CSIRO PUBLISHING Australian Systematic Botany, 2020, 33, 174–190 https://doi.org/10.1071/SB19005 Biogeographic characterisation of the Austral High Andean district, Patagonian province, based on vascular plant taxa Alfredo PadróA, Viviana Hechem A and Juan J. Morrone B,C A Facultad de Ciencias Naturales, Universidad Nacional de la Patagonia San Juan Bosco, Ruta 259 kilómetro 16.41, 9200 Esquel, Chubut, Argentina. Email: padroalfredo@outlook.com; vivianahf03@yahoo.com.ar. B Museo de Zoología ‘Alfonso L. Herrera’, Departamento de Biología Evolutiva, Facultad de Ciencias, Universidad Nacional Autónoma de México, 04510 Mexico City, Mexico. C Corresponding author. Email: juanmorrone2001@yahoo.com.mx Abstract. The Austral High Andean area extends from south-eastern Mendoza, Argentina, to the southernmost tip of South America in the form of isles on the peaks of the Andes range. The objective of this biogeographic regionalisation study was to characterise this area. Individual tracks were made on the basis of the distribution maps of 232 species of vascular plants present in the area, from which localities were identified and georeferenced. A parsimony analysis of endemicity (PAE) was used to obtain a generalised track. The results support an area of endemism located mainly in the Neuquén province, which is treated as a district of the Patagonian province that belongs to the Patagonian subregion of the Andean region. This track analysis is a preliminary contribution for understanding the distributional patterns of the High Andean biota within an evolutionary biogeographic framework. Additional keywords: biogeography, distributional patterns; High Andean area, parsimony analysis of endemicity, track analysis. Received 24 January 2019, accepted 5 August 2019, published online 5 February 2020 Introduction Panbiogeography, originally formulated by the Italian botanist Léon Croizat (1958, 1964), is a historical biogeographic approach that emphasises the role of location in the history of life. Croizat (1964) postulated that geographic barriers evolve alongside biotas, which he summarised with the metaphors ‘Life and Earth evolve together’ and ‘Space, time, form = biological synthesis’ (Llorente-Bousquets et al. 2000; Morrone 2004a). This approach highlights the relevance of the distribution of taxa in space so as to understand the history of life on earth (Crisci et al. 2003). According to Cabrera and Willink (1980), the High Andean biogeographic province includes the mountain peaks of the Andean mountain ranges, from Venezuela to Tierra del Fuego in the southern tip of South America. At tropical latitudes, this province appears above 4200 m above sea level, but towards the south it extends to lower altitudes. Around the 34S parallel, it is found above 3000 m, around the 40S parallel between 1500 and 1000 m, and around the 54S parallel at 500 m. According to the phytogeographic classification of Cabrera (1976) and Cabrera and Willink (1980), the High Andean province belongs to the Andean–Patagonian dominion together with the Patagonian and Punan provinces. This dominion, which belongs to the Neotropical region, extends from the mountain peaks of Venezuela and Colombia, covering the ranges and punas of Journal compilation Ó CSIRO 2020 Ecuador, Peru, Bolivia and Argentina, and reaching the Argentinean province of Tierra del Fuego, including the coastal deserts of Peru and Chile, and the Patagonian steppes of Neuquén, Río Negro, Chubut and Santa Cruz. More recent analyses have treated this area as part of the Andean region, which is closely related to South Africa, Australia, New Zealand and Antarctica (Morrone 2001, 2006, 2014a, 2015a, 2015b; Roig-Juñent et al. 2018). The Andean region (Morrone 1996, 2001) includes the Patagonian, Subantarctic and Central Chilean subregions and the South American transition zone. The High Andean areas of the northern Andes have been assimilated to the Paramo province, those of the central Andes were assigned to the Puna province, and those of the southern Andes from central Argentina have been treated as the Cuyan High Andean province (Morrone 2015a; Morrone and Ezcurra 2016). Here, we analyse the southernmost High Andean area, that in Argentina extends from central Mendoza, widening through the Neuquén, Río Negro, Chubut and Santa Cruz provinces up to northern Tierra del Fuego, reaching Chile in the Aysén and Magallanes regions. High-altitude environments are found as isles in the mountain peaks of the Andes (Cabrera and Willink 1980). They are found above the treeline, i.e. the altitude at which trees cannot grow (Ferreyra et al. 2006), which correlates with the 10C (54F) warmest-month isotherm (Daniels and Veblen 2003). In www.publish.csiro.au/journals/asb Austral High Andean district the northern Patagonian Andes, around the 40S latitude, the treeline is found between 1000 and 1500 m (Daniels and Veblen 2003), whereas south of the 50S in southern Santa Cruz and Tierra del Fuego, it descends to 500 m (Ferreyra et al. 2006). The treeline is not a continuous horizontal line but rather fluctuates because of changes in the stability of the ground, like landslides and avalanches, volcanism and fires, alongside other factors (Ferreyra et al. 2006). The treeline is composed almost in its entirety by Krummholz (flag trees) of species of Nothofagus Blume (Daniels and Veblen 2003). The High Andean area in the Southern Andes is characterised by a mountainous relief, with gentle or steep slopes, peaks, cirques and glacial troughs and plateaus. Soil is rocky or sandy, usually loose and shallow. Climate is polar (Köppen Climate Classification ETf), and can experience snow, hail or frost at any time of the year, has a low relative humidity, a large diurnal range, fierce winds and high solar irradiance. During winter, snow accumulation can be over 1 m deep (Ferreyra et al. 2006). The environment consists of microhabitats with different environmental conditions, determined by precipitation, altitude, insolation, soil type, presence of nurse plants and facing of the slopes. These microhabitats determine a very high environmental diversity, which has allowed the evolution and settlement of several species with unique adaptations (Ferreyra et al. 2006). Over 300 species of vascular plants have been identified in the area (Correa 1998; Ferreyra et al. 2006). Asteraceae and Poaceae are the best represented families, whereas Senecio L. and Nassauvia Comm. ex Juss. the best represented genera. Species belonging to the genus Nassauvia dominate the coldest, most adverse spots (Ferreyra et al. 2006). Perennial grasses are the dominant life form in the High Andean area, followed by subshrubs and shrubs. Perennial grasses are sparse because of the adverse conditions such as the short growing season. Most species are dwarf or cushion plants, sometimes forming woody shrubs compressed against the ground. This structure appears to be the most suitable to endure extreme conditions, because it provides protection against dehydration, solar radiation, wind and snow accumulation. These shrubs trap heat, moisture and organic matter, playing an important role as nurse plants by protecting other species that sprout and grow between their twigs. Most local plants possess highly developed underground structures, which allows them to grow in unstable substrates, reach moister soil and store substances used to rapidly awaken from their dormant state whenever climatic conditions become favourable (Ferreyra et al. 2006). The distribution of the High Andean plants is tightly related to longitude, so there are different species in the eastern-arid and western-humid areas of the Patagonian Andes (Ferreyra et al. 2006). The eastern mountains usually have a greater species richness because (1) the characteristic aridity of the mountains, associated with a pronounced precipitation gradient where precipitation decreases eastward, allowing the growth of both xeric and montane species; (2) the eastern mountains have been ice free for a longer time since the last ice ages and, therefore, plants have had more time to recolonise them; (3) the snow cover is thinner and less persistent on the eastern mountains and the substrate is more disintegrated, providing better conditions for plant growth; (4) eastern peaks have a higher diversity of environments, from dry slopes to always humid forest spots, Australian Systematic Botany 175 whereas western slopes lack extremely arid environments; and (5) the higher amount of persistent ice or snow covering some western peaks, on the top of the higher glacial erosion in the past, reduces the available area for plants. Currently, there is a growing tourist interest in the montane environments that is not limited to winters but also in summers, when vegetation is more vulnerable. It is worth noting that species that live on the peaks, despite impressive adaptations to adverse environmental conditions, cannot withstand anthropic disturbances, and some adaptations, such as morphological traits or longevity, may become disadvantages in these situations (Ferreyra et al. 2006). Research has shown that the transit of humans through areas where some delicate plants grow causes damage that takes years to recover, or may be even permanent. Considering the key role of some of these species as nurse plants, it is easy to imagine that disturbances affect the entire ecosystem, and that the appropriate sustainable use of these environments is of great relevance (Ferreyra et al. 2006). Our objective is to characterise biogeographically the Austral High Andean biota through the analysis of the distribution of vascular plants. We want to contribute to the identification of biogeographic patterns in the High Andean biota and to the study of its evolutionary biogeographic history. Materials and methods Area The study area includes the geographic areas of the southern Andean range and adjacent regions between 36 and 55S parallel, from central Mendoza, widening in the Neuquén, Río Negro, Chubut and Santa Cruz provinces and reaching Tierra del Fuego province (in Argentina) and the Aysén and Magallanes regions (in Chile). Altitudes range from 1500 m in the northern-most latitudes to 0 m in the southern-most latitudes (Fig. 1). Taxa The 381 species of vascular plants present in the area were analysed, from which 232 containing three or more registered localities were selected (Appendix 1) and 149 species with two or fewer localities were excluded. So as to establish the distribution of the species, data from previous works (Ferreyra et al. 2006; Hechem et al. 2011) and the revision of works from the databases of the herbaria BAB, BCRU, CONC, LP, SGO, SI (Index Herbariorum, see http://sweetgum.nybg.org/science/ih/, accessed 10 March 2019) were collected. Additionally, the databases of the Instituto de Botánica Darwinion (http:// www.darwin.edu.ar/Proyectos/FloraArgentina/FA.asp, accessed 20 December 2018) and Flora Patagónica (Correa 1998) were consulted to add distribution points. Data analysis The track analysis developed by Croizat (1958, 1964) aims to reconstruct the history of biotas. It is essentially a graphical method, which represents the distribution of a taxon by a map, where the localities are connected with a line so that the sum of segments has a minimum distance. The resulting line is called an individual track. If the individual tracks of different taxa overlap, they constitute a generalised track indicating the existence of an ancestral biota, which was fragmented by tectonic and geological 176 75ºW 33ºS A. Padró et al. Australian Systematic Botany 73ºW 71ºW 69ºW 67ºW 75ºW 65ºW 73ºW 71ºW 69ºW 67ºW 65ºW 0 0 33ºS 33ºS 33ºS 100 100 200 200 35ºS 35ºS 35ºS 35ºS 300 300 400 km 400 37ºS 37ºS 37ºS 37ºS 39ºS 39ºS 39ºS 39ºS 41ºS 41ºS 41ºS 41ºS 43ºS 43ºS 43ºS 43ºS 45ºS 45ºS 45ºS 45ºS 47ºS 47ºS 47ºS 47ºS 49ºS 49ºS 49ºS 49ºS 51ºS 51ºS 51ºS 51ºS 53ºS 53ºS 53ºS 53ºS 55ºS 55ºS 55ºS 55ºS 75ºW 75ºW 73ºW 71ºW 69ºW 67ºW 73ºW 71ºW 69ºW 67ºW 65ºW 65ºW Fig. 1. Location of the High Andean province in the Patagonian subregion (grey), based on Cabrera and Willink (1980). events. When two or more different generalised tracks overlap in an area, they constitute a node, which is a composite or hybrid area (Katinas et al. 1999; Crisci et al. 2003). Localities were georeferenced, maps for each species were made and the individual tracks were drawn. With the aim of simplifying the analysis, the area that contained the georeferenced localities was divided in cells of 1 degree of latitude by 1 degree of longitude and the cells were numbered from south to north and from east to west (Fig. 2). Then, the individual tracks were overlapped with the cells and a data matrix was built, where the rows and columns represent the cells and the tracks respectively. The presence of a track in a cell was coded with a ‘1’ and the absence with a ‘0’ (Morrone 2014b). One row with all ‘0’ was added to root the cladograms. The resulting matrix has 197 rows (cells) and 232 columns (species). Fig. 2. Cells used to build the data matrix for the parsimony analysis of endemicity. The matrix (Appendix 2) was analysed with a parsimony algorithm to generate an area cladogram, based on the taxa shared by the cells (Morrone, 1994, 2014b). Generalised tracks, which represent the overlap of multiple individual tracks, can be observed in the cladogram. This parsimony analysis of endemicity (PAE) groups areas or localities, on the basis of the presence of two or more species (Morrone 2014b). The matrix was analysed with the programs Winclada (ver. 1.00.08, K. C. Nixon, see http://www.cladistics.com/about_winc. htm, accessed 10 March 2019) and NONA (ver. 2.0, P. A. Goloboff, see http://www.cladistics.com/aboutNona. htm, accessed 10 March 2019). The search strategy to obtain the most parsimonious-area cladograms was through a tree bisection and reconnection heuristic search with 10 000 replications. Austral High Andean district Australian Systematic Botany 177 cell 123 cell 133 1 cell 143 2 cell 163 3 cell 153 4 cell 144 5 7 cell 154 6 cell 134 8 cell 114 9 cell 124 Fig. 3. Section of the consensus-area cladogram obtained, showing the generalised track. Species in the nodes and cells are listed in Table 1. Table 1. Presence, secondary absence (extinction) or presence in two independent areas (disjunct) of species in the nodes and cells of Fig. 3 Node or cell Presence Secondary absence Presence in two independent areas 1 2 17, 59, 118, 122 24, 65, 80, 137, 139, 203, 207, 219, 229 88, 111 – 3 4 21, 23, 49, 87, 140, 178, 180, 209 9, 40, 109, 124, 146, 225 144 19, 166 5 – 21, 23 38, 98, 113, 147, 170, 172, 176, 184, 206 7 56, 75, 77, 90, 97, 161, 186, 201, 205, 214, 224 3, 4, 13, 14, 36, 55, 66, 105, 112, 132, 156, 157, 162, 187, 193, 198, 208 – 5, 44, 91, 92, 101, 134, 166, 171, 175, 183 2, 28, 30, 37, 57, 67, 71, 81, 96, 100, 117, 131, 151, 163, 177, 179, 213 16, 33, 62, 69, 74, 79, 93, 154, 154, 195, 236 29, 64, 70, 72, 82, 84, 104, 119, 135, 143, 150, 190, 191, 197 53, 76, 88, 110, 142, 181, 228 42, 94 8 35, 115, 155, 160 2, 28, 30, 37, 43, 45, 59, 65, 100, 134, 136, 137, 149, 151, 165, 215 87, 140 9 Cell 123 Cell 133 Cell 143 Cell 163 1, 8, 52, 120, 133, 152, 158, 164 – – – – Cell 153 Cell 144 103, 116, 169, 195, 216 – Cell 154 85, 129, 199 Cell 134 Cell 114 – 11, 78, 107, 159, 189, 194, 204 Cell 124 – 6 209 43, 45, 149, 167 – 58, 83 2, 5, 24, 34, 45, 65, 80, 134, 139, 163, 167, 203, 207, 219, 229 195 7, 20, 34, 39, 44, 71, 81, 93, 101, 117, 129, 139, 179, 229 16, 32, 49, 62, 73, 79, 88, 150, 180, 209, 232 17, 44, 76, 101, 123 3, 4, 13, 32, 36, 38, 46, 49, 81, 83, 90, 100, 136, 146, 156, 161, 178, 180, 182, 229 59, 119, 215 48, 50, 51, 61, 89, 102, 106, 128, 185, 210, 212, 223 1, 22, 25, 26, 99, 121, 126, 144, 144, 188, 200 31, 38, 113, 119, 188 76, 222 42, 94, 148, 174, 184 – 27, 196 – 27 174 86, 111, 114, 127, 183, 229 31 178 A. Padró et al. Australian Systematic Botany Results The PAE produced 30 area cladograms (Fig. 3), with 1043 steps, consistency index of 0.22 and retention index of 0.68. The presence, secondary absence or presence in two independent areas of the species can be observed in its nodes and terminal units (Table 1). Secondary absence refers to a species that became extinct in a particular area, whereas presence in two independent areas refers to a species that has a disjunct distribution. Only one generalised track was found (Fig. 4) which is restricted to the Neuquén province, in the mountain range near the border with Chile. This generalised track is based on the individual tracks of the following 13 species: Arenaria serpens Kunth, Austroblechnum microphyllum (Goldm.) Gasper 75ºW 33ºS 73ºW 71ºW 69ºW 67ºW 65ºW 0 33ºS 100 200 35ºS 35ºS 300 37ºS 400 km 37ºS & V.A.O.Dittrich, Chaetanthera villosa D.Don, Combera paradoxa Sandwith, Erigeron schnackii Solbrig, Montiopsis polycarpoides (Phil.) Peralta, Moschopsis subandina (Speg.) Dusén, Nastanthus scapigerus (J.Remy) Miers, Noccaea magellanica (Comm. ex Poir.) Holub, Senecio pachyphyllos J. Rémy, S. portalesianus J.Rémy, Trisetum barbinode Trin. and Viola cotyledon Ging. Discussion The generalised track obtained corroborates a distributional pattern previously defined by Hechem et al. (2011) as an endemic Andean pattern. This generalised track suggests the existence of a single biotic unit, which is considered herein as a district within the Andean region, located in the mountain range of the Neuquén province, Argentina, near the Chilean border. So as to name this unit, the Austral High Andean district of Cabrera (1971) is available, although in its present circumscription, it is a more restricted biogeographic unit. As suggested by Rivas Martínez (1987) and Cabrera and Yepes (1940), a district has a moderate extension and is characterised by the existence of associations and peculiar species missing in nearby areas. The biogeographic classification of this district is as follows: * * * 39ºS 39ºS * * 41ºS 41ºS 43ºS 43ºS 45ºS 45ºS 47ºS 47ºS 49ºS 49ºS Andean region (Engler 1882). Patagonian subregion (Lorentz 1876). Patagonian province (de Mello-Leitão 1939). Subandean subprovince (Soriano 1950). Austral High Andean district (Cabrera 1971) In contrast, this district may be considered as belonging to the South American transition zone (Morrone 2004b, 2015a, 2018), as suggested by Roig-Juñent et al. (2018) for the whole Patagonian province. Further analyses are needed to explore this alternative hypothesis. A more thorough study should include taxa of the Neotropical region because of their possible boreal origin, as well as a thorough revision of the distributions of the taxa in Chile and field samplings of areas poorly studied. Conflicts of interest The authors declare that they have no conflicts of interest. Declaration of funding This research did not receive any specific funding. 51ºS 51ºS Acknowledgements 53ºS 53ºS 55ºS 55ºS 75ºW 73ºW 71ºW 69ºW 67ºW 65ºW Fig. 4. Generalised track in the study area, including the Andean region (Central Chilean, Patagonian and Subantarctic subregions), the Neotropical region and the South American transition zone, based on Morrone (2014a, 2015a, 2015b). We thank Cecilia Ezcurra and Adriana Ruggiero for critical comments on a preliminary version of this work. References Cabrera AL (1971) Fitogeografía de la República Argentina. Boletín de la Sociedad Argentina de Botánica 14, 1–42. 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Zootaxa 4132, 287–289. doi:10.11646/ zootaxa.4132.2.11 Rivas Martínez S (1987) ‘Memoria del Mapa de Series de Vegetación de España.’ (Ministerio de Agricultura, Pesca y Alimentación: Madrid, Spain) Roig-Juñent SA, Griotti M, Domínguez MC, Agrain FA, Campos-Soldini P, Carrara R, Cheli G, Fernández-Campón F, Flores GE, Katinas L, Muzón JR, Neita-Moreno JC, Pessacq P, San Blas G, Scheibler EE, Crisci JV (2018) The Patagonian Steppe biogeographic province: Andean region or South American transition zone? Zoologica Scripta 47, 623–629. doi:10.1111/zsc.12305 Soriano A (1950) La vegetación del Chubut. Revista Argentina de Agricultura 17, 30–66. Handling editor: Malte Ebach 180 A. Padró et al. Australian Systematic Botany Appendix 1. Geographic distribution of the species analysed ar, Argentina; ch, Chile Species Family Distribution 1 Abrotanella trichoachaenia Cabrera 2 Acaena antarctica Hook.f. 3 Acaena leptacantha Phil. 4 Acaena macrocephala Poepp. 5 Acaena magellanica (Lam.) Vahl 6 Adenocaulon chilense Hook. 7 Adesmia corymbosa Clos 8 Adesmia glomerula Clos. 9 Adesmia longipes Phil. 10 Adesmia parvifolia Phil. 11 Adesmia retusa Griseb. 12 Agoseris coronopifolia (d’Urv.) D.M.Moore 13 Agrostis inconspicua Kuntze 14 Agrostis meyenii Trin. 15 Anagallis alternifolia Cav. 16 Anemone multifida Poir. 17 Arenaria serpens Kunth 18 Arjona patagonica Hombr. & Jacq. ex Decne. 19 Arjona pusilla Hook.f. 20 Armeria maritima (Mill.) Willd. 21 Astragalus nivicola Gomez-Sosa 22 Astragalus palenae (Phil.) Reiche 23 Astragalus patagonicus (Phil.) Speg. 24 Austroblechnum microphyllum (Goldm.) Gasper & V.A.O.Dittrich 25 Azorella andina (Phil.) Drude 26 Azorella boelckei (Mathias & Constance) G.M.Plunkett & A.N.Nicolas 27 Azorella crassipes Philippi 28 Azorella echinus (DC.) G.M.Plunkett & A.N.Nicolas 29 Azorella lycopodioides Gaudich. 30 Azorella madreporica Clos 31 Azorella microphylla (Cav.) G.M.Plunkett & A.N.Nicolas 32 Azorella monantha Clos 33 Azorella nivalis Phil. 34 Baccharis magellanica (Lam.) Pers. 35 Baccharis nivalis (Wedd.) Sch. Bip. ex Phil. 36 Baccharis obovata Hook. & Arn. 37 Barneoudia major Phil. 38 Belloa nivea (Phil.) M.O.Dillon 39 Berberis empetrifolia Lam. 40 Blumenbachia prietea Gay 41 Bolax gummifera (Lam.) Spreng. 42 Boopis graminea Phil. 43 Bromus setifolius J.Presl. 44 Calandrinia affinis Gillies ex Arn. 45 Calandrinia caespitosa Gillies ex Arn. 46 Calandrinia colchaguensis Barnéoud 47 Calandrinia graminifolia Phil. 48 Calceolaria biflora Lam. 49 Calceolaria filicaulis Clos filicaulis 50 Calceolaria tenella Poepp. & Endl. 51 Calceolaria uniflora Lam. 52 Caltha appendiculata Pers. 53 Caltha sagittata Cav. 54 Cardamine cordata Barnéoud 55 Carex aphylla Kunth 56 Carex atropicta Steud. 57 Carex banksii Boott 58 Cerastium arvense L. 59 Chaetanthera villosa D.Don Asteraceae Rosaceae Rosaceae Rosaceae Rosaceae Asteraceae Fabaceae Fabaceae Fabaceae Fabaceae Fabaceae Asteraceae Poaceae Poaceae Primulaceae Ranunculaceae Caryoplyllaceae Schoepfiaceae Schoepfiaceae Plumbaginaceae Fabaceae Fabaceae Fabaceae Blechnaceae Apiaceae Apiaceae Apiaceae Apiaceae Apiaceae Apiaceae Apiaceae Apiaceae Apiaceae Asteraceae Asteraceae Asteraceae Ranunculaceae Asteraceae Berberidaceae Loasaceae Apiaceae Calyceraceae Poaceae Portulacaceae Portulacaceae Portulacaceae Portulacaceae Calceolariaceae Calceolariaceae Calceolariaceae Calceolariaceae Ranunculaceae Ranunculaceae Brassicaceae Cyperaceae Cyperaceae Cyperaceae Caryoplyllaceae Asteraceae ar, ch ar, ch ar, ch ar, ch ar, ch ar, ch ar, ch ar, ch ar, ch ar, ch ar, ch ar, ch ar, ch ar, ch ar, ch ar, ch ar, ch ar, ch ar, ch ar, ch ar, ch ar, ch ar, ch ar, ch ar, ch ar, ch ar, ch ar, ch ar, ch ar, ch ar, ch ar, ch ar, ch ar, ch ar, ch ar, ch ar, ch ar, ch ar, ch ar, ch ar, ch ar, ch ar, ch ar, ch ar, ch ar, ch ar, ch ar, ch ar, ch ar, ch ar ar, ch ar, ch ar, ch ar, ch ar, ch ar, ch ar, ch ar, ch (continued next page) Austral High Andean district Australian Systematic Botany 181 Appendix 1. (continued ) Species Family Distribution 60 Chiliotrichum diffusum (G.Forst.) Kuntze 61 Chloraea alpina Poepp. 62 Chloraea magellanica Hook.f. 63 Clinopodium darwinii (Benth.) Kuntze 64 Colobanthus lycopodioides Gris. 65 Combera paradoxa Sandwith 66 Cortaderia pilosa (d’Urv.) Hack. 67 Deyeuxia erythrostachya Desv. 68 Diplolepis biflora (Phil.) Hechem & C.Ezcurra 69 Diplolepis nummulariifolia (Hook. & Arn.) Liede & Rapini 70 Discaria chacaye (G.Don.) Tortosa 71 Draba gilliesii Hook. & Arn. 72 Empetrum rubrum Vahl ex Willd. 73 Ephedra chilensis C.Presl 74 Epilobium australe Poepp. & Hausskn. ex Hausskn. 75 Epilobium nivale Meyen 76 Erigeron andicola DC. 77 Erigeron cinereus Hook. & Arn. 78 Erigeron gilliesii (Hook. & Arn.) Cabrera 79 Erigeron leptopetalus Phil. 80 Erigeron schnackii Solbrig 81 Erythranthe lutea (L.) G.L.Nesom 82 Escallonia alpina Poepp. ex DC. 83 Euphorbia collina Phil. 84 Euphrasia meiantha Clos 85 Festuca acanthophylla E.Desv. var. acanthophylla 86 Festuca pallescens (St.-Yves) Parodi 87 Gamocarpha alpina (Poepp. ex Less.) H.V.Hansen 88 Gamocarpha selliana Reiche 89 Gamochaeta alpina (Poepp.) S.E.Freire & Anderb. 90 Gamochaeta neuquensis Cabrera 91 Gamochaeta nivalis Cabrera 92 Gaultheria caespitosa Poepp. & Endl. 93 Gaultheria pumila (L.f.) D.J.Middleton 94 Gentiana sedifolia Kunth 95 Gentianella magellanica (Gaudich.) Fabris ex D.M.Moore 96 Geranium sessiliflorum Cav. 97 Grindelia prunelloides (Poepp. ex Less.) A.Bartoli & Tortosa 98 Gunnera magellanica Lam. 99 Gutierrezia baccharoides Sch. Bip. 100 Haplopappus grindelioides (Less.) DC. 101 Hordeum comosum J.Presl 102 Hypochaeris arenaria Gaud. 103 Hypochaeris montana (Phil.) Reiche 104 Hypochaeris tenuifolia (Hook. & Arn.) Griseb. 105 Leucheria glacialis (Poepp. ex Less.) Reiche. 106 Leucheria millefolium Dusén & Skottsb. 107 Leucheria papillosa Cabrera 108 Lobelia oligophylla (Wedd.) Lammers 109 Luzula chilensis Nees & Meyen ex Kunth 110 Luzula racemosa Desv. 111 Marsippospermum grandiflorum (L. f.) Hook. 112 Marsippospermum philippii (Buchenau) Hauman 113 Marsippospermum reichei Buchenau 114 Menonvillea nordenskjoeldii (Dusén) Rollins 115 Menonvillea rigida Rollins 116 Montiopsis cistiflora (Gillies ex Arn.) D.I.Ford 117 Montiopsis gayana (Barnéoud) D.I. Ford 118 Montiopsis polycarpoides (Phil.) Peralta 119 Montiopsis umbellata (Ruiz & Pav.) D.I.Ford 120 Moschopsis caleofuensis (Speg.) Dusén Asteraceae Orchidaceae Orchidaceae Lamiaceae Caryoplyllaceae Solanaceae Poaceae Poaceae Apocynaceae Apocynaceae Rhamnaceae Brassicaceae Empetraceae Ephedraceae Onagraceae Onagraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Calceolariaceae Euphorbiaceae Fabaceae Calceolariaceae Poaceae Poaceae Calyceraceae Calyceraceae Asteraceae Asteraceae Asteraceae Ericaceae Escalloniaceae Gentianaceae Gentianaceae Gerianiaceae Asteraceae Gunneraceae Asteraceae Asteraceae Poaceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Campanulaceae Juncaceae Juncaceae Juncaceae Juncaceae Juncaceae Brassicaceae Brassicaceae Portulacaceae Portulacaceae Portulacaceae Portulacaceae Calyceraceae ar, ch ar, ch ar, ch ar, ch ar, ch ar, ch ar, ch ar, ch ar, ch ar, ch ar, ch ar, ch ar, ch ar, ch ar, ch ar, ch ar, ch ar, ch ar, ch ar, ch ar ar, ch ar, ch ar, ch ar, ch ar, ch ar, ch ar, ch ar, ch ar ar, ch ar, ch ar, ch ar, ch ar, ch ar, ch ar, ch ar, ch ar, ch ar, ch ar, ch ar, ch ar, ch ar, ch ar, ch ar, ch ar, ch ar, ch ar, ch ar, ch ar, ch ar, ch ar, ch ar, ch ar, ch ar ar, ch ar, ch ar, ch ar, ch ar (continued next page) 182 A. Padró et al. Australian Systematic Botany Appendix 1. (continued ) Species Family Distribution 121 Moschopsis rosulata (N.E.Br.) Dusén 122 Moschopsis subandina (Speg.) Dusén 123 Nardophyllum bryoides (Lam.) Cabrera 124 Nassauvia aculeata (Less.) Poepp. & Endl. 125 Nassauvia argentea Phil. 126 Nassauvia argyrophylla Speg. ex. Cabrera 127 Nassauvia darwinii (Hook. & Arn.) Hoffm. & Dusén 128 Nassauvia dentata Griseb. 129 Nassauvia digitata Wedd. 130 Nassauvia glomerata (Gill.) Wedd. 131 Nassauvia lagascae (D.Don) F.Meigen 132 Nassauvia planifolia Wedd. 133 Nassauvia pulcherrima Cabrera 134 Nassauvia pygmaea (Cass.) Hook. f. 135 Nassauvia revoluta D.Don 136 Nastanthus patagonicus Speg. 137 Nastanthus scapigerus (J.Remy) Miers 138 Nicoraepoa andina (Trin.) Soreng & L.J.Gillespie chonotica 139 Noccaea magellanica (Comm. ex Poir.) Holub 140 Ochetophila nana (Clos) J.Kellerm., Medán & Aagesen 141 Olsynium frigidum (Poepp.) Goldblatt 142 Olsynium junceum (E.Meyer ex Presl.) Goldblatt 143 Onuris graminifolia Phil. 144 Oreobolus obtusangulus Gaudich. 145 Oreopolus glacialis (Poepp.) Ricardi 146 Ourisia alpina Poepp. & Endl. 147 Ourisia breviflora Benth 148 Ourisia fragrans Phil. 149 Ourisia ruelloides (L.f.)Kuntze 150 Oxalis adenophylla Gillies ex Hook & Arn. 151 Oxalis erythrorhiza Gillies ex Hook. et Arn. 152 Oxalis magellanica G.Frost. 153 Oxychloë bisexualis Kuntze 154 Patosia clandestina (Phil.) Buchenau 155 Perezia bellidifolia (Phil.) Reiche 156 Perezia capito (Phil.) Reiche 157 Perezia delicata Vuilleumier 158 Perezia fonckii (Phil.) Reiche 159 Perezia lactucoides (Vahl) Less. 160 Perezia linearis Less. 161 Perezia lyrata (J.Rémy) Wedd. 162 Perezia pedicularidifolia Less. 163 Perezia pilifera (D.Don) Hook. & Arn. 164 Perezia prenanthoides Less. 165 Perezia recurvata (Vahl) Less. 166 Pinnasa nana (Phil.) Weigend & R.H.Acuña 167 Plantago barbata G.Forst. 168 Plantago uniglumis Wallr. ex Walp. 169 Poa holciformis J.Presl 170 Poa obvallata Steud. 171 Polygala salasiana Gay 172 Polystichum andinum Phil. 173 Pozoa coriacea Lag. 174 Pozoa volcanica Mathias & Constance 175 Primula magellanica Lehm. 176 Quinchamalium chilense Molina 177 Ranunculus peduncularis Sm. 178 Rhodolirium andicola (Poepp.) Ravenna 179 Rhodophiala mendocina (Phil.) Ravenna 180 Rytidosperma pictum (Nees & Meyen) Nicora 181 Rytidosperma virescens (Desv.) Nicora Calyceraceae Calyceraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Calyceraceae Calyceraceae Poaceae Brassicaceae Rhamnaceae Iridaceae Iridaceae Brassicaceae Cyperaceae Rubiaceae Calceolariaceae Calceolariaceae Calceolariaceae Calceolariaceae Oxalidaceae Oxalidaceae Oxalidaceae Juncaceae Juncaceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Loasaceae Plantaginaceae Plantaginaceae Poaceae Poaceae Polygalaceae Aspidiaceae Apiaceae Apiaceae Primulaceae Santalaceae Ranunculaceae Amaryllidaceae Amaryllidaceae Poaceae Poaceae ar, ch ar, ch ar, ch ar, ch ar, ch ar ar, ch ar, ch ar, ch ar, ch ar, ch ar ar, ch ar, ch ar, ch ar ar, ch ar, ch ar, ch ar, ch ar, ch ar, ch ar, ch ar, ch ar, ch ar, ch ar, ch ar, ch ar, ch ar, ch ar, ch ar, ch ar, ch ar, ch ar ar, ch ar ar, ch ar, ch ar, ch ar, ch ar, ch ar, ch ar, ch ar, ch ar ar, ch ar, ch ar, ch ar, ch ar, ch ar, ch ar, ch ar, ch ar, ch ar, ch ar, ch ar, ch ar ar, ch ar, ch (continued next page) Austral High Andean district Australian Systematic Botany 183 Appendix 1. (continued ) Species Family Distribution 182 Sanicula graveolens Poepp. ex DC. 183 Saxifraga magellanica Poir. 184 Schoenus andinus (Phil.) H.Pfeiff. 185 Senecio argyreus Phil. 186 Senecio aspericaulis J.Rémy 187 Senecio baccharidifolius DC. 188 Senecio boelckei Cabrera 189 Senecio carbonensis C.Ezcurra, Ferreyra & Clayton 190 Senecio chilensis Less. 191 Senecio chionophilus Phil. 192 Senecio covasii Cabrera 193 Senecio crithmoides Hook. & Arn. 194 Senecio diemii Cabrera 195 Senecio fistulosus Poepp. ex Less. 196 Senecio gilliesii Hook. & Arn. 197 Senecio gnidioides Phil. 198 Senecio hieracium J.Rémy 199 Senecio jobii Cabrera 200 Senecio kingii Hook.f. 201 Senecio linariifolius Poepp. ex DC. 202 Senecio magellanicus Hook. & Arn. 203 Senecio pachyphyllos J.Rémy 204 Senecio parodii Cabrera 205 Senecio peteroanus Phil. 206 Senecio poeppigii Hook. & Arn. 207 Senecio portalesianus J.Rémy 208 Senecio subdiscoideus Sch. Bip. ex Wedd. 209 Senecio subumbellatus Phil. 210 Senecio trifurcatus (G.Forst.) less. 211 Senecio triodon Phil. 212 Silene andicola Gill. ex Hook. et Arn. 213 Silene chilensis (Naud.) Bocquet 214 Silene cuspidata Pedersen 215 Sisyrinchium arenarium Poepp. 216 Sisyrinchium macrocarpum Hieron. 217 Solenomelus segethii (Phil.) Kuntze 218 Symphyotrichum glabrifolium (DC.) G.L.Nesom 219 Trisetum barbinode Trin. 220 Tristagma nivale Poepp. 221 Tristagma patagonicum (Baker) Traub 222 Valeriana boelckei Rossow 223 Valeriana carnosa Sm. 224 Valeriana fonckii Phil. 225 Valeriana macrorhiza DC. 226 Valeriana philippiana Briq. 227 Vicia bijuga Gillies ex Hook. et Arn. 228 Viola columnaris Skottsb. 229 Viola cotyledon Ging. 230 Viola dasyphylla Becker 231 Viola maculata Cav. 232 Weberbauera colchagensis (Barnéoud) Al-Shehbaz Apiaceae Saxifragaceae Cyperaceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Asteraceae Caryoplyllaceae Caryoplyllaceae Caryoplyllaceae Iridaceae Iridaceae Iridaceae Asteraceae Poaceae Alliaceae Alliaceae Valerianaceae Valerianaceae Valerianaceae Valerianaceae Valerianaceae Fabaceae Violaceae Violaceae Violaceae Violaceae Brassicaceae ar, ch ar, ch ar, ch ar, ch ar, ch ar, ch ar ar ar, ch ar, ch ar, ch ar, ch ar ar, ch ar, ch ar, ch ar, ch ar ar, ch ar, ch ar, ch ar, ch ar, ch ar, ch ar, ch ar, ch ar, ch ar, ch ar, ch ar, ch ar, ch ar, ch ar, ch ar, ch ar ar, ch ar, ch ar, ch ar, ch ar, ch ar, ch ar, ch ar, ch ar, ch ar, ch ar, ch ar ar, ch ar ar, ch ar, ch 184 Australian Systematic Botany A. Padró et al. Appendix 2. Data matrix for the parsimony analysis of endemicity Each 0 (absence) or 1 (presence) represents 1 of the 232 species, as per the order listed as in Appendix 1, that are present within the numbered cells (C1–C196) in Fig. 2 (e.g. only one species, Berberis empetrifolia Lam., is found in cell C193) Root 00000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000 00000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000 C1 01001000000000000001000000001000000000001000000000000000110100000000000100000000000000000010100001000000000000001000000 00010000000000000000000010000100000000000000000000000001000000010000000000000000000000000000000000000000000000000 C2 01001000000000000001000000001000000000001000000000000000110100000000000100000000000000000010100001000000000000001000000 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00000000000000000000000000000000000000000000000000000000000000001000000000000000000000000000000000000000000000000 C21 01000000000000000001010000000001000000100000000000100000010000000000000000000000000000000001000000000000000000001000000 00010000000000000000000010000100000000000000010100000000000001000000000000000000100000000000000000000001000100000 C22 01001000000000010101000000000000000000000000000100100000010000000000010000000000000000000000101000000000000000101000000 00000000000000000000000000010100000000000000010000001001000001010000000000000000001000000000000000000001000000010 C23 01001000000000010101000000001000000000000000000100100000100100010000010101000010000000000000101001000000000001000000000 01000000000000000000000000010100000000000000000000001001000000010000000000000000001000000000000000000000000000010 C24 0000000000000000000100000000100001000000000000010010100010010001001000010000001000000000001010100100000000000100000 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(continued ) C26 0000000000000000000000000000000000000000000000010000000000000000000000000000000000000000000010100000000000000000000 000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000 C27 0000000000000000000000000000000000000000000000010000000000000000000000000000000000000000000010000000000000000000000 000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000 C28 0000000000000000000101000000000100000010000000000000000000000000000000000000000000000000000100000000000000000000000 000000010000000000000000000010000000000000000000000100000000000000000000000000000000100000000000000000001000000100000 C29 0000000000000000000100000000000100000000000000000000000001000000000000000000000000000000000010000000000000000010100 000000000000000000000000000000000000000000000000010000000000000001000000000000000000000000000000000000000001000000000 C30 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0100000000000000000100000000100001000000000000000100100010010001000000000000001000000000000000100000000000000000100 000000000000000000000000000000000000000000000000000000000000000000011100000000000000000000000000000000000000000000000 (continued next page) 186 Australian Systematic Botany A. Padró et al. Appendix 2. (continued ) C52 0000000000000000000000000000000001000000000000000000000000000000000000000000000000000000000000000000000000000000000 000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000 C53 0000000000000000000000000001000001000000000000000000000000000000000000000000000000000000000000000000000000000000000 000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000 C54 0000000000000000010001000001000001000000000000000000000000000000000000001000000000000000000000000000000000000000000 000000000000000000000000001000000000000000000001000000000000000000000000000000000000000000000000000000000000000000000 C55 00000010000000000101010000010000010000100000000000000000000000000000010010000000010000000001100000000000000000000000 00000000000000000001000000000000000000000000000000100000000000000000000000000000000000000000000000000000000000100000 C56 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00000000000000000000000000000000000000000000000001000000000000000000000000000000000000000000000000000000000000001000 00000000000000000000000000000000000000000000000000000000000000000001100000000000000000000000000000000000000000000000 (continued next page) Austral High Andean district Australian Systematic Botany 187 Appendix 2. (continued ) C79 00000000000000000100000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000 00000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000 C80 00000010000000000100000000000000000000000000000000000000000000000000000000000000001000000000000000000000000000000000 00000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000 C81 00000010000000000000000000010000000000000000000000000000000000000000000000000000001000000000000000000000000000000000 00000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000 C82 00000000000000000001000000010000000000000000000000000000000000000000000000000000000000000000000000000000000000000000 00000000000000000000000001000000000000000000001000000000000000000000000000000000000000000000000000000000000000000000 C83 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Padró et al. Appendix 2. (continued ) C109 0000000000000000000100000000000000000000000000000000000000000000000000000000000000000100000000000000000000000000000 000000000000000000001000000100000000000000000000000000000000000000000000000000000000000000000000000000000000000000000 C110 0000000000000000000100000000001000000000000000000000000000000000000000000000000000000100000000000000000000000000000 000000000000000000001000000100000000000000000000000000000000000000000000000000000000000000000000000000000000000000000 C111 0000000000000000000000000000001000000000001000000000000000000000000000000000000000000100000000000000100000000000000 000000000000000000001000000100000000000000000000000000000000000000000000000000001000000000000000000000000000000000000 C112 0000000000000000000000000000001000000000001000000000000000000000000000000000000000100100000000000000100000000000000 000000000000000000001000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000100000 C113 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0000000000000000000000000000000000000000000000000000000000000000000000000000000000100000000000000000000000000000000 000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000 (continued next page) Austral High Andean district Australian Systematic Botany 189 Appendix 2. (continued ) C151 0000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000 000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000100000000000 C152 0000000000000000000000000000000000000000000010000000000000000000000000000000000000000000000000000000100100000000000 000000000000000000000000000000000000000000000000000000000000000000000000000000001000000000000000000000000100000100000 C153 01001010110100011101101100111101110010110011101010001001111101111011111111111011111100110110101110011011000011000001 11100011000001100111111111100100111011000000101010111010001011111010010001110001100010101010101011110010100110011011 C154 0011101011001110100100010010100011010111010101000000111111010001011111110111100111111010011111111100100110011101100 010100111000011110010001111100110000011011000111000100101101111101101011001101010111010101111001011001111000111011000 C155 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0000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000100000000000 000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000 (continued next page) 190 A. Padró et al. Australian Systematic Botany Appendix 2. 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