16. Nothofagetea pumilionis-antarcticae & Wintero-Nothofagetea shared taxa: Asteranthera ovata + in 19; Blechnum magellanicum + in 17; Hymenophyllum sp. + in 13; Nothofagus betuloides 1 in 15; Uncinia tenuis + in 13. Companion taxa: Carex darwinii + in 6; Gaultheria pumila + in 14; Senecio chilensis + in 14.

Senecio acanthifolius · · · · · Nothofagetea pumilionis-antarcticae & Wintero-Nothofagetea shared taxa Osmorhiza chilensis 1 1 1 1 + Ribes magellanicum · + · + · Myoschilos oblonga · · · 1 1 Dysopsis glechomoides · · · · · Blechnum penna-marina 1 + 1 · · Polystichum plicatum · · · · · Embothrium coccineum · · 1 · · Myrceugenia chrysocarpa · · · r · Berberis trigona · · · · · Grammitis magellanica · · · · · Companion taxa Acaena ovalifolia 1 1 1 1 + Gaultheria poeppigi · · · · · Cortaderia egmontiana r + · · · Festuca thermarum · + + 1 2 Poa obvallata + · · · · Perezia foncki · + · · 1 Chrysosplenium valdivicum · · · · · Berberis microphylla · · · · ·



at latitude 39º 40’ and southwards, based on which some associations have been proposed that we consider advisable to re-order. In the first place FREIBERG (1985) proposed a Carici minutissimae-Nothofagetum pumilionis with relevés from lenga forests on slopes of several Chilean volcanoes, although most come from Antillanca (PNP); his proposal was published with a provisional name, although HILDEBRAND-VOGEL ET AL. (1990) subsequently considered it to be one of the recognisable associations in Chilean territory, assimilated to a suballiance Valeriano-Nothofagenion pumilionis. But it was not until later that FLORES-TORO & HILDEBRAND-VOGEL (2006), arguing that this was a nomen nudum, proposed a new name for this association: Drimydo andinae-Nothofagetum pumilionis, choosing as the holotype for the new association an relevé by HILDEBRAND-VOGEL ET AL. (op. cit.), which was the only relevé in this publication that included the presence of Carex minutissima. In our opinion this proposal should be rejected for two reasons: 1) the floristic composition of most of the relevés published under the name of Carici minutissimae-Nothofagetum is identifiable with the association proposed for Argentinian territory as Macrachaenio-Nothofagetum; and 2) the proposal of FLORES-TORO & HILDEBRANDVOGEL (2006) as an independent association is based on a type relevé that is very poor in species for a forest (only14 species), among which there are three erroneous identifications –the most important being that of the supposed Carex minutissima, as noted by ESKUCHE (2002) and recently defended by AMIGO & CASTRO (2015). In our opinion the oceanic lenga forests in Chilean territory from the start of the region of Los Ríos to the northern part of the region of Los Lagos are identifiable with Macrachaenio-Nothofagetum. Our own data from latitude 39º 46’ to the interior of the VPRNP in 41º 05’ have been included in Table 5. ● The identity of Macrachaenio-Nothofagetum pumilionis When the name of Macrachaenio-Nothofagetum pumilionis was first validly published it was done with only three relevés, one of which was already designated “typicum”; the lectotype should thus have been ratified in this table and with this relevé instead of proposing a new holotype (ESKUCHE 2002). There can therefore be no doubt that the name and type should be: Macrachaenio gracilis-Nothofagetum pumilionis Eskuche 1973, lectotypus ESKUCHE 1973: Table 1, rel. M4 (page 72) in hoc loco. Indeed the identification of this relevé “typicum” served as the basis for the interpretations made by HILDEBRAND-VOGEL ET AL. (1990) to frame it in a specific sub-alliance, and by FINCKH (1996) when he identified it as present in his study territory in the VNP. The second publication with Argentinian relevés (ESKUCHE 2002; see Table 1, cols. 18 and 19) characterised a subassociation escallonietosum alpinae –also invalid due to the application of Art. 16 of the ICPN– with which its author wished to denote a slightly greater oceanic influence (the relevés in this subassociation came from areas bordering Chile). Comparing the relevés of ESKUCHE (op. cit.) with those published previously, we can assume that the Macrachaenio-Nothofagetum pumilionis can be interpreted as the association

found in Chilean territory in the regions of Los Ríos and the northern part of the region of Los Lagos. Our own relevés shown in Table 5 serve as an example (col. 21, Table 1). The syntaxonomical rationale we propose for the interpretation of this association and its modifications is as follows: A) The nomenclatural type of the association corresponds to a relevé from the more orophilous part of the lenga forests, without reaching the band of dwarf lenga forest; it has an absence of shrubs with greater biomass (Drimys andina, Escallonia alpina and Chusquea culeou) and presents some species that are more typical of the upper edges of the forest such as Senecio argyreus, Chiliotrichum rosmarinifolium and Poa obvallata. This is the typical association. B) There is another version of this association at slightly lower altitudes characterised by the presence of Drimys andina, Chusquea culeou and several herbaceous species. Only two relevés were assigned to this lenga formation by ESKUCHE (1973), based on which we accept the name of Macrachaenio-Nothofagetum pumilionis subass. chusqueetosum culeou Eskuche 1973, lectotypus ESKUCHE 1973: Tab. 1, rel. M5 in hoc loco. Most of the relevés subsequently collected by its author (ESKUCHE 2002) correspond to this subassociation, where the common species he recognised as present were Valeriana lapathifolia, Ribes nitidissimum (sub R. densiflorum) and several species of the order and the class, in addition to the constant Andean canelo (which in the relevés in this publication appeared as D. winteri “strauchförmig”, and its interpretation as Drimys andina is therefore indisputable). A number of relevés in this work (ESKUCHE op. cit., Table 4) can be classified as a “Variant of Escallonia alpina”, such as those from Puyehue Pass on the border between Argentina and Chile. C) As the whole set of Chilean lenga forests in the territory in the region of Los Ríos and the northern part of Los Lagos shows a clear and even more pronounced oceanic influx, it is proposed here to differentiate these forests on the western Andean side as Macrachaenio-Nothofagetum pumilionis subass. ovidietosum andinae subassociatio nova in hoc loco. (holotypus rel. 7, Table 5). In addition to the common presence of the shrubs Escallonia alpina and Drimys andina, good differential species of the shrubby subassociation are Ovidia andina and the hemicryptophyte Ranunculus peduncularis, to which can be added the occasional presence of Nothofagus betuloides (these last three are absent from the Argentinian relevés of ESKUCHE op. cit.) (Figure 5). Also at lower altitudes in the subass. ovidietosum andinae there is a massive presence of colihues –the bambusoideae in the genus Chusquea– which can attain significant coverage in the understory and lead to a lower diversity of herbaceous species, and even affect the regeneration dynamic of lengas (VEBLEN 1982). A syntaxonomical modification with the rank of “variant of Chusquea montana” could be considered for these situations, of which there are numerous examples (see Table 5, rel. 8-13); however as noted in the case of the variability of lenga-pehuén forests, it is also necessary to


31

determine whether the Chusquea that appear on the Argentinian side are all Ch. culeou, or whether on the other hand they can be identified with Ch. montana, as appears to be the most frequent case in the lenga forests on the Chilean side, as indicated by AMIGO & CASTRO (2015).

Figure 5. Lenga-forests of Macrachaenio gracilis-Nothofagetum pumilionis subas. ovidietosum andinae show a dense shrub understory; southern slope of Volcano Choshuenco (Los Ríos Region, CHI)

Figure 6A. Ultrahyperhumid lenga-forest of Senecioni acanthifolii-Nothofagetum pumilionis; in the foreground, Senecio acanthifolius. Cerro Riggi, VPRNP (Los Lagos Region, CHI).

According to our proposed interpretation, most of the lenga forests in Chilean territory to the south of latitude 39º 45’ to latitude 41º south –where we have our own data– correspond to Macrachaenio-Nothofagetum pumilionis subass. ovidietosum andinae; the minority proportion of lenga forests represented by relevés lacking in differential species (Escallonia alpina, Ovidia andina, Nothofagus betuloides) are at least identifiable with the subass. typicum. With this we defend the superfluous and expendable character of the names proposed for many of these orotemperate forests in the VP: Carici minutissimae-Nothofagetum pumilionis, Drimydo andinaeNothofagetum pumilionis and Valeriano lapathifoliaeNothofagetum pumilionis should be downgraded to synonyms. ● Ultrahyperhumid lenga forests In his study on the vegetation in the VPRNP, VILLAGRÁN (1980) proposed a different association for the lenga forest, namely Senecioni-Nothofagetum pumilionis. Although herrelevés came from the same southern latitude as those of ESKUCHE (1973) when he described Macrachaenio-Nothofagetum –as they came from the western slope of the Andes– their floristic composition revealed a greater degree of moisture due to the presence of Senecio acanthifolius, Gunnera magellanica, Acaena magellanica and Ranunculus chilensis. We interpret this as an ultrahyperhumid association encompassing the lenga forests in the rainier areas of the VP and which extend southwards from this VPRNP; all the testimonies attributable to this association come from the Chilean territory. Its composition is shown in Table 1 col. 28, along with other relevés of FLORES-TORO & HILDEBRAND-VOGEL (2006) published under the same name – although not all of them are correctly interpreted (Table 1, cols. 30 and 31)–, and some of our own relevés (Table 1, col. 29) made in the same classic locality as Villagrán: Cerro Riggi.

Figure 6B. At the same latitude but 23’ to the East, lengaforest of Anemono antucensis-Nothofagetum pumilionis at Cerro Catedral, NHNP (Río Negro Province, ARG).

The name proposed by VILLAGRÁN (op. cit.), Senecioni-Nothofagetum pumilionis (sic) was published as a “provisional name” without specifying which species of Senecio gave the name to the association (two appeared in the composition: S. acanthifolius and S. prenanthifolius); based on this FLORES-TORO & HILDEBRANDVOGEL (op. cit.) assumed it as nomen nudum and proposed a new interpretation under their authorship [Senecioni acanthifolii-Nothofagetum pumilionis Flores-Toro & Hildebrand-Vogel], presenting a nomenclatural type designated by them among their own relevés. This proposal is completely invalid as prior to their work ESKUCHE (2002) lectotypified the association based on relevés of VILLAGRÁN (op. cit.), and the nomenclature was therefore clear: Senecioni acanthifolii-Nothofagetum pumilionis Villagrán ex Eskuche 2002 (Figure 6).

32 J. Amigo & M.A. Rodríguez-Guitián Table 6: Chilean lenga-forests of Senecioni acanthifolii-Nothofagetum pumilionis subass. gunneretosum magellanicae Column number 1 2 3 1390 1415 1455 Altitude (m.a.s.l.) 25 15 40 Slope (º) NE SE SE Aspect 95 90 95 Canopy cover (%) 12-20 15-20 6-8 Heigh of tree layer (m) 200 150 120 Plot area (m2) 27 21 22 Number of taxa Characteristic taxa of Association & Alliance Senecio acanthifolius 1 2 1 Gunnera magellanica 1 1 1 Ribes nitidissimum 1 1 1 Ranunculus peduncularis + 2 1 Valeriana lapathifolia 4 · 4 Acaena antarctica · 1 1 Blechnum microphyllum 1 r · Carex lateriflora r · · Dysopsis glechomoides · 2 1 Caltha appendiculata · 1 + Oxalis magellanica · · r Characteristic taxa of Order & Class Nothofagus pumilio 5 5 5 Macrachaenium gracile 2 2 1 Adenocaulon chilense 3 2 r Maytenus disticha 2 + 2 Perezia prenanthoides + + 1 Berberis montana 1 + + Rubus geoides 1 · 1 Escallonia alpina 1 · 2 Codonorchis lessoni + 1 · Perezia pedicularidifolia 1 r · Viola reichei 1 · 1 Lagenophora hariotii + · 1 Lycopodium magellanicum 1 · · Gavilea lutea 1 · · Berberis serrato-dentata 1 · · Nothofagetea pumilionis-antarcticae & Wintero-Nothofagetea shared taxa Myoschilos oblonga + r · Osmorhiza chilensis + 1 · Companion taxa Gaultheria poeppigi + · · Acaena ovalifolia + · · Perezia foncki · 2 + Marsippospermum philippii · 1 +

4 1405 15 SSE 90 15-20 150 22

5 1400 25 S 95 15-20 250 22

6 1375 10 SSE 100 10-22 150 17

7 1230 30 SSW 90 20-22 400 20

1 + 1 + · 1 + r · · ·

1 1 1 + 1 + · 1 · · ·

1 1 1 1 2 · 2 · · · ·

+ 2 + · 1 · · r 1 · ·

5 3 3 2 2 1 2 2 + 2 + · · · ·

5 3 2 3 2 1 1 2 + 2 · · + · ·

5 3 + 1 + · 1 1 · · · · + + ·

5 5 1 · · 1 + · 1 · 1 + · r +

+ ·

+ ·

· ·

· 1

+ · 1 ·

+ · · ·

+ + · ·

r · · ·

Taxa present in only 1 relevè: Characteristic taxa of Nothofagetea pumilionis-antarcticae: Olsynium scirpoideum subsp. luridum + in 1; Uncinia negeri + in 5; Nothofagetea pumilionis-antarcticae & Wintero-Nothofagetea shared taxa: Blechnum penna-marina + in 7; Drimys andina + in 7. Companion taxa: Cortaderia egmontiana + in 2; Misodendron sp. 1 in 4; Ourisia breviflora subsp. uniflora + in 3; Poa obvallata + in 5. Localities: All relevés from Cerro Riggi (VPRNP, Los Lagos-CHI) 1) 41° 04' 39''// 71° 50' 12''. 2) 41° 04' 44''// 71° 50' 20''. 3) 41° 04' 41''// 71° 50' 21''. 4) 41° 04' 45''// 71° 50' 16''. 5) 41° 04' 40''// 71° 50' 13''. 6) 41° 04' 39''// 71° 50' 10''. 7) 40º 42’ 55”// 71º 56’ 06”.

VILLAGRÁN (op. cit.) also distinguished two different aspects within Senecioni acanthifolii-Nothofagetum pumilionis: one at lower altitudes with a dense coverage of Andean canelo which he designated “typicum”; and another which he called subass. gunneretosum magellanicae, in which this shrub practically disappeared. The lectotype chosen by ESKUCHE (op. cit.) is among the first, although in our opinion the association is manifestly more recognisable in its subass. gunneretosum, as the high coverage of Drimys andina offers little opportunity for the low-growing hemicryptophytes that serve as good differential species for Senecioni acanthifolii-

Nothofagetum. As it was not done by either VILLAGRÁN (op. cit.) or ESKUCHE (op. cit.), and although it was proposed by FLORES-TORO & HILDEBRAND-VOGEL (2006) but supported on an invalid name, we propose here the correct typification of this association: Senecioni acanthifolii-Nothofagetum pumilionis subass. gunneretosum magellanicae Villagrán ex Amigo & RodríguezGuitián [lectotypus VILLAGRÁN (1980), Tab. 2, Aufnahme 78] in hoc loco. The floristic set of the association is characterised by Senecio acanthifolius, Gunnera magellanica, Acaena magellanica and Ranunculus chilensis. Table 6 shows


some typical relevés of this subass. gunneretosum magellanicae that we collected in the classic locality of the association: Cerro Riggi in the VPRNP. According to this we consider that other differential species for the subass. gunneretosum are Acaena antarctica and Oxalis magellanica. The distinction between Ranunculus chilensis and R. peduncularis may also have led to a somewhat erroneous interpretation of a characteristic species; in our relevés collected in 2015 we found only R. peduncularis to be participating in these lenga forests.Senecioni acanthifolii-Nothofagetum pumilionis is found predominantly to the south of the VPRNP; FLORES-TORO & HILDEBRAND-VOGEL (2006) contributed a number of relevés from points such as Sierra Escondida (41º 45’), the Hornopirén Volcano (41º 53’), Futaleufú Pass (43º 12’) and Cerro Moraga (43º 21’) which can largely be interpreted as indicative of this association. More to the south of the region of Los Lagos there is scarcely any data to show the complete floristic composition of these forests. We only include here the association described by HILDEBRAND-VOGEL ET AL. (1990) as Berberido ilicifoliae-Nothofagetum pumilionis (col. 34 in Table 1); it points to a clearly impoverished floristic content which can be explained by its 45º latitude south, the participation of Nothofagus betuloides in the tree layer, and the good differential species Berberis ilicifolia which does not reach more northerly latitudes than parallel 40º 10’ (LANDRUM 2003). ● Dwarf lenga forests Various studies on lenga forests in different territories have highlighted the presence of a band of varying widths at their upper altitudal limits comprising dwarf lengas of between barely 1.5 and 3 m in height, the result of the mechanomorphosis exerted by the snow dynamic on specimens of Nothofagus pumilio in the areas most exposed to prolonged snow coverage. ESKUCHE (1973) provided a detailed explanation of the origin of these dwarf forms –popularised in the scientific literature under the German name (krummholz)– and proposed an ass. Pernettyo-Nothofagetum pumilionis to distinguish them, for which he published a single relevé on Argentinian territory whose main characteristic species was identified as “Pernettya cf. pumila”. FINCKH (1996) subsequently took up this idea and combined the relevé of ESKUCHE (op. cit.) with others of his own from the VNP and some from FREIBERG (1985) to compile a set of relevés which he designated “Pernettyo-Nothofagetum pumilionis Eskuche 1973”, assigning Pernettya pumila as the differential species; this set is shown in Table 1, col.7. ESKUCHE (2002) later retracted his initial proposal and chose instead to interpret these situations as subassociations, for which he defined an Anemono-Nothofageteum pumilionis subass. pernettyetosum nanae and another Macrachaenio-Nothofageteum pumilionis subass. pernettyetosum nanae; in spite of the binomen used to name these subass. the author admitted that in both cases he was uncertain as to whether the differential taxon was Pernettya myrtilloides var. nana or Pernettya pumila var. crassifolia ESKUCHE (op.cit.: 26). In our opinion these are normal situations at the upper limit of these Nothofagus pumilio forests, where it

33

is the lengas themselves that act as the edge of the mature forest. The flora beneath these dwarf lengas consists of very few characteristic species of the order or class Nothofagetea pumilionis-antarcticae that may fortuitously survive, along with the presence of species of the communities of supraforest camephytes and hemicryptophytes. The two Pernettya species mentioned by ESKUCHE (2002) belong to these communities, which can be framed in the class Quinchamalio-Pernettyetea, although today they are considered part of the genus Gaultheria (sub Gaultheria poeppigi and G. pumila; see TEILLIER & ESCOBAR 2013). We judge it preferable to interpret these situations with the syntaxonomical rank of “variant”, which allows the distinction of ecological situations that can be differentiated and mapped, but without incurring in the hypertrophication of forest syntaxonomy. Conclusions In the Andean territory in Chile and Argentina between latitudes 38º and 42º south, and over a period of half a century of studies in lenga forests, as many as 15 different associations have been proposed, many of which are supported by relevés that are poor in species or by the presence of taxa typical of dynamic succession stages or the supraforest vegetation with which they contact. To a large degree these situations can be interpreted in terms of lower ranking syntaxonomical units (subassociations, variants) more closely associated with the factors causing this floristic poverty or particular floristic composition. Throughout the whole of the VP there are continuous orotemperate forest dominated by Nothofagus pumilio. This predominance is shared with Araucaria araucana between parallel 38º and 39º 40’ south on both sides of the Andes, and occasionally with Nothofagus betuloides from 40º 40’ on only the Chilean side. When lenga forests are formed with other Nothofagus species that attain the same tree height, this indicates a descent to the supratemperate belt, and the communities formed should preferably be linked to nemoral associations in the class Wintero-Nothofagetea. We propose here to recognise and interpret all lenga forest formations based on the redefinition (including cases of lectotypification) of the four oldest associations described. Another two associations described for territories below parallel 43º which present an impoverishment in the total content of flora and characteristic species are also accepted. Several authors have incurred in inaccuracies or incorrect floristic identification which have led to the perception that there are more differences than identities among lenga forests in different territories. The identification of species of Chusquea genus present in the VP lenga forests still require a more documented analysis. Phytosociological associations –and particularly in the case of climactic communities– must be understood as being associated to a specific territory. This does not imply that after a detailed study of a reduced territory the floristic variations observed within one type of forest should be described as different associations (as in the case of FINCKX 1996). Nor does it appear reasonable to


interpret the presence/absence of a single species as serving to identify an association “A” as opposed to another “B”, in the absence of an ecological-dynamic reason to justify it. This has given rise to a situation where the same three associations of different lenga forests recognised throughout the whole of the Chilean region of Los Lagos are interpreted as being present on the same mountain (the case of FLORES-TORO & HILDEBRAND-VOGEL 2006). Acknowledgements: We would like to thank Dr. Carlos Ramírez and Professor Cristina San Martín from the UACh for their kind welcome and collaboration; Mr Mario Maturana Arévalo and the forest ranger Mr Jeremías Cárdenas Mira in the CONAF 10th Region for allowing us access to the VPRNP; and Eduardo Castro Rodríguez for his photographic skills and assistance in field research. References: Amigo J & Castro E. (2015).- Notas taxonómicas sobre la flora de los bosques orotemplados de la provincia biogeográfica valdiviana. Chloris Chilensis 18 (1). URL://www.chlorischile.cl Amigo J & Rodríguez-Guitián M.A. (2011).- Bioclimatic and phytosociological diagnosis of the species of the Nothofagus genus (Nothofagaceae) in South America. International Journal of Geobotanical Research, 1: 1-20. Amigo J, Rodríguez-Guitián M.A & Ramírez C. (2010).- The lleuque forests of South Central Chile: a phytosociological study and syntaxonomical classification within Southamerican temperate forests. Lazaroa 31: 85-98. Besoaín E. (1985).- Capítulo I, Los suelos. In: Tosso J. (ed.), Suelos volcánicos de Chile. Talleres Gráficos del I.N.I.A: 25106. Santiago. Braun-Blanquet J. (1979).- Fitosociología. Bases para el estudio de las comunidades vegetales. H. Blume Ediciones. 820 pp. Barcelona. Cabrera AL. (1976).- Regiones Fitogeográficas Argentinas. Encic. Argent. Agric., 2ª Edic. II (I): 1-85. Buenos Aires Cabrera AL & Willink A. (1973).- Biogeografía de América Latina. Secretaría General de la Organización de los Estados Americanos, Washington. 117 pp Donoso C. (1998).- Bosques templados de Chile y Argentina. Variación, estructura y dinámica.4ª Edic. Editorial Universitaria, Santiago de Chile: 484 pp. Donoso C (editor). (2006).- Las especies arbóreas de los bosques templados de Chile y Argentina. Autoecología. Marisa Cuneo Ediciones, Valdivia. 678 pp. Eskuche U. (1973).- Estudios fitosociológicos en el norte de Patagonia. I. Investigación de algunos factores de ambiente en comunidades de bosque y de chaparral. Phytocoenologia 1(1): 64-113. Eskuche U. (1999).- Estudios fitosociológicos en el norte de la Patagonia. II. Los bosques de Nothofagion dombeyi. Phytocoenologia 29(2): 177-252. Eskuche U. (2002).- Pflanzensoziologische Untersuchungen in Nordpatagonien. IV. Die Wälder des Nothofagion pumilionis. Folia Botanica et Geobotanica Correntesiana 16: 1-47. Finckh M. 1996. Die Wälder des Villarrica-Nationalparks (Südchile) – Lebengemeinschaften als Grundlage für ein Schutzkonzept. Diss. Bot. 259: 195 pp.

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Wheeler GA. (1989).- The Taxonomy of Carex sect. Aciculares (Cyperaceae) in South America. Syst. Bot. 14 (2): 168-188. Wheeler GA (1997a).- Two new species of Uncinia (Cyperaceae) from austral South America and a first report of U.negeri in Argentina. Hickenia 45(2): 215-222. Wheeler GA (1997b).- Two new species of Uncinia (Cyperaceae) from Chile. Aliso 15: 1-6

APPENDIX I: Taxonomic names cited along the text and tables are in nomenclatural accordance with ZULOAGA except Ribes nitidissimum Neger

ET AL.

2009,

APPENDIX II: Phytosociological typology of the syntaxa cited in the text. NOTHOFAGETEA PUMILIONIS-ANTARCTICAE Oberd. 1960 Adenocaulo chilensis-Nothofagetalia pumilionis Oberd. 1960 em. Hildebrand-Vogel, Godoy & Vogel 1990 Vicio nigrescentis-Nothofagion pumilionis ( Hildebrand-Vogel et al. 1990) all. nova in hoc loco (Typus: Anemono antucensis-Nothofagetum pumilionis) + Anemono antucensis-Nothofagetum pumilionis Oberd. 1960 Variant of Festuca pallescens Variant of Gaultheria pumila + Carici lateriflorae-Araucarietum araucanae Oberd. 1960 araucarietosum araucanae Oberd. 1960 drimydetosum andinae subass. nova quinchamalietosum chilensis (Finckh 1996) subass. nova + Chiliotricho rosmarinifolii-Nothofagetum pumilionis Eskuche 2002 Senecioni acanthifolii-Nothofagion pumilionis all. nova in hoc loco (Typus: Senecioni acanthifolii-Nothofagetum pumilionis) + Macrachaenio gracilis-Nothofagetum pumilionis Eskuche 1973 nothofagetosum pumilionis Eskuche 1973 Variant of Escallonia alpina Variant of Gaultheria pumila chusqueetosum culeou Eskuche 1973 ovidietosum andinae subass. nova + Senecioni acanthifolii-Nothofagetum pumilionis Villagrán ex Eskuche 2002 nothofagetosum pumilionis Villagrán ex Eskuche 2002 gunneretosum magellanicae Villagrán 1980 ex Amigo & Rodríguez- Guitián + Berberidi ilicifoliae-Nothofagetum pumilionis Hildebrand-Vogel, Godoy & Vogel 1990 Violo magellanicae-Nothofagetalia pumilionis (Roig et al. 1985) Hildebrand-Vogel, Godoy & Vogel 1990

Rejected names: Class Nothofagetea antarcticae Freiberg 1986 (Art. 5) Alliance Lagenophoro-Nothofagion pumilionis Oberd. 1960 em. Freiberg 1986 (Art. 5) Alliance Valeriano lapathifoliae-Nothofagenion pumilionis : its nomenclatural Typus (Dombeyo-Nothofagetum pumilionis) is here considered as belonging to the Wintero-Nothofagetea class. Superfluous homonyms (Art. 22): Ass. Nothofago pumilionis-Araucarietum araucanae // Ass. Senecioni linariifolii-Nothofagetum pumilionis // Ass. Carici minutissimae-Nothofagetum pumilionis // Ass. Drimydo andinae-Nothofagetum pumilionis // Ass. Valeriano lapathifoliae-Nothofagetum pumilionis // Ass. Festuco scabriusculae-Nothofagetum pumilionis // Ass. PernettyoNothofagetum pumilionis // Ass. Chusqueo-Nothofagetum pumilionis.