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The lleuque forests of South Central Chile: a phytosociological study and syntaxonomical classification within South American temperate forests.

INTRODUCTION

The lleuque [Prumnopitys andina (Poepp. ex Endl.) de Laub] is a gymnosperm, a practically en demic plant of Chile, which occurs in some temperate forests located on a strip of land extending approximately from parallel 35[degrees]50' to 40[degrees] 30' south (CONAF 1998). It grows mostly in the lower valleys of the Andean Cordillera and, to a much lesser extent, in rain shadow sites of some areas of the Cordillera de la Costa (Nahuel Buta and Cordillera Pelada), under extremely difficult conditions for its survival (Figure 1). In a comprehensive assessment following UICN criteria (Hechenleitner & al., 2005), the lleuque populations have recently been ranked as vulnerable, VU (B2ab).

[FIGURE 1 OMITTED]

Prumnopitys andina, the most suitable name for this plant species belonging to the Podocarpaceae family, should prevail over Podocarpus andina or Prumnopitys spicata (see MILL & Quinn 2001). General works, such as those by Donoso (1983), Rodriguez & al. (1983), Rodriguez (1988) or Rodriguez & Quezada (1995), have already sufficiently described this Chilean (and only in very specific places) Argentinian tree (Zuloaga & Morrone 1996) with regards its botanical and morphological characteristics (leaves, flowers, etc.). However, these descriptive approaches deal very little with the tree's sociological behaviour or companion species (apart from some brief comments on its ecological requirements). As a result of the ecological preferences of the tree, it seldom gives rise to extensive formations, a fact which has made the tree appear to be of minor importance not only for descriptive works dealing with Chilean vegetation (GAJARDo, 1994), but also for works to classify it for forestry purposes (Donoso, 1981).

Recent studies confirmed by experts such as Hechenleitner & al., (2005) suggest that the lleuque is geographically an extremely restricted coniferous tree, after only the pehuen (Araucaria araucana) and the alerce (Fitzroya cupressoides), the most restricted of all. Consequently, we must dismiss prior interpretations which allegedly recorded its occurrence as far as parallel 45[degrees] south (Woltz, 1985) as incorrect and obviously based on confusion with another species endemic to Chile: Podocarpus nubigena. Whereas alerce and pehuen are large-sized, coniferous tree species (adult individuals can grow up to 30 metres or more) capable of dominating forest masses over their angiosperm companions, the lleuque barely reaches 20 m at the most, and usually only reaches 12-16 m in height. The lleuque is therefore forced to co-exist in its distribution area with other potentially taller tree species such as the roble (Nothofagus obliqua), the coihue (Nothofagus dombeyi) and even the cipres de la cordillera (Austrocedrus chilensis).

More than two decades ago, one of the authors proposed an association characterized by the lleuque. In a phytosociological synthesis, Ramirez & Figueroa (1985) suggested the association named Prumnopito-Nothofagetum obliquae nomen nudum, which was basically characterized by the occurrence of Chilean oaks associated with these coniferous trees. Since there were no supportive releves at the time, the proposed association was invalid according to the requirements of the Code of Phytosociological Nomenclature (Weber & al., 2000) and remained as a nomen nudum. The name was hardly ever used again in any descriptive botanical work. The only exception was a compilation by Luebert & Pliscoff (2006), who mentioned it as one of the zonal communities which could be found in the territorial area of their unit 54: "Temperate, caducifolious Nothofagus obliqua and Laurelia sempervirensforest".

Aware of the content vacuum around the nomen nudum status, we took up the topic again and carried out a number of releves in order to properly define the floristic composition, map the chorological distribution and determine the bioclimatic preferences, dynamic aspects and possible exploitation of these wood formations where lleuque occurs. Our records suggest that all these characteristics indicate a genuine phytosociological association.

METHODOLOGY

We explored the areas of Chile where the occurrence of lleuque had been massively recorded. For this purpose we visited different areas of the valleys at the foot of the Andean Cordillera throughout the regions of Bio-Bio and La Araucania (VIIIth and IXth regions, respectively, in the Chilean administrative arrangement; see Figure 1). Our aim was to sample plant formations with a considerable presence of lleuque trees, with a more or less natural appearance and a homogeneous wood mass, dismissing any man-induced discontinuities. In these sites we made releves using the phytosociological method of BraunBlanquet (1979), later updated by Gehu & Rivas-Martinez (1981).

To determine the species growing in each sampling area we followed the nomenclature suggested by Marticorena & Quezada (1985) and Marticorena & Rodriguez (1995, 2001, 2003 and 2005). For syntaxonomical purposes, we complied with the directives of the 3rd edition of the Code of Phytosociological Nomenclature (Weber & al., 2000).

RESULTS

The lleuque populations in the Cordillera de la Costa are so small and their natural environment has been so dramatically altered by Pinus reforestation that we will omit the forest releves including this species in this area of Chile. In the distribution area, in the foothills of the Andean Cordillera, there are occasional forest patches dominated by lleuque trees (and variably shared with broad-leaved species, mostly belonging to the Nothofagus genus). As we will discuss later, Prumnopitys andina tends to grow on the lower levels of the mountain slopes, that is, where it can most easily form extensive populations on considerably large sites. However, in the pre-Cordilleran valleys where the community has its optimum, natural environments have also been dramatically altered by the native mapuche population. For this reason, although scattered mature trees and sometimes even tree copses are frequently found, the understorey has often been reduced to a minimal expression by cattle-farming (goats, cows and horses) activities. Examples are abundant in all the river valleys with headwaters in the Andean pre-Cordillera. We have found lleuque copses, for instance, in the valleys of the rivers Nuble, Queuco, Bio-Bio and Tolten.

In spite of all this, we were also able to sample some sufficiently natural formations in which, as they were in topographical locations which were fairly inaccessible to cattle, human action seems to have had little or no impact on the configuration of the forest. Table 1, with a total of 14 releves, comprises all these samples supporting our proposal for the new association Nothofago obliquae-Prumnopitydetum andinae Amigo, Rodriguez-Guitian & Ramirez ass. nova hoc loco (holotypus rel. 7, Table 1).

The new community shows not only the constant affinity of lleuque for roble (Nothofagus obliqua) but also the ability of lleuque to join up with other trees having rather opposite ecological preferences. An example is coihue (Nothofagus dombeyi), with which it forms more shady arrangements as a result of their shared evergreen character. It also joins up with the cipres de la cordillera (Austrocedrus chilensis), a more heliophilous and to a certain extent xerophilous species. This last matching reveals a somewhat sub-Mediterranean influence in the formations shared by these coniferous trees.

DISCUSSION

About the habitat

We think that it is correct to say that Prumnopitys andina grows "mostly at the bottom of valleys near large rivers" (Hechenleitner & al., 2005). However, the forest patches in which this coniferous tree may become dominant show an edaphotopographical peculiarity: they tend to be located on colluvial deposits of large rocky blocks, often at the foot of cliffs or steep slopes (Figure 2), in an ecological environment which resembles the European forests belonging to the Tilio-Acerion Klika 1955 (Ellenberg, 1988) alliance. The lleuque's ability to germinate and develop on deep, very stony soils has probably provided it with efficient survival options to defend itself against the human colonization of the valley floors and the usual activities that ensue: land deforestation, farming and cattle-raising and human settlement. It is well known that in sites which have been partly deforested for cattle-farming activities but where adult lleuque trees have been maintained, these trees no longer regenerate because pigs and goats eat their fleshy seeds (Hechenleitner & al., op. cit., Figure 3).

About the chorological and bioclimatic diagnosis

The new association suggested is endemic to the temperate territory, which is more or less coincidental with the Subantarctic Province in the biogeographical arrangement by Cabrera & Willink (1973), or the Valdivian-Magallanic Region, according to the chorological scheme suggested by Rivas-Martinez (see in Costa, 2004). In default of other more detailed biogeographical sectorization models, in the mapping suggested by Luebert & Pliscoff (2006) this association would be part of the "Caducifolious forest" plant formation and of the vegetation belt called "Caducifolious temperate Nothofagus obliqua and Laurelia sempervirens forest" (hereafter "forest no. 54 roble-laurel").

[FIGURE 2 OMITTED]

The bioclimatic indexes of sites not only on the mountain peaks but also on the foothills of the Cordillera cannot be accurately calculated as there are no suitable weather stations in the Andean areas of Chile. Nevertheless, the records measured at the nearby Lonquimay weather station (Amigo & al., 2007a), at an altitude of 900 m, 38[degrees] 26' south and 71[degrees] 15' west, can be taken as approximate values. According to the scheme suggested by Rivas-Martinez (1993, 2007, On Line), these bioclimatic records clearly belong to the upper supratemperate (Tp = 1005) and upper hyperhumid (Io = 19.1) ranges. Taking into account these records and the sampled altitudinal range (630-925 m), the geographical areas colonized by the Nothofago obliquae-Prumnopitydetum andinae association show a bioclimatic profile which corresponds to a supratemperate thermotype. Consequently, we categorically reject the subordination of the name "PrumnopytoNothofagetum obliquae" with its vegetation belt "forest no. 54 roble-laurel" suggested by Luebert & Pliscoff (op.cit.), since these authors ascribe fluctuation ranges of Itc = 223-273 and Tp = 1238-1440 to that belt. These values clearly correspond to a bioclimatically mesotemperate belt.

The clearest bioclimatic peculiarity associated with the distribution of lleuque forests is a certain degree of continentality. Most of the territories where they occur (we exclude here the vestigial sites of the Cordillera de la Costa) are clearly semihyperoceanic, sometimes even euoceanic. At the Lonquimay weather station, the Ic = 14.0, that is, the threshold value between semihyperoceanic and euoceanic conditions. All these temperate territories are also bioclimatically characterized by their sub-Mediterranean character, that is, they undergo a certain degree of water deficit in summer time. Although there are no accurate weather records available, the distribution of Prumnopitys andina tends clearly to coincide with the non-co astal area described by Luebert & Pliscoff (op.cit.: 66, Figure 14) as having a temperate subMediterranean bioclimate.

About the floristic composition

Although the analysis shown in Table 1 reveals some variability, the community as a whole presents a constant occurrence of nemoral species. In other words, the community is able to maintain an acceptable forest structure which becomes less defined as the sub-Mediterranean conditions become more apparent, and the copse physiognomy changes due to the progressively dominant presence of Austrocedrus chilensis. The combination of a coniferous tree, with a higher cover rate, and a deciduous species, such as Nothofagus obliqua, gives rise to a myriad of understorey microhabitats with extraordinarily changing light conditions (Figure 4) and very different contributions of dead leaves. The phenomenon is particularly prominent because it also frequently takes place on very irregular microtopographical terrains with abundant large rocky outcrops. These microhabitats induce the occurrence of species belonging to the whole range of biotypes peculiar to a temperate forest. Included in this last group, the scrub species which present their optimum in the wood mantles of temperate forests sensu lato are the most noteworthy. Given their autoecological profile, these species occur very frequently, albeit with a low cover ratio, in woodland understoreys such as those of Nothofago-Prumnopitydetum, where light availability varies considerably from one place to another. In Table 1 we have highlighted the occurrence of these nanophanerophytes by placing them apart as representatives of communities belonging to the Aritotelienea chilensis subclass. The association which is usually noteworthy as its mantle scrub is the Azaro microphyllae-Aristotelietum chilensis, whose distribution in continental areas not subjected to extreme low temperatures had previously been related to the occurrence of roble-lleuque forests (Amigo & al., 2007b).

[FIGURE 3 OMITTED]

There are also many examples of species which grow in the dense shade at the foot of the lleuques and therefore grow less vigorously or at a lower phenological rate, and do not develop all their flower or fruit organs. Consequently, they cannot be properly identified as distinct species.

[FIGURE 4 OMITTED]

Table 1 also reveals that the northernmost releves show the smallest number of species. This is probably because not only is there a smaller surface suitable for making releves in these sites but also because the coniferous trees " Prumnopitys andina and Austrocedrus chilensis form marginal forests at the dryness limit of the forest region north of 37,5[degrees] south" (Hildebrand-Vogel, 2002: 119). Not surprisingly, in Table 1 releves 13 and 14, taken on land plots of just 100 m2 in sites north of parallel 37[degrees] , have not only the poorest flora of all but also, as a result of their extreme northern location, some of the more spurious species with respect to the usual profile of the association. Releve no. 13 shows, for example, the occurrence of species such as Aextoxicon punctatum, Laurelia sempervirens, Gevuina avellana or Podocarpus saligna, all belonging to Nothofago-Eucryphion cordifoliae, the most clearly thermophilous flora arrangement within the Wintero-Nothofagetea class. In releve no. 14 the influence of a markedly sub-Mediterranean temperate bioclimate is not only revealed by the dominance of the cipres de la cordillera (Austrocedrus chilensis) but also by the presence of species peculiar to the sclerophyllous forests belonging to the Lithraeo-Cryptocaryetea class, such as Quillaja saponaria, Sophora macrocarpa, Chusquea cumingii, Gochnatia foliolosa, or its succesional stages.

About the syntaxonomical classification

With regard to the syntaxonomical classification, we think that the largest number of species relates the Nothofago obliquae-Prumnopitydetum to the Wintero-Nothofagetea class, although for a correct ascription to this class we accept the following assumptions:

1) The interpretation by Pollmann (2001) is correct in extending the Wintero-Nothofagetea class in order to include a series of forests dominated by species which belong to the Nothofagus genus, and present a colder and more continental profile due to the fact that they grow in Argentinian territory. This author defined the Berberido trigonae-Nothofagetalia dombeyi order to comprise several associations suggested by Eskuche (1968, 1973) as belonging to the Nothofagetea pumilionis-antarcticae class.

2) The supratemperate Argentinian forests dominated by different species belonging to the Nothofagus genus, and even by Austrocedrus chilensis in temperate territories, should be more closely related to their vicariant instances growing in Chile. This can easily be done by subordinating a series of species peculiar to the Nothofagus forests growing on the temperate strip of land in Argentinian territory to the Wintero-Nothofagetea class. However, in the past some researchers of that trans-Andean territory (Eskuche, 1968, 1973 and 1999; Conticello & al., 1996) have opted to include these series in the Nothofagetea pumilionis-antarcticae class.

3) Nothofago obliquae-Prumnopitydetum should therefore be included in the Berberido trigonae-Nothofagetalia dombeyi order and, within this order, temporarily in the Austrocedro-Nothofagion dombeyi alliance, which comprises the most clearly sub-Mediterranean communities of the order. Nevertheless, a correct interpretation of the lleuque forests will require still more accurate research on the locations of the Chilean-Argentinian deciduous forests dominated by robles, from Dioscoreo brachyobotryae-Nothofagetum obliquae, described for the territory in Argentina, to Elymo andiniNothofagetum macrocarpae, described for the Chilean sites of the region known as Libertador O'Higgins (parallel 34[degrees] south).

Appendix 2 shows some flora arrangements which we interpret as characteristic or territorial differentials of three phytosociological units whose profile in supratemperate ArgentinianChilean territories is worth clarifying:

+ The first group shows the species which we consider as characteristic of continentalized, supratemperate Valdivian forests included in the Berberido trigonae-Nothofagetalia dombeyi order. Many of these species have previously been interpreted by Argentinian authors as belonging to the Nothofagetea pumilionis-antarcticae class, although none of them were included as such by the author suggesting the syntaxa (Oberdorfer, 1960). To these we have also added some others which are more usually found in other syntaxa of the Wintero-Nothofagetea class, but whose presence in the Berberido trigonaeNothofagetalia dombeyi forests can be used as a differential to highlight their non-inclusion in Nothofagetea pumilionis-antarcticae.

+ The second group is made up of scrub and microforest species representing scrub succesional stages of the forests belonging to the above mentioned order. Although we support the name Berberido-Nothofagetalia antarcticae suggested by Eskuche (1969) for this unit, we think that some associations originally ascribed to this order are spurious. Ideally this order should be combined with those mantle scrubs and preforests defined for the temperate forests of Chile, that is, included in the Aristotelienea chilensis subclass. As occurs with the preceding case, some of the species chosen for this second group have their optimum in other orders belonging to that subclass. Consequently, their occurrence in the continental area must be interpreted as a differential species.

+ Finally, the third group comprises those species genuinely characteristic of the Nothofagetea pumilionis-antarcticae class, although some others, peculiar to the Nothofagus pumilio forests in the Magallanic Province, could also be added. In this third group, the proportion of species (almost 50%; those marked as +W-N) more or less frequently found in the supratemperate WinteroNothofagetea forests is particularly worth noting. Consequently, their role as characteristic species must be interpreted as less significant. We think that the distinction Nothofagetea pumilionis-antarcticae versus Wintero-Nothofagetea must be identified with the limit between orotemperate versus supratemperate forests. To properly recognize this limit, the Nothofaguspumilio forests become particularly relevant for diagnostic purposes, given their role as bioindicators for the bioclimatic orotemperate belt, as suggested by Amigo & al. (2007a).

CONCLUSIONS

Consequently, we propose the new forest association dominated by the roble (Nothofagus obliqua) and the lleuque (Prumnopitys andina). In the Chilean plant syntaxonomy this association must be included in the Wintero-Nothofagetea class. As a result, we also suggest subordinating to this phytosociological class some syntaxa interpreted in the 20th century by some Argentinian authors as belonging to the Nothofagetea pumilionis-antarcticae class.
APPENDIX 2

Characteristic or differential species of the three
Valdivian-Magallanic syntaxonomical units of Chile and Argentina.

Berberido trigonae-Nothofagetalia dombeyi

Anemone multifida Poir.             Cystopteris fragilis (L.) Bernh.

Austrocedrus chilensis (D.Don)      Dioscorea brachyobotrya Poepp.
Pic.Ser. & Bizzarri                 (W-N)

Berberis trigona Kunze ex           Elymus andinus Trin.
Poepp.& Endl. (W-N)

Bowlesia tropaeolifolia             Galium chilensis Endl.
Gill. & Hook.

Calceolaria filicaulis Clos         Gavilea glandulifera (Poepp.)
                                    Correa

Carex patagonica Speg.              Gavilea odoratissima Poepp.

Diplolepis descolei (T.Mey.)        Maytenus chubutensis (Speg.)
Lieder& Rapini                      Lourt., O'Don.& Sleum.

Berberido-Nothofagetalia antarcticae

Azara microphylla Hook.f. (+ Ach)   Geranium patagonicum Hook.f.

Berberis darwinii Hook.             Lomatia hirsuta (Lam.) Diels ex
                                    J.F.Macbr. (+ Ach)

Berberis microphylla G.Forst.       Maytenus boaria Mol. (+ Ach)
(+ Ach)

Diostea juncea (Gill. & Hook.)      Mutisia decurrens Cav.
Miers

Festuca purpurascens Banks & Sol.   Mutisia spinosa Ruiz & Pavon
ex Hook.f.

Fragaria chiloensis (L.)Duchesne    Myrceugenia ovata (Hook. & Arn.)
                                    Berg var. nannophylla (Burret)
                                    Landrum

Nothofagetea pumilionis-antarcticae

Adenocaulon chilense Less. (+W-N)   Escallonia alpina Poepp. ex DC.

Alstroemeria aurea Graham (+W-N)    Gavilea lutea (Pers.) M.N.Correa

Anemone antucensis Poepp.           Hypochaeris tenuifolia
                                    (Hook.& Arn.) Griseb.

Arachnitis uniflora Phil. (+W-N)    Lagenophora hirsuta Less. (+W-N)

Araucaria araucana K.Koch (+W-N)    Leucheria thermarum (Phil.) Phil.

Berberis montana Gay (+W-N)         Macrachaenium gracile Hook. f.

Berberis serrato-dentata Lechler    Maytenus disticha (Hook.f.) Urban
(+W-N)

Cardamine glacialis (G.Forster)     Nothofagus pumilio (Poepp. &
DC.                                 Endl.) Krasser (+W-N)

Codonorchis lessonii (Brongn.)      Perezia pedicularifolia Less.
Lindl.

Drimys andina (Reiche) R.A.         Perezia prenanthoides Less.
Rodriguez & Quezada (+W-N)

Embothrium coccineum J.R.           Ranunculus peduncularis Sm.
Forster & G.Forster (+W-N)

Berberido trigonae-Nothofagetalia dombeyi

Anemone multifida Poir.             Myoschilos oblonga Ruiz & Pavon
                                    (W-N)

Austrocedrus chilensis (D.Don)      Polystichum plicatum (Poepp. ex
Pic.Ser. & Bizzarri                 Kunze) Hicken

Berberis trigona Kunze ex           Relchela panicoides Steud.
Poepp.& Endl. (W-N)

Bowlesia tropaeolifolia             Trisetum caudulatum Trin.
Gill. & Hook.

Calceolaria filicaulis Clos         Vicia nigricans Hook. & Arn.

Carex patagonica Speg.              Viola maculata Cav.

Diplolepis descolei (T.Mey.)
Lieder& Rapini

Berberido-Nothofagetalia antarcticae

Azara microphylla Hook.f. (+ Ach)   Nothofagus antarctica (G.Forster)
                                    Oerst. (+ Np-a)

Berberis darwinii Hook.             Ovidia andina (Poepp. & Endl.)
                                    Meisn.

Berberis microphylla G.Forst.       Relbunium hypocarpium (L.)
(+ Ach)                             Hemsl.(+ Ach)

Diostea juncea (Gill. & Hook.)      Ribes cucullatum Hook.et Arn.
Miers                               (+ Np-a)

Festuca purpurascens Banks & Sol.   Ribes valdivianum Phil. (+ Ach)
ex Hook.f.

Fragaria chiloensis (L.)Duchesne    Schinuspatagonicus (Phil.) Johnst.

Nothofagetea pumilionis-antarcticae

Adenocaulon chilense Less. (+W-N)   Ribes magellanicum Poir. (+W-N)

Alstroemeria aurea Graham (+W-N)    Ribes nitidissimum Neger

Anemone antucensis Poepp.           Rubus geoides J.E.Sm. (+W-N)

Arachnitis uniflora Phil. (+W-N)    Senecio acanthifolius Hombr. &
                                    Jac-.

Araucaria araucana K.Koch (+W-N)    Seneciopilquensis H.Buek

Berberis montana Gay (+W-N)         Senecio prenanthifolius Phil.

Berberis serrato-dentata Lechler    Uncinia negeri Kuckenthal
(+W-N)

Cardamine glacialis (G.Forster)     Valeriana lapathifolia Vahl (+W-N)
DC.

Codonorchis lessonii (Brongn.)      Viola reichei Skottsb. ex
Lindl.                              Macloskie (+W-N)

Drimys andina (Reiche) R.A.
Rodriguez & Quezada (+W-N)

Embothrium coccineum J.R.
Forster & G.Forster (+W-N)

Top: continental, supratemperate, Valdivian-Magallanic forests
(Chile and Argentina): Berberido trigonae-Notho- fagetalia dombeyi.
The species which tend ideally to occur in other woodland
communities belonging to different orders of the
Wintero-Nothofagetea class are marked with (W-N).

Middle: continental scrub and microforests:
Berberido-Nothofagetalia antarcticae. The species peculiar to other
Aristotelietalia chilensis communities and acting as differentials
are marked with (Ach). Some cases marked with (+ Np-a) are
considered by other authors as characteristic of the
Nothofageteapumilionis-antarcticae class.

Below: orotemperate, Valdivian-Magallanic forests: Nothofagetea
pumilionis-antarcticae. The species which are usually found in
supratemperate Wintero-Nothofagetea forests are marked with
(+ W - N).


doi: 10.5209/rev_LAZA.2010.v31.5

ACKNOWLEDGMENTS

We would like to thank the staff of the Instituto de Botanica of the Austral University of Valdivia for their valuable help and support, Eduardo Castro Rodriguez for his photographic skills and assistance in field research, and the Xunta de Galicia and the university of Santiago for partially financing the time spent in Chile of one of the Spanish authors.
SYNTAXONOMICAL SCHEME OF THE MENTIONED COMMUNITIES

Wintero-Nothofagetea Oberd. 1960
  Wintero-Nothofagenea Oberd. 1960
    Laurelietalia philippianae Oberd. 1960
      Nothofago-Eucryphion cordifoliae oberd. 1960
    Berberido trigonae-Nothofagetalia dombeyi Pollmann 2001
     Myrceugenio-Nothofagion dombeyi (Eskuche 1999) Pollmann 2001
      Dioscoreo brachyobotryae-Nothofagetum obliquae Eskuche (1973)
      1999
     Austrocedro-Nothofagion dombeyi Eskuche 1968
      Nothofago obliquae-Prumnopitydetum andinae Amigo,
      Rodriguez-Guitian & Ramirez ass. nova
     Elymo andini-Nothofagion obliquae oberd. 1960 prov.
     Elymo andini-Nothofagetum macrocarpae oberd. 1960 prov. corr.

   Aristotelienea chilensis Amigo, Ramirez & Quintanilla 2007
    Aristotelietalia chilensis (Oberd. 1960) Hildebrand 1983
     Berberidion buxifoliae oberd. 1960
      Azaro microphyllae-Aristotelietum chilensis Amigo, Ramirez &
      Quintanilla 2007
     Berberido-Nothofagetalia antarcticae Eskuche 1969 (excl.
      Discarietum articulatae)


Recibido: 3 septiembre 2009 Aceptado: 25 noviembre 2009

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Gehu, J.M. & Rivas-Martinez, S.--1981--Notions fondamentaux de phytosociologie--In: Dierschke, H. (Ed.). Syntaxonomie. Pp. 5-53. J.Cramer, Vaduz.

Hechenleitner, P., Gardner, M.F., Thomas, P.I., Echeverria, C., Escobar, B., Brownless, P. & Martinez, C.--2005--Plantas Amenazadas del Centro-Sur de Chile. Distribucion, Conservacion y Propagacion, 1st ed.--Univ. Austr. Chile. Jard. Bot. Edimburgo. 188 pp.

Hildebrand-Vogel, R.--2002--Structure and dynamics of southern Chilean natural forests with special reference to the relation of evergreen versus deciduous elements--Folia Geobot. 37: 107-128.

Luebert, F. & Pliscoff, P.--2006--Sinopsis bioclimatica y vegetacional de Chile-- Ed. Univ. 316 pp.

Marticorena, C. & Quezada, M.--1985--Catalogo de la flora vascular de Chile--Gayana Bot. 42 (1-2): 5-157.

Marticorena, C. & Rodriguez, R.--1995--Flora de Chile. Vol. I. Pteridophyta-Gymnospermae--Univ. Concepcion. Ed. Anibal Pinto. 352 pp.

Marticorena, C. & Rodriguez, R.--2001--Flora de Chile. Vol. 2. Winteraceae-Ranunculaceae--Univ. Concepcion. Trama Impresores S.A. 99 pp.

Marticorena, C. & Rodriguez, R.--2003--Flora de Chile. Vol. 2(2). Berberidaceae-Betulaceae--Univ. Concepcion. Impresos Siglo XXI Ltda., Concepcion. 93 pp.

Marticorena, C. & Rodriguez, R.--2005--Flora de Chile. Vol. 2(3). Plumbaginaceae-Malvaceae--Univ. Concepcion. Impresos Siglo XXI Ltda., Concepcion. 128 pp.

Mill, R.R. & Quinn, C.J.--2001--Prumnopitys andina reinstated as the correct name for Lleuque, the Chilean conifer recently renamed P. spicata (Podocarpaceae)--Taxon 50: 1143-1154.

Oberdorfer, E.--1960--Pflanzensoziologische Studien in Chile-- Flora & Vegetatio Mundi 2: 1-208.

Pollmann, W.--2001--Caracterizacion floristica y posicion sintaxonomica de los bosques caducifolios de Nothofagus alpina (Poepp. & Endl.) Oerst.en el centro-sur de Chile--Phytocoenologia 31(3): 353-400.

Ramirez, C. & Figueroa, H.--1985--Fitosociologia de los Nothofagus de la zona higromorfica chilena--Bosque 6: 127-132.

Rivas-Martinez, S.--1993--Clasificacion bioclimatica de la Tierra--Folia Bot. Matritensis 11: 1-20

Rivas-Martinez, S.--2007--Mapa de series, geoseries y geopermaseries de vegetacion de Espana (Memoria del Mapa de Vegetacion Potencial de Espana. Parte 1)--Itinera Geobot. 17: 5-435

Rivas-Martinez, S.--On-Line--www.globalbioclimatics.es.

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Rodriguez, R. & Quezada, M.--1995--Prumnopitys Phil.--In Marticorena, C. & Rodriguez, R. (Eds.). Flora de Chile, vol I. Pteridophyta-Gymnospermae. Pp. 321-322. Univ. Concepcion.

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Javier Amigo (*), Manuel A. Rodriguez-Guitian (**) & Carlos Ramirez (***)

* Laboratorio de Botanica, Facultad de Farmacia, Universidad de Santiago de Compostela. E-15782 Santiago de Compostela (Galicia, Espana). E-mail: javier.amigo.vazquez@usc.es ** Departamento de Produccion Vexetal. Escola Politecnica Superior de Lugo. Universidad de Santiago de Compostela. 27002-Lugo *** Instituto de Botanica, Universidad Austral de Chile. Casilla 567, Valdivia (Chile).
Table 1
Nothofago obliquae-Prumnopitydetum andinae ass. nova
(Austrocedro-Nothofagion dombeyi, Berberido trigonae-Nothofagetalia
dombeyi, Wintero-Nothofagetea)

Altitude
(m a.s.l.)                    630     735     800     925
Slope ([degrees])             16      30      35      10
Aspect                        SE     NE A     WSW     NNE
Cover (%)                     100     100     90      95
Vegetation bight (m)          18     12-14   12-22   20-28
Plot area ([m.sup.2])         200     200     500     500
N[degrees] of species         32      26      22      23
N.                             1       2       3       4
Dominant trees
Prumnopitys andina             5       4       4       4
Nothofagus obliqua             2       2       2       1
Austrocedrus chilensis         .       .       .       +
Nothofagus dombeyi             2       2       3       3
Characteristics of association and upper units
Chusquea culeou                1       1       3       2
Osmorhiza chilensis            2       +       1       2
Blechnum hastatum              1       2       1       1
Alstroemeria aurea             +       r       +       +
Lomatia hirsuta                .       +       2       2
Myoschilos oblonga             .       1       1       1
Maytenus chubutensis           .       .       +       1
Polystichum plicatum           .       +       1       1
Relchela panicoides            .       .       1       +
Viola maculata                 .       1       r       +
Calceolaria filicaulis         .       .       .       +
Mutisia decurrens              .       .       +       r
Schinus patagonicus            .       .       .       +
Lomatia dentata                r       .       .       .
Vicia nigricans                .       .       1       .
Lathyrus subandinus            .       .       +       +
Sanicula crassicaulis          +       .       .       .
Pseudopanax laetevirens        r       .       .       .
Adenocaulon chilense           1       .       .       .
Bowlesia tropaeolifolia        .       .       .       .
Cystopteris fragilis           .       .       .       .
Trisetum caudulatum            .       .       .       .
Carex patagonica               .       .       .       .
Aristotelienea species
Relbunium hypocarpium          +       .       +       +
Azara microphylla              2       1       1       1
Aristotelia chilensis          1       1       .       .
Ribes gr. valdivianum          1       +       +       +
Berberis darwini               +       +       1       .
Rosa rubiginosa                +       .       .       .
Berberis microphylla           .       .       .       .
Boquila trifoliolata           1       r       .       .
Myrceugenia nanophylla         1       1       .       .
Azara integrifolia             .       .       .       .
Fuchsia magellanica            +       +       .       .
Companion species              .       .       .       .
Adiantum chilense              .       1       .       .
Dactylis glomerata             1       1       .       .
Acaena ovalifolia              +       1       .       .
Prunella vulgaris              +       +       .       .
Solanum cyrtopodium            +       +       .       .
Carex aphylla                  .       .       .       +
Leucheria caerulescens         .       .       +       .
Valeriana cf. valdiviana       .       .       .       .
Gaultheria phyllireifolia      +       .       .       .
Lapsana communis               .       +       .       .
Adiantum scabrum               .       .       .       .
Gavilea sp.                    .       .       .       .
Dioscorea sp.                  1       r       .       .
Asplenium dareoides            +       .       .       .
Agrostis leptotricha           .       .       +       .
Nasella chilensis              .       .       .       .
Acaena argentea                .       .       .       .
Cynanchum pachyphyllum         .       .       .       .
Equisetum bogotense            .       1       .       .

Altitude
(m a.s.l.)                    725     800     800     865
Slope ([degrees])             34      40      40      35
Aspect                        NE      NE      SSW     NNE
Cover (%)                     100     95      90      90
Vegetation bight (m)         16-18   15-26   15-20   15-17
Plot area ([m.sup.2])         300     300     250     250
N[degrees] of species         23      28      30      25
N.                             5       6       7       8
Dominant trees
Prumnopitys andina             5       5       5       5
Nothofagus obliqua             2       1       3       1
Austrocedrus chilensis         .       +       +       +
Nothofagus dombeyi             +       .       .       .
Characteristics of association and upper units
Chusquea culeou                1       1       +       3
Osmorhiza chilensis            1       1       1       2
Blechnum hastatum              2       1       .       .
Alstroemeria aurea             +       +       +       .
Lomatia hirsuta                .       .       3       2
Myoschilos oblonga             .       +       +       1
Maytenus chubutensis           .       .       +       1
Polystichum plicatum           +       .       1       .
Relchela panicoides            .       +       +       1
Viola maculata                 .       .       +       +
Calceolaria filicaulis         r       +       +       +
Mutisia decurrens              .       .       +       .
Schinus patagonicus            r       .       .       r
Lomatia dentata                .       .       .       .
Vicia nigricans                .       +       +       .
Lathyrus subandinus            .       .       .       .
Sanicula crassicaulis          .       1       .       .
Pseudopanax laetevirens        +       .       .       .
Adenocaulon chilense           .       .       .       .
Bowlesia tropaeolifolia        .       +       1       .
Cystopteris fragilis           .       +       +       .
Trisetum caudulatum            .       .       1       .
Carex patagonica               .       .       .       1
Aristotelienea species
Relbunium hypocarpium          r       1       1       +
Azara microphylla              +       1       +       .
Aristotelia chilensis          1       1       +       .
Ribes gr. valdivianum          +       .       +       r
Berberis darwini               .       .       +       .
Rosa rubiginosa                .       +       r       +
Berberis microphylla           .       +       r       +
Boquila trifoliolata           1       .       .       .
Myrceugenia nanophylla         .       .       .       .
Azara integrifolia             .       .       .       .
Fuchsia magellanica            .       .       .       .
Companion species              .       .       .       .
Adiantum chilense              +       +       .       .
Dactylis glomerata             1       +       .       .
Acaena ovalifolia              .       .       .       .
Prunella vulgaris              +       +       .       .
Solanum cyrtopodium            .       +       .       .
Carex aphylla                  .       .       .       .
Leucheria caerulescens         .       .       1       .
Valeriana cf. valdiviana       1       .       +       .
Gaultheria phyllireifolia      .       .       r       .
Lapsana communis               +       .       .       .
Adiantum scabrum               .       .       .       .
Gavilea sp.                    .       +       .       .
Dioscorea sp.                  .       .       .       .
Asplenium dareoides            +       .       .       .
Agrostis leptotricha           .       .       .       .
Nasella chilensis              .       .       .       .
Acaena argentea                .       +       .       .
Cynanchum pachyphyllum         .       .       .       .
Equisetum bogotense            .       .       .       +

Altitude
(m a.s.l.)                    760     750     830     730
Slope ([degrees])             15      10      30      26
Aspect                        NE      SSE      W      NNW
Cover (%)                     90      100     95      95
Vegetation bight (m)         12-15   12-14   14-20   15-20
Plot area ([m.sup.2])         120     200     400     300
N[degrees] of species         25      24      25      15
N.                             9      10      11      12
Dominant trees
Prumnopitys andina             5       3       5       4
Nothofagus obliqua             +       4       3       4
Austrocedrus chilensis         .       1       2       .
Nothofagus dombeyi             .       2       .       .
Characteristics of association and upper units
Chusquea culeou                1       .       2       1
Osmorhiza chilensis            +       .       1       1
Blechnum hastatum              2       1       .       2
Alstroemeria aurea             +       r       .       .
Lomatia hirsuta                1       1       2       .
Myoschilos oblonga             .       1       1       .
Maytenus chubutensis           .       1       +       .
Polystichum plicatum           +       .       +       .
Relchela panicoides            .       .       1       .
Viola maculata                 .       .       +       .
Calceolaria filicaulis         .       .       .       .
Mutisia decurrens              .       +       r       .
Schinus patagonicus            .       +       r       .
Lomatia dentata                .       .       .       +
Vicia nigricans                .       .       +       .
Lathyrus subandinus            .       .       .       .
Sanicula crassicaulis          .       .       .       +
Pseudopanax laetevirens        .       .       .       .
Adenocaulon chilense           +       .       .       .
Bowlesia tropaeolifolia        .       .       .       .
Cystopteris fragilis           .       .       .       .
Trisetum caudulatum            .       .       1       .
Carex patagonica               .       .       r       .
Aristotelienea species
Relbunium hypocarpium          +       +       1       r
Azara microphylla              +       1       1       .
Aristotelia chilensis          1       2       1       1
Ribes gr. valdivianum          +       +       .       .
Berberis darwini               1       +       +       .
Rosa rubiginosa                .       +       +       +
Berberis microphylla           .       +       .       .
Boquila trifoliolata           1       .       .       .
Myrceugenia nanophylla         1       .       .       .
Azara integrifolia             .       +       .       1
Fuchsia magellanica            .       .       .       .
Companion species              .       .       .       .
Adiantum chilense              +       .       .       +
Dactylis glomerata             1       .       .       .
Acaena ovalifolia              +       .       r       .
Prunella vulgaris              .       .       .       .
Solanum cyrtopodium            .       .       .       .
Carex aphylla                  .       .       .       +
Leucheria caerulescens         .       .       .       .
Valeriana cf. valdiviana       1       .       .       .
Gaultheria phyllireifolia      +       .       .       .
Lapsana communis               1       .       .       .
Adiantum scabrum               .       .       .       +
Gavilea sp.                    .       .       .       .
Dioscorea sp.                  .       .       .       .
Asplenium dareoides            .       .       .       .
Agrostis leptotricha           .       .       .       .
Nasella chilensis              .       .       .       1
Acaena argentea                .       .       .       .
Cynanchum pachyphyllum         +       1       .       .
Equisetum bogotense            .       .       .       .

Altitude
(m a.s.l.)                    860     750
Slope ([degrees])              5      32
Aspect                        SW      ESE
Cover (%)                     80      70
Vegetation bight (m)         10-18   8-13
Plot area ([m.sup.2])         100     100
N[degrees] of species         18      21       Frequency
N.                            13      14
Dominant trees
Prumnopitys andina             3       +          14
Nothofagus obliqua             1       2          14
Austrocedrus chilensis         2       4           8
Nothofagus dombeyi             .       .           6
Characteristics of association and upper units
Chusquea culeou                .       .          11
Osmorhiza chilensis            .       .          11
Blechnum hastatum              .       +          10
Alstroemeria aurea             .       .           9
Lomatia hirsuta                .       1           9
Myoschilos oblonga             .       .           8
Maytenus chubutensis           .       +           7
Polystichum plicatum           .       .           7
Relchela panicoides            .       .           6
Viola maculata                 .       .           6
Calceolaria filicaulis         .       .           5
Mutisia decurrens              .       .           5
Schinus patagonicus            .       .           5
Lomatia dentata                .       1           4
Vicia nigricans                .       .           4
Lathyrus subandinus            .       .           3
Sanicula crassicaulis          .       .           3
Pseudopanax laetevirens        .       .           2
Adenocaulon chilense           .       .           2
Bowlesia tropaeolifolia        .       .           2
Cystopteris fragilis           .       .           2
Trisetum caudulatum            .       .           2
Carex patagonica               .       .           2
Aristotelienea species
Relbunium hypocarpium          .       .          11
Azara microphylla              .       .          10
Aristotelia chilensis          2       .          10
Ribes gr. valdivianum          +       .          10
Berberis darwini               .       .           7
Rosa rubiginosa                .       .           7
Berberis microphylla           .       .           4
Boquila trifoliolata           .       .           4
Myrceugenia nanophylla         .       .           3
Azara integrifolia             .       1           3
Fuchsia magellanica            .       .           2
Companion species              .       .           .
Adiantum chilense              .       .           5
Dactylis glomerata             .       .           5
Acaena ovalifolia              .       .           4
Prunella vulgaris              .       .           4
Solanum cyrtopodium            .       .           3
Carex aphylla                  .       1           3
Leucheria caerulescens         .       1           3
Valeriana cf. valdiviana       .       .           3
Gaultheria phyllireifolia      .       .           3
Lapsana communis               .       .           3
Adiantum scabrum               .       .           2
Gavilea sp.                    .       .           2
Dioscorea sp.                  .       .           2
Asplenium dareoides            .       .           2
Agrostis leptotricha           .       .           2
Nasella chilensis              +       .           2
Acaena argentea                +       .           2
Cynanchum pachyphyllum         .       .           2
Equisetum bogotense            .       .           2

Other species: Characteristics species Azara lanceolata r, Blechnum
mochaenum, Blechnumpenna-marina and Rhaphithamnus spinosus +, Luma
apiculata 1 in 1; Elymus andinus + in 4; Anemone multifida + in 8;
Embothrium coccineum 1 in 9; Berberis rotundifolia r in 11; Gevuina
avellana r, Aextoxicon punctatum +, Drimys winteri and
Muehlenbeckia hastulata 1, Laurelia sempervirens and Podocarpus
saligna 2, Hydrangea serratifolia 3 in 13; Diostea juncea + in 14.
Companion species: Polypodium feuillei + in 5; Loasa sp. r,
Geranium robertianum and hemicryptophyte unknown + in 6; Asteracea
sp. and Stellaria sp. + in 7; Holcus lanatus and Hypericum
perforatum +, Lactuca serriola 1 in 8; Festuca scabriuscula +,
Lotus uliginosus 1 in 9; Pernettya myrtilloides and Poacea sp. +,
Rumohra adiantiformis 1 in 10; Poa gr. trivialis + in 11; Cirsium
arvense, Solanum gr. gayanum and Tristerix tetrandrus + in 13;
Baccharis concava, Cheilanthes glauca, Chusquea cumingii, Oxalis
araucana and Satureja gilliesi +, Sophora macrocarpa 1, Gochnatia
foliolosa and Quillaja saponaria 2 in 14.

Localities (author's code for releves is included): 1, 2, 5, 9 and
10: La Araucania, Cautin, from Cahuilelfun towards Curarrehue,
39[degrees]15'-71[degrees]26' (070124/1); from Reigolil towards
Curarrehue, 39[degrees]11'-71[degrees]27' (070123/4);
39[degrees]12'-71[degrees]27' (070123/5); from Reigolil towards
Curarrehue, 39[degrees]11'-71[degrees]27' (090122/3); Conguillio
National Park, Southwards exit to Melipeuco,
38[degrees]45'-71[degrees]37' (070119/2); 3, 4, 6-8 and 11: La
Araucania, Malleco, Bio-Bio river valley, Casas de Lolco,
38[degrees]08'-71[degrees]24' (090120/3); from Troyo Northwards,
38[degrees]10'-71[degrees]18' (090120/2); between Contraco and
Lolco, 38[degrees]08'-71[degrees]20' (070118/5); Bio-Bio river
valley, from Troyo Northwards, 38[degrees]11'- 71[degrees]18'
(070118/3), holotypus ass. Bio-Bio river valley, from Lonquimay
towards Troyo, 38[degrees]19'-71[degrees]19' (090119/1); Nalcas
National Reserve, from Casas de Lolco towards Lonquimay.
38[degrees]11'-71[degrees]26' (090120/4); 12: Bio-Bio, Bio-Bio:
river Queuco valley upwards, close to Comuna Caunico,
37[degrees]43'-71[degrees]23' (070117/1); 13 and 14: Bio-Bio,
Nuble: entrance to Nuble National Reserve,
36[degrees]58'-71[degrees]30' (080119/1); Nuble valley upwards, a
little downwards from Los Sauces, 36[degrees]40'-71[degrees]16'
(070115/2).
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