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Areas of endemism in the southern central Andes.

INTRODUCTION

Located in the centre of the South American dry diagonal, the southern central Andes have received relatively little attention in terms of quantitative phytogeographic studies. At a continental scale, hotspots and areas of endemism have mainly been defined and explored in more humid portions of the Andes, such as the northern Andes or northern central Andes (e.g. Kier et al., 2005; Orme et al., 2005; Sklenar et al., 2011) and in the southern Andes (Arroyo et al., 1999, 2004; Rodriguez-Cabral et al., 2008).

Within the southern cone of South America the endemic vascular flora is concentrated in the well documented Chilean hotspot (Arroyo et al., 1999, 2004; Bannister et al., 2012 and references therein). However, the arid southern central Andes appear to be the second most important area of endemism, as checklists and catalogues show that north-western Argentina contains far more endemic species than the remaining of the region including Uruguay, southern Brazil, and southern Paraguay (Zuloaga et al., 1999, 2008).

During the Tertiary, the central Andes became relatively dryer than the northern and southern part (Simpson & Todzia, 1990). Comparative floristic studies along the Andes have identified the flora of the southern central Andes as a separate unit among other arid plant formations (Lopez et al., 2006) and found more similarities at generic and family level between the high Andean flora of the humid northern and southern Andes than between these and the high Andean flora of the dry central Andes (Simpson & Todzia, 1990). Summer rainfall is sufficiently high on the eastern slopes to support humid subtropical Yungas forest that is one of the most diverse habitats within the southern cone (Brown, 1995). However, no quantitative studies have examined whether endemism in the southern central Andes is related to the Yungas forest or found within the more arid vegetation on the inner slopes and valleys.

In their biogeographic analysis of South America, Cabrera & Willink (1973) and Cabrera (1976) emphasised the distinctiveness of the slope vegetation in the southern central Andes by differentiating the vegetation between 2400-3500 m asl as the Prepuna phytogeographic province. The Prepuna was originally restricted to north-western Argentina (Cabrera, 1976), but later expanded to include the dry inter-Andean valleys of southern Bolivia (Lopez, 2000, 2003). Since the phytogeographic units in the traditional scheme of Cabrera are based on the presence of endemic taxa (Cabrera, 1951, 1953; Ribichich, 2002) it is reasonable to expect that at least part of the endemic species are distributed along the Prepuna, therefore defining the limits of this phytogeographic unit.

Here we explore the distribution pattern of the vascular plant species endemic to the southern central Andes including the Prepuna province in its entire extension. Defining areas of endemism is considered fundamental for historical and ecological biogeography (Crisp et al., 2001), but also for conservation studies especially if endemism is found out of the high diverse vegetation types as protected area systems focus on diversity hotspots (Orme et al., 2005). Our main aims are to explore how the endemic species are distributed within the southern central Andes, identify areas of endemism, and examine to which extent these main areas of endemism correspond to pre- established phytogeographic units such as the Prepuna slope vegetation, or found within the high diverse Yungas forest. We used the optimality criterion developed by Szumik et al. (2002) and Szumik & Goloboff (2004) and implemented in the program NDM/VNDM (Goloboff, 2005) to analyze the distribution of the species endemic to the north-western Argentina and south-western Bolivia.

MATERIALS AND METHODS

Study Region

The study region comprises all west Argentinean provinces between the Bolivian border and ~32[degrees]S as well as the adjacent southern Bolivia departments of Tarija, Chuquisaca, and Potosi (see maps in Fig. 1-2). The Chaco basin towards the east (Fig. 1) and the hyper arid northern Chile to the west make natural borders of the study region while the southern limit follows the northern limit of the Patagonian biogeographic province (Cabrera, 1979; Cabrera & Willink, 1973; Morrone, 2001). The northern limit follows Lopez (2000) and Lopez & Beck (2002), authors who analyzed the floristic composition of the Bolivian Prepuna and stressed the floristic relationship between the southern Bolivia and the north-western Argentina (see also Ibisch et al., 2003). The study region comprises approx. 720.600 [km.sup.2].

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Rainfall mainly occurs during the southern hemisphere summer where the South American Monsoon System (SAMS) brings moist air to the region (Zhou & Lau, 1998). The influence of the SAMS decreases by an east-west and north-south gradient (Fig. 2A). In the southernmost part of the study region rainfall due to the SAMS is low while winter rain is higher due to the influence of the Pacific westerlies that brings winter rain to central Chile (Montecinos & Aceituno, 2003).

Traditionally the study region has been divided in a phytogeographic scheme developed by Cabrera (1951, 1953, 1976) and Cabrera & Willink (1973-see Fig. 1) that is widely used by South American biologists, as well as by governmental and non-governmental organizations (Ribichich, 2002). The study region includes xerophytic Chaco vegetation in north-eastern lowlands, and arid Monte vegetation in the south-eastern lowlands. Subtropical moist broadleaf forests, locally known as Yungas, are found at the eastern slopes north of 28[degrees]S. The Andes vegetation is furthermore divided in Puna vegetation between approx. 3500m-4500m above which extends the high Andean grasslands that reaches the line of perpetual snow. The Prepuna province--which is the only phytogeographic unit confined to the study region--extends north-south through the region on the dry mountain slopes approx. between 2400 m and 3500 m asl (Fig. 1 and 2B). The present study follows the phytogeographic scheme of Cabrera (1951, 1953, 1976), and that of Ibisch et al. (2003) for the Yungas forests. Ibisch et al. (2003) recognized the north-south oriented Yungas forests that are found within our study region as the Tucumano-Bolivian Yungas. Due to differences in temperature, precipitation seasonality, and taxonomic composition the Tucumano-Bolivian Yungas were considered a phytogeographic unit distinct from the more tropical northwest-southeast oriented Bolivian-Peruvian Yungas forests found north of the Andes bend (~18[degrees]S).

Data set

According to the catalogue of the Southern Cone flora (Zuloaga et al., 2008) 589 species of vascular plants are strictly endemic to the Argentinean portion of the study region which is approx. 1/3 of the species endemic to Argentina in general (Zuloaga et al., 1999). We compiled the distribution of 513 endemic species excluding 40 species that were only known from localities that could not be georeferenced with precision, while 36 species were excluded as they were found outside the study region while we were compiling their distribution. In addition to species endemic to north-western Argentina we added 27 species endemic to the Bolivian Prepuna or shared between the Bolivian Prepuna and north-western Argentina. These species were selected with assistance from Dr. Beck (the National Herbarium, La Paz, Bolivia) as a checklist of vascular plants of Bolivia has not been completed yet. All endemic species as well as author names are found in Table 2.

Species distribution were compiled from collections deposited in the herbaria BA, BAA, LIL, LP, LPB, and SI. We considered a species sufficiently sampled when the distribution data reflected the distribution published in the Catalogue of the Plants of the Southern Cone (Zuloaga et al., 2008). Redundant locations were only georeferenced if the collections differed sufficiently in altitude to suggest that the species appears in several phytogeographic strata. To facilitate the task of georeferencing, specimens with altitude noted in the field by the collector were preferred. If the herbarium vouchers were not sufficient to cover the published distributions we added published exsiccatae from the original descriptions or monographies. Finally we used the Missouri Botanical Garden data base TROPICOS(r) (http://www.tropicos.org/) to complete distribution data or to exclude species collected outside the study region.

Distribution data without corresponding vouchers deposited in public herbaria were only used for some Bolivian endemics, where we added field observations from R. P. Lopez, and also when compiling species distribution within the Cactaceae. The Cactaceae are particularly species rich in the study region (Mourelle & Ezcurra, 1996) but also one of the most underrepresented families in herbaria; as a result, specimens deposited in public herbaria are far from covering the distribution of an individual species. Since the Cactaceae is one of the main targets for commercial collectors the information on collection sites, found at Cactus-enthusiastic homepages, is overwhelming and in sharp contrast with the sparse information available from traditional academical sources. We choose to include information from a private collector homepage (http://ralph.cs.cf.ac.uk/Cacti/finder.html). This procedure was done after consulting a specialist on the family, Dr. Roberto Kiesling, in order to avoid species that are not easily recognized in the field. Data from this source were also considered only when the location, and altitude, agrees with the distribution range defined in the Catalogue of the Plants of the Southern Cone (Zuloaga et al., 2008).

A total of 3262 records were compiled and georeferenced. This gives an unimpressive average of six registers/species, which both reflect that species were not georeferenced, in the study, for redundant or proximate locations, and that endemic species of north-western Argentina and southern Bolivia are usually rare and poorly collected, many of them being only known from the type material. All specimens without GPS recorded coordinates were georeferenced according to the point-radius method of Wieczorek et al. (2004). We carefully respected the altitudes of the collection sites in order to extract climate data with as much accuracy as possible. All data have been submitted to the online version of Flora Argentina (http://www.floraargentina.edu.ar/).

Analyses

We searched for areas of endemism using the program NDM/VNDM ver. 2.7c (Goloboff, 2005). VNDM is a grid based method that identifies an area of endemism as the congruent distributions of two or more species (Szumik et al., 2002; Szumik and Goloboff, 2004, 2007). The general outputs in VNDM are area sets (groups of cells in the grid) that are supported by the presence of two or more species. Each area receives an endemism-score according to the optimality criterion developed for the method. The optimality criterion measures the 'fit' of each species to the area, e.g. how well each species found within the area adjust to this. The optimality criterion penalizes both absence in part of the area as well as presence in adjacent cells outside the area (if a species is present outside the area in a non-adjacent cell it is not considered among the species supporting the area). The endemism score for each area is the sum of the 'fit' of the supporting species. The endemism score is therefore influenced both by the number of species supporting an area as well as the distribution of the supporting species within and around the area (Szumik et al., 2002; Szumik and Goloboff, 2004). The algorithms in NDM have lately been found to outperform other and more widespread methods for delimiting areas of endemism such as hierarchical clustering (Carine et al., 2008; Escalante et al., 2009). One of the advantages of NDM is its ability to recognize overlapping distribution patterns if these are defined by different species groups. Overlapping patterns may be independent if defined by different sets of species (Szumik & Goloboff, 2004), and are to be expected when distribution analysis are based on grids if more than one environments are found in the same cell.

Grid and cell size. As discussed by Linder (2001) the choice of grid size is important. The use of small cells would result in a finer and more detailed resolution but at the same time increase the number of artificially empty cells where species occur but have not been recorded. We used three different cell sizes to explore both distribution patterns at different scale as well as the robustness of the resulting areas to changes in grid size (Aagesen et al., 2009; Navarro et al., 2009). Grids with cell sizes 0.2[degrees]x0.2[degrees], 0.5[degrees]x0.5[degrees], and 0.5[degrees]x1.0[degrees]were used where 1[degrees]is approximately equal to 100 km. In the last grid we used rectangular cells rather than square cells because the north-south running Andes Mountains form a steep east-west altitude gradient that causes various habitat changes within 100 km. The rectangular cells were used to explore north-south running patterns of widely distributed but poorly collected species without lumping too many habitats into the same cell.

Fill option. The problem of artificial empty cells is somewhat alleviated in NDM by a user defined fill option (radius size) that assumes the presence of a given species if it has been collected close to the limit within a neighbouring cell (Szumik & Goloboff, 2004; Aagesen et al., 2009; Escalante et al., 2009). We used two radius sizes (see Table 1). The smallest radius size reflects approximately the error of the hand georeferenced records as determined by the point-radius method (Wieczorek et al., 2004; see Aagesen et al., 2009) while the second and wider radius size was used to explore dependence on the fill options in the final areas of endemism.

Search procedure. The searches were done using default settings with the following changes: swap two cells at a time; discard superfluous sets as they are found; replace a set if improved during swapping; pre-check duplicates; keep overlapping subsets if 20% of species unique. Searches were done by changing the seed for each search, without replacing existing sets and deleting duplicate sets after each replicate. This search sequence was repeated until numbers of sets were stable.

Consensus rules. As we kept all overlapping subsets if they differed by 20% in species composition, we found several area sets that were supported by 5-20 species but differing only by one or two species. Area sets can be merged into consensus areas if they share at least some of their defining species (Szumik & Goloboff, 2007). Two consensus rules are available in VNDM. The strictest consensus rule merges area sets if they all share a user defined percentages of their defining species --(in the program called 'against every included area' hereafter abbreviated as consensus rule ae). The second more relaxed criterion includes an area set in a consensus if it shares a user defined percentage of its defining species with at least one other area set in the consensus--(called 'against any included area' hereafter abbreviated as consensus rule aa; see Aagesen et al., 2009).

Ideally, the strict consensus rule ae should be used for identifying areas of endemism as at least some species are shared among all the individual area sets, thus ensuring a certain coherency within the area. Consensus areas derived from the laxer consensus rule aa, are efficient to outline gradual species overlap between area set, but the distribution of the species defining the consensus are likely to be found in just a small part of this. However, due to the high number of ae consensus produced by our data (Table 1) it was unfeasible to base the discussion on the ae consensus areas only. Consequently, we explored areas of endemism through the laxer aa rule that produced less, and more inclusive consensus areas.

Environmental data and aridity

Environmental data for the study region that includes both Argentina and Bolivia is limited to the data available in WorldClim (Hijmans et al., 2005). We used the program Diva-Gis 7.4.0.1 (Hijmans et al., 2005-2010) to extract altitude, temperature, and precipitation of the collection sites. As the collection sites span from 500 m asl to 5000 m asl and an annual mean temperature from -3[degrees]C to 23[degrees]C we used humidity rather than rainfall when comparing different sites. We adopted the aridity index of De Martonne (1927) that can be calculated through the limited environmental variables available for the study region, and furthermore provides a quantitative measure for the useful but somewhat imprecise terms arid, semi-arid, humid, etc.

Aridity Index of de Martonne. Accumulated annual precipitation / (annual mean temperature + 10) We use the following categories (De Martonne, 1927; Almorox, 2003): 0-5: desert; 5-10: semi desert; 10-20: semi arid; 20-30: subhumid; 30-60: humid; >60: wet. Distribution of aridity within the study region is shown in Fig. 2C.

RESULTS

The results are summarized in Table 1-3. In general, the number of area sets increased with cell size, but the bigger cells also produced more overlapping distribution patterns that grouped into fewer consensus areas under the laxer aa rule (Table 1). The two different fill radius produced nearly identical results in number of area sets and consensus areas, hence support for the resulting areas did not depend on a specific fill ratio. The discussion is based on the analyses using the widest fill ratios.

Five hundred-nineteen (519) species supported some area under at least one grid size, while 21 species did not support any area (Table 2). Of these, eight species are only known from a single or two localities, while the remaining species are widely distributed within the region. The study region has been sufficiently sampled to analyse species distribution under cell sizes 0.5[degrees]x0.5[degrees]where most cells of the study region are assigned to one or more areas of endemism (Fig. 3A). When the cell size is reduced to 0.2[degrees]x0.2[degrees]only the cores of the main areas are recovered even if presence of one or more endemic species is recorded in most cells (Fig. 3B). Consequently, we base our discussion on the results from analysing cells of 0.5[degrees]x0.5[degrees]. The two alternative cell sizes with either smaller or longer cells were only used to explore the distributions of species that did not support distribution patterns under the 0.5[degrees]x0.5[degrees]grid.

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When using a grid size of 0.5[degrees]x0.5[degrees], the lax aa consensus rule produced seven distribution patterns, under both the 5% and 10% consensus criterion (Table 1). About 2/3 of the endemic species were gathered in a single main consensus area (Fig. 4A), while five distinct distributions, with a total of 49 species, did not form part of the main pattern (see discussion below). A single area consisting of four species was considered an artefact and not discussed any further (the species from this pattern are considered part of the complex Jujuy-Tucuman area, see Discussion and Appendix). The remaining 120 endemic species did not support any area under this grid size.

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In order to explore sub-areas of the main consensus area in Fig. 4. A this was decomposed by gradually increasing the consensus criterion. By this procedure sub-areas separate successively from the main area with those defined by less overlapping species separating first. We decomposed the main area both to extract the individual high endemic areas as well as to assign all endemic species to sub-areas with a closer association between area extension and distributions of the defining species (see, Consensus rules). All final sub-areas (Table 3, Fig. 4) are also found under the strictest ae consensus rule.

A total of 17 areas of endemism are discussed below. The 120 species that did not support areas in the 0.5[degrees]x0.5[degrees]grid were assigned to one of these 17 areas according to results found under the 0.2[degrees]x0.2[degrees]or 0.5[degrees]x1.0[degrees]grids (Table 2). Two areas are based exclusively on results obtained by analysing cells sizes of 0.5[degrees]x1.0[degrees]; these areas were not found under the smaller cell sizes. Only 21 species did not support any of the final areas.

We have named areas of endemism according to the political division in which the core of the area occurs. The political names do not indicate that the areas fit any political border or that the political borders have influenced the analysis. The names are simply useful for a quick spatial orientation. In some cases, such as Famatina and Ambato, the political names coincide with the mountain range in which the endemic species from the core areas have been found. In other cases such as Jujuy and Tucuman several mountain ranges and valleys are included in the core of the areas with some of these situated in the neighbouring province Salta.

Even when subdividing the main area into subareas, most of the defining species will be distributed only in part of the area. One example is Aphelandra lilacina that has only been collected in 'El Rey' National Park in the province of Salta. The cell including 'El Rey' National Park lies in the south easternmost low endemic part of the Jujuy border area (Fig. 4C). Consigning Aphelandra lilacina to the Jujuy area is therefore slightly misleading, but a necessity unless a prohibited high number of consensus areas are shown. However, as our aim is to provide an overview of the distribution of the endemic species in north-western Argentina we concentrate the discussion on main distribution patterns. Further details of the individual species distribution can be consulted in the online version of the Flora Argentina (http://www.floraargentina.edu.ar/) that includes dot distribution maps and label information for all specimens included in the present study.

DISCUSSION

The vast majority of the vascular flora endemic to the southern part of the central Andes outlines small, successively overlapping sub-areas that combine into a single main consensus area only under the lax consensus rule and a very lax consensus criterion (Fig. 4A). Endemism is far from evenly distributed among the sub-areas but declines gradually towards the south and west of the study region. Peaks of endemism are found near the humid Andes slopes in Jujuy, Tucuman/Ambato, and to a lesser extent in the isolated Sierra de Famatina in La Rioja.

Although the main portion of the endemic species conform to the patchy distribution patterns that form the consensus in Fig. 4A, the southern part of the central Andes can still be recognized as a single area of endemism, defined by at least 53 species that are widely distributed within the region (the NOA area in Fig. 4B).

Here we discuss taxonomic composition and habitat of the endemic species from the study region in general, as well as the individual main distribution areas such as the general NOA pattern and sub- areas of the main consensus in Fig. 4A. Sub-areas were separated from the main consensus by gradually raising the consensus criterions, which causes sub-areas to separate successively. A total of 18 different sub-areas were identified (Table 2-3) among which we discuss selected high-endemic areas. A description of the remaining areas is found in Appendix 1.

Altitude and Aridity

Our results indicate that 473 of the endemic species, nearly 90%, have been collected at middle altitudes between 1500-3500 m asl (Table 2), a pattern that is consistent with the notion that Prepuna slope vegetation harbours the main part of the endemic species. Only 40 species are restricted to altitudes below 1500 m asl while 33 species are restricted to high Andean environments above 4000 m asl (mainly from the genera Senecio L., Nototriche Turcz., and Viola L.). The relative low number of endemic high Andean species is perhaps unexpected and may be underestimated as collection of high Andean localities above 4000 m asl are sparse. However, part of the high Andean flora were left out of the analyses as it is shared with the neighbouring Chile.

In accordance with the general arid climate of the central Andes, the semi-arid environments (AI=10- 20, Fig. 2C) appear as the principal habitat type for the endemic species. Semi-arid environments harbour about 80% of the endemic flora with 30% (159 species) being restricted to this habitat type. Semi-arid environment are mostly extended at lower altitudes (compare Fig. 1 and 2A), but only 14 endemic species are restricted to altitudes below 1000 m asl (Table 2). Consequently the relatively narrow band of semiarid habitats between 1500-3500 m asl (compare Fig. 2 B and C) includes the main portion of the endemic species, making the topographic complex plateau, slope, and valley system of the southern central Andes the main locations for endemism.

As few as 36 species are restricted to subhumid or humid environments (AI=20-60, Fig. 2C) indicating that the diverse Tucumano-Bolivian Yungas forest is surprisingly poor in endemic species at a local scale. This is in accordance with Ibisch et al. (2003) who considered the endemism of the Tucumano- Bolivian Yungas to be unimportant at a local Bolivian scale and moderate at the inclusive scale considering the Tucumano-Bolivian Yungas forests in their entire extension. The Bolivian-Peruvian Yungas forests were in contrast considered the main area of endemism for Bolivia. Endemism in the south-western Bolivia were, as in the present study, mainly found in the Prepuna vegetation and in the arid interandean forests (Ibisch et al., 2003). In this study the mountain grasslands above the tree line are rich in local endemics [note that these grasslands were considered the highest strata of the Yungas by Cabrera (1976), while we follow Ibisch et al. (2003) who included these grasslands in the subhumid Puna restricting the term Tucumano-Bolivian Yungas to forest vegetation].

Local endemics are also nearly lacking from desert environments with only seven species found exclusively in habitats with an aridity index below 5 (Table 2). Although desert environments are found throughout the study region, they become part of the distribution pattern only in the areas that include patches of desert climate separated from the Andes (i.e. in the Catamarca, La Rioja, and San Juan provinces--Table 3, Fig. 2C). Only a single species, Gentianella punensis, is collected above 3500 m asl where it is endemic to vegas in the deserts of northern Salta. Dry adapted endemic desert species are consequently restricted to desert parches of middle or low altitudes, such as the inner basins of the Andes that are covered by Monte vegetation. High Andean desert environments are common on both sites of the Andes in the northern part of the study region (compare Fig. 2 B and C), hence desert endemics may be lacking from the northern areas because the endemic species are shared between Argentina and Chile, hence excluded from our analyses.

Taxonomic groups

The Asteraceae outnumbers all other families in the region, with 121 endemic species from 35 genera (Table 2 and 4); furthermore, it is the only family with endemic species supporting all areas (except Valle Fertil--Fig. 4R). The endemic Asteraceae are found in all environments ranging from species restricted to humid habitat (e.g., Mikania siambonensis and Vernonia novarae), to desert environment (e.g., Senecio lilloi), and from strict high Andean (e.g., Senecio kunturinus and S. delicatulus) to lowland species (e.g. Eupatorium tucumanense and Isostigma molfinianum).

Nearly 1/3 of the endemic Asteraceae belong to the genus Senecio with 39 endemic species in the study region--twice as many species as the second most common genera Lobivia Britton & Rose (Cactaceae), Nototriche (Malvaceae), and Solanum L. (Solanaceae) (see Table 4). Nonetheless, the remaining 2/3 of the endemic Asteraceae are highly diverse belonging to 16 of approx. 48 Asteraceae tribes (Funk et al., 2009), the most numerous being the Astereae Cass., Eupatorieae Cass., and Heliantheae Cass. including 17, 13, and 10 species each.

Although both the origin and early diversification of the Asteraceae are likely to have occurred in southern South America (Funk et al., 2009) the main radiations of Asteraceae within the study region belong to clades that are principally found in the northern hemisphere, e.g., Senecio, Hieracium L., Flourensia DC., Baccharis L., and Stevia Cav. (see Funk et al., 2009 for biogeography of the family). Still, all tribes from the basal Asteraceae grade (Mutisieae s. l. sensu Cabrera; Ortiz et. al. 2009) are represented, although in lower numbers, among the local endemic species, with Hyalideae Panero being the only exception. The Asteraceae endemics are consequently not only numerous within the region but also phylogenetically diverse, both reflecting the general family trend of being especially diverse in arid environments (Funk et al., 2009) as well as the fact that the Asteraceae is the most diverse family both in number of genera and species within the southern cone (Zuloaga et al., 1999; Moreira-Munoz & Munoz-Schick, 2007).

Cactaceae is the second most numerous family with 74 endemic species and 15 genera within the study region (Table 3). Unlike the Asteraceae the main part of the endemic Cactaceae belong to a single clade (55 species), the tribe Trichocereeae Buxb. that principally consists of high Andean species from the eastern slopes of the central Andes (Ritz et al., 2007, Hernandez-Hernandez et al., 2011). The remaining Cactaceae species are either close relatives to the Trichocereeae from the Core Cactoideae II clade that also includes the Trichocereeae (Hernandez-Hernandez et al., 2011) or from early diverging lines of the subfamily Opuntioideae (Griffith & Portery, 2009).

Like the Asteraceae, the Cactaceae are considered to be of South American origin with the early diversification of the tribes Trichocereeae and Opuntioideae situated in the Central Andes (Nyffeler, 2002; Edwards et al., 2005; Griffith & Porter 2009). While the extreme succulence of most Cactaceae species makes the family well adapted to the aridity of the southern central Andes, the Cactaceae endemics occupy nearly as wide an array of habitats as the Asteraceae including strict Altoandine species restricted to altitudes above 4000 m (Lobivia marsoneri), with endemic species only lacking from the most humid part of the study region.

Following Asteraceae, Poaceae is the second most diverse family in the southern cone (Zuloaga et al., 1999) and reaches the study region as the third most numerous family with 45 endemic species from 20 different genera (Table 3). The endemic species belong to both of the main Poaceae clades, the BEP and the PACMAD clades (GPWG 2001; Sanchez-Ken et al., 2007; GPWG II, 2012). The BEP clade generally appears in colder climates than the PACMAD clade (Edwards & Smith, 2010), and it is more species rich in the study region, with 34 endemic species versus 11 from the PACMAD.

Like Asteraceae, the endemic Poaceae species are found in most of the available habitat types (see Table 2) with two species restricted to subhumid or humid environments (Aristida pedroensis and Chusquea deficiens), high Andean species restricted to altitudes above 4000 m asl (Anatherostipa henrardiana and Poa nubensis) and lowland species only found below 1000 m asl (e.g., Digitaria catamarcensis and Neobouteloua paucirracemosa). Only strict desert endemics are lacking although a single species from semi-desert environments enter desert habitats (Pappostipa hieronymusii).

In addition to the main families discussed above, the Solanaceae and Malvaceae both include genera with considerable radiations within the region. The genus Solanum with 18 endemic species occupies a wide array of habitats from semi-arid to humid environments between 500 and 4000 m asl. Several of the Solanum endemics reach sub-humid and humid sites, with four of the species restricted to these habitats, while desert environments are occupied by another Solanaceae genus, Sclerophylax Miers with six endemic species in the region (Table 2).

As opposed to the taxa mentioned above, the radiation of Nototriche (Malvaceae) is exclusively high Andean, with all 19 endemic species found in semi-desert to semi-arid environments above 4000 m asl. Other well documented high Andean radiations such as Gentianella Moench, Lupinus L., and Valeriana L. (von Hagen & Kadereit, 2001; Bell & Donoghue, 2005; Hughes & Eastwood, 2006) are less important in the region with eleven, seven, and five endemic species, all found at lower altitudes than Nototriche.

Main areas of endemism

The NOA (Nor-Oeste-Argentina) area. The NOA area (Fig. 4B) is the general area of endemism for the southern part of the central Andes. The area is supported by species that are widely distributed within the study region and overlaps with all below mentioned sub-areas segregated from the main consensus in Fig. 4A. Four species reach the south of San Juan (Table 2) and consequently define a slightly wider pattern than the NOA area shown in Fig. 4B.

The northern limit of the NOA area coincides with the political border between Argentina and Bolivia hence the pattern probably extends into southern Bolivia. However, the scarcity of intensive botanical explorations in this part of Bolivia prevents us to speculate on the exact northern limit of the NOA pattern. So far among the defining NOA species only Mirabilis bracteosa, Mutisia kurtzii, and Trichocereus terscheckii have been collected in Bolivia, but endemic Argentinean taxa widely distributed in Jujuy are most probably also present in southern Bolivia.

Since the distribution pattern spans approximately 1000 km from north to south, the pattern is unsurprisingly not recovered when using cell sizes of 0.2[degrees]x0.2[degrees], a feature that requires collection sites every 20 to 40 km depending on the settings of the fill option (see Material and Methods). The NOA pattern emerges when using cell sizes of 0.5[degrees]x0.5[degrees]in which case 24 species are sufficiently well collected to define the area (Table 2). Other common, but less frequently collected species such as the columnar cactus Trichocereus terscheckii, support the north-south running pattern when the grid is changed to a cell size of 0.5[degrees]x1.0[degrees], allowing for even wider distances between the collecting sites. A total of 54 species support the area when using a cell size of 0.5[degrees]x1.0[degrees](Table 2).

All NOA endemics are found in desert, semi-desert and semi-arid habitats with several species also reaching either desert or sub-humid sites. Humid environments are not part of the pattern, which is consistent with the north-southern range of the area, e.g. the distributions of the defining species that reach out of the region where the humid habitats are found. Consequently, the moist requiring species are restricted to the northern portion of the study region and support sub-areas in Fig. 4A but not the general NOA pattern. The broader distribution pattern reaching San Juan is slightly dryer and does not include sub-humid habitats.

The NOA pattern includes all altitudinal layers of the vegetation as the defining species are found from 500 m to above 4500 m asl. However, slopes vegetation seem to be an important part of the distribution pattern as all species appear between 1500 m and 3500 m asl. Most species have broad ranges with some ranging nearly 3000 m asl (e.g. Oxalis famatinae see Table 2). The highest part of the NOA pattern is defined by species restricted to high Andean vegetation above 3000 m asl (e.g., Mulinum famatinense and Pycnophyllum convexum). The lowest part of the distribution pattern includes several plants that are common to Monte vegetation (Cabrera, 1976) and is delimited by Tephrocactus weberi that is found between 500-2000 m asl in desert and semi-desert habitats.

Several of the species that define the NOA pattern are common elements in the vegetation of the study region. These include the bushes Bulnesia schickendantzii, Plectocarpa rougesii (Zygophyllaceae), Mutisia kurtzii, and Cersodoma argentina (Asteraceae), as well as the grasses Panicum chloroleucum and Sporobolus maximus. Common but less frequently collected species that support the north-south running pattern when the grid is changed to cell size 0.5[degrees]x1.0[degrees], include the columnar cactus Trichocereus terscheckii, the opuntioid cactus Tephrocactus weberi, the cushion forming Bromeliaceae Deuterocohnia haumanii and Hieronymiella marginata (Amaryllidaceae) (Table 2).

High endemic areas: Jujuy, Tucuman, and Jujuy-Tucuman. Within the study region, the cells of highest endemism score include the cores of the Jujuy area, the Tucuman area, and the combined Jujuy-Tucuman area (Fig. 4C-E). These three areas are jointly supported by nearly 30% of the endemic species found in the region, and are furthermore by far the most diverse both in number of genera and families (Table 2). When also considering the species that define the diffuse border of the core areas (Fig. 4C, Table 2), about 45% of the endemic species are confined to these three distribution patterns that undoubtedly form the main area of endemism in the southern cone east of the Andes.

Common for the cores of both the Jujuy and Tucuman areas are that they include some of the study regions most variable cells both in terms of altitude, temperature, and precipitation. The available habitats within these areas range through all phytogeographic unites described for north-western Argentina (Cabrera, 1976). While variability in temperature and altitude do not decline along the north-south gradient of the study region, amount of rainfall does decline (Fig. 2A). Humid sites with an aridity index above 30 are mainly found at the eastern Andean slopes in both the Jujuy and Tucuman areas (Fig. 2C). Consequently, endemic species from humid environments, such as the subtropical Tucumano-Bolivian Yungas forests are only found in the three high endemic areas (Table 2).

Nevertheless, and although the Tucumano-Bolivian Yungas forest is one of the most diverse phytogeographic units in Argentina (Cabrera, 1976; Zuloaga et al., 1999), it should be noted that only twelve species are restricted to humid sites in the present study (Table 2), indicating that the Yungas vegetation includes few endemic species at this local scale. The main part of the endemic species are distributed through the broad array of available habitats both in terms of altitude and aridity (Table 3), with no species occupying the entire range (Table 2).Variability in precipitation rather than high rainfall might therefore be the main factor that causes high level of endemism within the Jujuy and Tucuman areas.

The decline in endemism in the center of the Jujuy-Tucuman area is consistent with the patterns found in general diversity studies (Szumik et al., 2012), and distribution maps of the Yungas forest (Hawkes & Hjerting, 1969; Cabrera, 1976; Brown, 1995). It is unclear whether the decline in diversity and endemism is caused by lack of collections or whether some habitats--e.g., humid high Andean grasslands--are lacking in this part of the area.

Minor areas of endemism: San Juan, Famatina, and south-western Bolivia. Several minor areas of endemism separate from the general area consensus in Fig. 4A. Here we discuss two southern most areas in San Juan, the central Famatina area, as well as an area related to the vegetation in south- western Bolivia. The full list of areas is found in Appendix 1.

San Juan: Two areas of endemism are defined by species restricted to the southern most part of the study region, the Andes of La Rioja-San Juan (Fig. 4F) and the San Juan area (Fig. 4G, Table 2). These two areas are well defined and do not combine with the general consensus area even under very lax consensus criteria, hence the areas do not share species with any of the other areas of endemism. This distinct distribution pattern of the endemic species from the southern part of the study region may reflect that the San Juan areas lie close to the limit of the Patagonian phytogeographic unit sensu Cabrera (1976), hence close to a general change in vegetation types that is widely recognized among biogeographers (e.g., Olson et al. 2001; Morrone, 2006). The San Juan areas are among the most arid areas within the study region. Although some of the species that define the Andes area of La Rioja-San Juan (Fig. 4F) reach sub-humid locations according to Table 2, these sub-humid locations fall at sites where the annual mean temperature is below 0[degrees]C rather than at sites with high rainfall (note that for the same annual precipitation the aridity index of de Martonne will increase with falling mean annual temperature).

Famatina. The Famatina areas both separate from the main area when raising the consensus criterion to 25% (Fig. 4H). The two partly overlapping areas include a highly endemic area of species found mainly above 1500 m asl as well as a low endemic border area of species found at lower altitudes. The areas do not share species, hence, these areas do not combine into a single consensus area. Combined both Famatina areas are supported by 28 endemic species found in desert/semi-desert and semi-arid environments between 700 m and 4000 m asl.

The area with highest endemism score contains primarily the isolated Nevados de Famatina that reaches above 6000 m asl. Mining activity at 4.600 m asl has ensured long standing accessibility to the high peaks where botanical exploration has been relatively constant since the late 19th century. The Famatina is, consequently, one of the best sampled high Andean locations south of the high- endemic areas in Jujuy and Tucuman. Nonetheless, Famatina is defined by much less endemic species than the areas Jujuy and Tucuman. Furthermore, diversity within this area is low as more than half of the endemic species belong to three of the most species rich radiations within the study region: Senecio, Nototriche at altitudes above 3000 m asl, and Gymnocalycium Pfeiff. ex Mittler in the border area (Fig. 4H) below 2000 m asl.

South-western Bolivia-northern Argentina. Towards the northern part of the study region four species delimit a shared, well defined Bolivian-Argentinean area that do not combine with any other distribution pattern even under very lax consensus criteria (Fig. 4I, Table 2-3). The area overlaps partly with the NOA pattern (Fig. 4B) but stretches further north into Bolivia. The Bolivian portion of this area corresponds to the Bolivian Prepuna as defined by Lopez (2000) while the Argentinean part only includes the northern most part of the Prepuna province sensu Cabrera (1976). The area defined in Fig. 4I includes semi-desert and semi-arid habitats between 200-4000 m. When increasing the cell size to 0.5[degrees]x1.0[degrees]two additional species (Deuterocohnia strobilifera and Oxalis cotagaitensis) are incorporated into the area.

Although only six species support the Bolivian-Argentinean area, our study did not sample exhaustively this region, therefore it is most likely that other species share a similar distribution pattern. According to the Catalogue of the Plants of the Southern Cone (Zuloaga et al., 2008) at least 70 species are distributed from Salta to Bolivia many of which may approximately fit the area in Fig. 4I. This distribution pattern stresses the similarity between the flora of southern Bolivia and north- western Argentina as mentioned by Lopez (2000, 2003), but further sampling within the region is needed to establish the size and importance of the area.

The Bolivian Prepuna, sensu Lopez (2000), is nested within the above south-western Bolivia-northern Argentina distribution pattern but combined with the main distribution pattern shown in Fig. 4A. The Bolivian Prepuna segregates from the main area in Fig. 4A when the consensus criterion is raised to 25% or higher (Fig. 4J). The Bolivian Prepuna is formed by two overlapping distribution patterns one of which enters northern Jujuy (Argentina). Both patterns are supported by species that are mainly found in semi-arid environments from 2500 m up to 4000 m asl with the highest collection sites lying in the Argentinean side of the pattern. Since we did not compile an exhaustive list of species endemic to southern Bolivia, the Bolivian Prepuna may also be supported by more species that those included in our study.

Phytogeographic divisions: the Prepuna province and areas of endemism

Although Cabrera aimed to base his classic phytogeographic scheme on the presence of endemic taxa from family to species level (Cabrera 1951, 1953, 1976), the system was not based on quantitative studies and did not apply consistent criteria for defining the individual phytogeographic units (Ribichich, 2002). It is therefore not surprising that quantitative analyses, based on endemic species distribution, result in areas of endemism that do not correspond to Cabrera's phytogeographic divisions. In our analyses all areas were defined by species from a wide array of altitudes in most cases including lowland, slopes, and high Andean species (Table 2). Moreover, as the areas with highest endemism were found in regions where several climatic regimes meet (Jujuy and Tucuman), they include several phytogeographic units within shortest distances of each other. As a result, the species that define the high endemic areas are just as variable as the areas themselves in their altitude and aridity ranges (Table 2).

Cabrera (1976) defined the Prepuna province as xeric slope vegetation with emphasis on the distribution of emblematic species such as columnar cacti and cushion forming Bromeliaceae species, mentioning several species from different families as common on the slopes. Lopez (2000, 2003) later extended the Prepuna to include a similar xeric vegetation in the south-western Bolivia. The Prepuna, which is here included in its entire extension, does not appear among the resulting areas of endemism, hence the Prepuna province sensu Cabrera (1976) is not definable simply by the presence of two or more unique Prepuna species.

The partly overlapping south-western Bolivia/northern NOA and the NOA area (Figs. 7 and 12) cover geographically the Prepuna in its entire extension. Both areas are defined by several of Cabrera's (1976) Prepuna species including columnar cacti and cushion forming Deuterocohnia Mez species. Notably three of the species mentioned by Cabrera: Aphylloclados spartioides, Cercidium andicola, and Prosopis ferox define the joint Southern-Bolivia/northern NOA (Fig. 4I) supporting the northern Prepuna as delimited by Lopez (2000, 2003) in which the Bolivian Prepuna separates as an independent area of endemism (Fig. 4J). Columnar Trichocereus (A. Berger) Riccob. species define both the Bolivian Prepuna and the NOA area while most of Cabrera's (1976) remaining Prepuna species support the NOA area. The NOA distribution pattern does, however, also include species from other altitude strata, hence, although it appears to include the Prepuna it defines a broader area of endemism in the southern central Andes.

CONCLUSIONS

Located in the center of the South American dry diagonal, the aridity of southern central Andes is reflected in the distribution of its endemic vascular flora. Of 540 endemic plant species, more than 2/3 are restricted to semi-desert and semi-arid habitats including some of these regions main radiations, such as the ultra high Andean Nototriche (Malvaceae) and the radiation of Gymnocalycium (Cactaceae) found below 2000 m. Even the 39 endemic species from the cosmopolitan genus Senecio (Asteraceae) are almost exclusively found in semi-desert or semi-arid habitats with only two species entering subhumid or humid locations.

The distributional bias of the endemic species towards arid sites is in contrast with that of vascular plant diversity in the region, since the Tucumano-Bolivian Yungas forests, on the humid eastern Andes slopes, undoubtedly include the most diverse vegetation of the region (Brown, 1995; Ibisch et al., 2003). The Tucumano-Bolivian Yungas forests in north-western Argentina are, however, a patch of a more extensive area of endemism whose northern limit is found south of the Andes bend at approx. 18[degrees]S (Ibisch et al., 2003). Emblematic Yungas species, such as Juglans australis Griseb., Podocarpus parlatorei Pilg, and Duranta serratifolia (Griseb.) Kuntze appear to be endemic to the Tucomano-Bolivian Yungas. Other species such as Alnus acuminata Kunth and Fuchsia boliviana Carriere are widely distributed along the eastern slopes and further north to Mexico. These forests may consist of several nested or partly overlapping distribution patterns as is the case of the endemic vegetation of the arid slopes of the southern central Andes. Any attempt to evaluate endemism of the Tucomano-Bolivian Yungas should include these forests in their entire extension.

Although the Yungas forests are of little importance for endemisms at the scale of our study, the high endemic areas of the region lies in juxtaposition, west of the main rainfall zones (Fig 1a). The endemic species of these areas are found in a broad and variable range of aridity and altitude (Table 2), hence, rather than high rainfall itself, the gradual decreasing rain-veil west of the main rainfall zone caused by the complex topography of the region, may be the main factor allowing for the elevated number of endemic species to coexist in these areas.

We did not find any relationship between the main areas of endemism and phytogeographic units previously defined, as the main areas of endemism appeared in cells that included a widest array of habitats. The Prepuna province sensu Cabrera (1976) and Lopez (2000, 2003) was not defined by the endemic species as a unit; instead, two partly overlapping areas covered the entire extension of the Prepuna (Figs. 6 and 14). Both areas include several Prepuna characteristics sensu Cabrera (1976). Nevertheless, of these two areas, the NOA area should be considered a general area of endemism for the southern central Andes since it also included high Andean and lowland species.

ACKNOWLEDGEMENTS

We thank Claudia Szumik for helpful discussions and support. We also thank the staff at The Lillo Foundation (Tucuman) as well as Dr. Beck and the staff at the National Herbaria of Bolivia for help and hosting us during our visits to these Institutes. We also thank the staff at IBODA for the identification of all newly collected material, and Fernando Biganzoli for accompanying us on several collecting trips. We gratefully acknowledge GBIF and John Wieczorek for the georeferencing workshop in Buenos Aires. CONICET (PIP-CONICET 0207) and National Geographic (Grant N[degrees]8862-10) funded the project and field collections respectively.

APPENDIX 1

NOA (Fig. 4B)

The NOA pattern is described in detail in the discussion. Here we only add that the area is composed by three different main patterns. In its widest version the pattern runs from the limit between Argentina and Bolivia to San Juan supported by four species (see Table 2) the most notably being Adesmia cytisoides and Jarava media. Several species are distributed as shown in the consensus area in Fig. 4B e.g., Panicum chloroleucum and Polygala argentiniensis, while the third group of species are lacking in the northernmost part of the area, e.g., Bulnesia schickendantzii, Plectrocarpa rougesii and Sporobolus maximus.

Yavi-Santa Victoria (Fig. 4K)

The Santa Victoria-Yavi area separates as an individual area of endemism in the north-eastern corner of the Jujuy border. The Santa Victoria-Yavi area is supported by 17 endemic species with distributions tightly adjusted to the area, that separates from the general area of endemism at 15% aa, hence the high endemism scores is due to little species overlap between this area and other distribution patterns rather than a high number of endemic species. Nearly all defining species have been collected in the surroundings of Yavi in semi-arid Puna or along the high Andean road between Yavi and Santa Victoria in sub-humid high Andean, or montane grassland. We suspect that the Santa Victoria-Yavi area is artificially delimited both towards the north and south. Botanical collections are incomplete in southern Bolivia but the eastern slopes of the Eastern Andes range in Salta area almost unexplored botanically except for the road leading to Santa Victoria.

Jujuy (Fig. 4C)

The core of the area that lies within the Eastern Andes range is supported by 67 endemic species from 48 genera and 23 families (Table 2). The endemic species are found in a wide range of available habitats in semi-desert to humid environments between 500->4500 m asl that includes the tropical moist Yungas forest (Sierra Calilegua), mountain grasslands (Sierra de Calilegua, Sierra de Zenta), as well as the slopes of the inner dry valleys (Quebrada de Humahuaca). The few species that are found exclusively above 3500 m, in this area, are from well to sparsely collected high peaks such as Mina Aguilar and Nevado de Chani, as well as Nevado del Castillo and Cerro de Fundicion in Salta.

The Jujuy area is not easily delimited as the core is surrounded by a diffuse border with a gradually decreasing endemism score (Fig. 4C). The high number of sub-sets included in the consensus also indicates that the distribution patterns supporting this area are far from uniform (Table 3). A total of 35 species are endemic to the border (Table 2) that includes same habitats as the core as well as Puna environment between the Eastern and Western Andes range. The main part of the species defining the border area are either species found in the Puna (e.g., Lobivia einsteinii, Mancoa venturii, and Senecio punae) or species present along the the mountain grasslands reaching the Santa Victoria area (e.g., Macropharynx meyeri, Microliabum humile, and Silene bersieri). Part of the area enters Bolivia as four of the defining species have been found in the Tarija department (Parodia stuemeri, Psychotria argentinensis, Puya micrantha, and Solanum calileguae). More of the border species are likely to appear in the southern Bolivia as botanical inventories become more complete.

Salta (Fig. 4L)

Between the high-endemic Jujuy and Tucuman areas lies the Salta area that separates from the main consensus under the 50% consensus criterion. A total of 29 species are endemic to the core area mainly found in montane grasslands, and slopes of the inner dry Valleys.

Like the Jujuy area, the Salta area has diffuse borders that overlap and shares species with the border of the Jujuy area and the northern part of the Tucuman area. The high Andean endemics are restricted to this part of the area where endemic species from Nevado del Castillo and Cerro del Cajon also supports the southern Jujuy or northern Tucuman area respectively. The high mountains peaks of the core area such as Cerro Malcante (5226 m) and Nevados de Palermo (6200m) have not been explored botanically to our knowledge and could add new endemics to this area.

Tucuman (Fig. 4D)

Like the Jujuy area the Tucuman area lies in the eastern Andes range and contains mainly the same habitat types as in the core-Jujuy area. In terms of altitude, temperature, and precipitation, the Tucuman area is just as variable as the core area of Jujuy. A total of 62 species are endemic to the area from a total of 41 genera and 25 families (Table 2).

The area includes tropical moist Yungas forest (eastern slopes of Sierra del Aconquija and Cumbres de Tafi), mountain grasslands (Cumbres del Tafi, Cumbres Calchaquies and Sierra del Aconquija), as well as the slopes of the inner dry Valleys (Valles Calchaquies). Species found exclusively above 3500 m asl are mainly from peaks of the two main ranges (Sierra Aconquija and Cumbres Calchaquies) or from Cerro del Cajon in the northern extreme of Sierra de los Quilmes.

Unlike the Jujuy area, the Tucuman area has well defined limits without diffuse border mainly because species distribution towards the west can be defined as an independent area of endemism, see the Ambato area below.

Jujuy-Tucuman (Fig. 4E)

The combined Jujuy-Tucuman area is defined by species that are found in the cores and/or borders of both the Jujuy and Tucuman areas. The Jujuy-Tucuman area contains the most complex distribution patterns of the study region, being the left over of the main consensus area when all sub-areas are extracted at a consensus criterion of 50%. Like the Jujuy area, the high amount of individual sub-sets that are included in this area indicates that distribution patterns supporting this area are not uniform and variations among these are the main source of the high number of subsets found in the analysis in general.

Ambato (Fig. 4M)

West of the Tucuman area and partly overlapping with this lies the Ambato area (Fig. 4M) that is defined by species that reach further south and/or east than the species defining the Tucuman area.

The high endemic core of the area incudes 16 endemic species (35% aa) found in semi-desert to semi-arid environments from 500-3500 m. These species are mainly found in the montane grasslands along the eastern slopes of Sierra Ambato and in high Andean habitats of Cerro Manchado.

The border of the Ambato area extends from the core towards the north and west partly overlapping with the Belen- Tinogasta area (Fig. 4N).

Belen-Tinogasta (Fig. 4N)

The Belen-Tinogasta area (Fig. 4N) is the western most and least species rich of the three partly overlapping areas of Tucuman, Ambato, and Belen-Tinogasta. The area is supported by 12 species from rare collections in arid or hyper-arid regions of Sierras de Belen or northern Tinogasta. Only a single species [Tephrocactus geometricus (A. Cast.) Backeb.] have been collected repeatedly within the area in Sierras de Belen and Sierra de Zapata. The area furthermore includes rare collection of endemic species from the valleys southeast of Cerro Chucula (Tinogasta) and from the eastern slopes of Sierra de Fiambala northwest of Belen.

Famatina (Fig. 4H)

Endemism in the Famatina area are mainly from the isolated Nevados de Famatina but some species are also found in the surrounding valleys of Chilcito and Vinchina as well as Cuesta de Miranda. The core of the area is supported by 20 endemic species mainly found in semi-desert to semi-arid environments from 1000-4000 m. Twelve of the defining species are endemic to Nevados de Famatina with nine species restricted to altitudes at approx. 4000 m asl including five species of the ultra high Andean Nototriche genera. The border of the Famatina areas is composed of a single area mainly defined by four Gymnocalycium species from low altitudes in valleys north-east of the Nevados de Famatina.

Andes La Rioja-San Juan (Fig. 4F)

Includes high Andes in La Rioja and San Juan and is mainly defined by species found above 3000 m in the regions of Laguna Brava (La Rioja) and Parque Nacional San Guillermo (San Juan) or collected along the few high Andean roads south of San Guillermo. Few species from the Andean foothill, e.g., Aphyllocladus ephedroides and Solanum glaberrimum also support the area.

San Juan (Fig. 4G)

The southern San Juan area is found outside the Andes range. The area includes valleys and minor peaks of the Pampeanas Ranges with all defining species found below 3000 m. Highest altitudes of the defining species are found in Sierra Pie de Palo and Sierra del Tontal. The southern San Juan area is one of the driest areas with 50% of the specimens collected in desert environments.

Inner valleys and Pampeanas Range

The four partly overlapping areas below are defined by species mainly found in the inner valleys, slopes and summits of the Sierras Pampeanas. Three of the four areas include species that reach above 4000 m asl with the lower part of the area set by the altitude of the valley button in the respective areas. Species from the areas are restricted to semi- desert and semi-arid habitats while few enter desert environments. Sub-humid locations are only reached in these areas at high altitudes with low annual mean temperature rather than higher rainfall.

Salta-Catamarca summits and dry inner valleys (Fig. 4O). The northernmost of the Pampeanas Range areas are defined by species distributed along valleys and high Andean locations from Santa Rosa de Tastil (Quebrada del Toro, Salta) to Sierra Ambato and Quebrada de Belen (Catamarca).

Catamarca-La Rioja summits and dry inner valleys (Fig. 4P). This area includes mainly the locations Hualfin, Andalgala, Londres, Cuesta de Zapata, and Mina Capillitas in Catamarca as well as Los Corrales in Famatina, La Rioja. Several species are found in the semi-desert of the inner valleys, e.g., the five Cactaceae species that defines the area as well as Sclerophylax cynocrambe from the arid radiation of Solanaceae. The high Andean part of the area is defined by species distributed along the summits of the Calchaquies, Ambato and Famatina e.g., Poa plicata and Viola triflabellata.

La Rioja-San Juan summits and dry inner valleys (Fig. 4Q--obtained under cell size 0.5[degrees]x1.0[degrees]). The area is mainly defined by species distributed in valleys from southern Catamarca to San Juan. Several of the endemic species from a wide array of families reach desert environments in this area, e.g., Flourensia hirta and Senecio sanagastae (Asteraceae), Lobivia famatimensis and Tephrocactus alexanderi (Cactaceae), Senna fabrisii (Fabaceae), Guindilia cristata (Sapindaceae), and Sclerophylax kurtzii (Solanaceae). Only few species in this area reach high Andean environments, e.g., Zephyranthes diluta (Amaryllidaceae) and Adesmia nanolignea (Fabaceae) that are found along the high Andes of La Rioja and San Juan.

Valle Fertil (Fig. 4R--obtained under cell size 0.5[degrees]x1.0[degrees]). This minor area of endemism only appears when using longitudinal cells of 0.5[degrees]x1.0[degrees]in which case four species from different families define the area (Table 2). The main part of the area consists of the valley between Sierra de Valle Fertil/Sierra de La Huerta in San Juan and Sierra de Los Llanos/Sierra de las Minas in La Rioja. This area also reaches the northern part of San Luis, where nearly all defining species have been confirmed.

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Lone Aagesen [1], Maria J. Bena [1], Soledad Nomdedeu [1], Adela Panizza [1], Ramiro P. Lopez [2, 3] & Fernando O. Zuloaga [1]

[1] Instituto de Botanica Darwinion (ANCEFN-CONICET), Labarden 200, Casilla de Correo 22, B1642HYD San Isidro, Buenos Aires, Argentina; laagesen@darwin.edu.ar (author for correspondence).

[2] Herbario Nacional de Bolivia, Campus Universitario, Cotacota, calle 27 s/n, Casilla 3-35121 La Paz, Boliva.

[3] Laboratorio de Ecofisiologia-IEB, Departamento de Biologia, Universidad de La Serena, Casilla de correo 599, La Serena, Chile.
Table 1. General results for all analyses under different grid sizes,
consensus criterion (%), and consensus rules (ae and aa). For more
information on consensus rules and criterion see Materials and
Methods.

             0.2[degrees] x 0.2[degrees]   0.5[degrees] x 0.5[degrees]
               25%-50%        50%-75%        10%-20%        25%-50%
Sub-sets          72            125            783            695
(number of       -144           -192           -365           -425
  defining
  species)
ae 50%            48             64            305            264
ae 33%            39             52            211            184
ae 10%            29             40            104            106
ae 5%             28             37             87             82
aa 50%            40             48             80             65
aa 33%            25             26             29             22
aa 10%            15             17             9              7
aa 5%             14             17             9              7

             0.5[degrees] x 1.0[degrees]
             10-20%-5-10%     25%-50%
Sub-sets         958            826
(number of       -425           -505
  defining
  species)
ae 50%           355            286
ae 33%           248            203
ae 10%           123             92
ae 5%             92             70
aa 50%            81             64
aa 33%            7              8
aa 10%            2              2
aa 5%             2              2

Table 2a. List of endemic species. Each species is assigned to the
area of endemism in which it obtained the best score. Altitudinal
ranges were obtained from the georeferenced locations and therefore
are suggestive--not observed field data. For area names see Table 2b.

Familiy             Species (number of
                    georeferenced records)

Alstroemeriaceae    Alstroemeria bakeri Pax (5)
Amaryllidaceae      Habranthus pictus Ravenna (1)
Apocynaceae         Philibertia coalita (Lillo) Goyder (3)
Asteraceae          Hieracium lorentzianum Zahn (8)
Asteraceae          Hieracium sordidum Griseb. (8)
Asteraceae          Hieracium vervoorstii Sleumer (1)
Asteraceae          Luciliocline catamarcense (Cabrera) Anderb. &
                      S.E. Freire (1)
Asteraceae          Senecio ambatensis Cabrera (1)
Bromeliaceae        Puya harmsii (A. Cast.) A. Cast. (5)
Cactaceae           Acanthocalycium glaucum F. Ritter (3)
Cactaceae           Gymnocalycium hybopleurum (K. Schum.) Backeb. (7)
Cactaceae           Lobivia crassicaulis Backeb. ex R. Kiesling (1)
Calyceraceae        Boopis castillonii (Hicken) Pontiroli (1)
Convolvulaceae      Cuscuta argentinana Yunck. (6)
Dioscoreaceae       Dioscorea trifurcata Hauman (1)
Fabaceae            Lupinus burkartianus C.P. Sm. (2)
Fabaceae            Senna pachyrrhiza (L. Bravo) H.S. Irwin &
                      Barneby (1)
Iridaceae           Sisyrinchium biflorum Griseb. (2)
Poaceae             Agrostis ambatoensis Asteg. (1)
Poaceae             Digitaria catamarcensis Rugolo (2)
Poaceae             Nassella catamarcensis Torres (1)
Poaceae             Nassella ragonesei Torres (4)
Solanaceae          Solanum crebrum C.V. Morton ex L.B. Sm. (1)
Solanaceae          Solanum mortonii Hunz. (4)
Verbenaceae         Verbena andalgalensis Moldenke (1)
Violaceae           Viola joergensenii W. Becker (1)
Asteraceae          Aphyllocladus ephedroides Cabrera (6)
Asteraceae          Huarpea andina Cabrera (3)
Asteraceae          Senecio cremnophilus I.M. Johnst. (3)
Boraginaceae        Cryptantha latefissa R.L. Perez-Mor. (1)
Brassicaceae        Sarcodraba andina O.E. Schulz (2)
Fabaceae            Adesmia sanjuanensis Burkart (2)
Fabaceae            Astragalus boelckei Gomez-Sosa (4)
Fabaceae            Astragalus nelidae Gomez-Sosa (6)
Fabaceae            Astragalus pulviniformis I.M. Johnst. (1)
Juncaceae           Oxychloe castellanosii Barros (7)
Malvaceae           Nototriche copon Krapov. (4)
Poaceae             Nassella famatinensis Torres (4)
Solanaceae          Solanum glaberrimum C.V. Morton (4)
Valerianaceae       Valeriana corynodes Borsini (3)
Violaceae           Viola flos-evae Hieron. (2)
Asteraceae          Aphyllocladus spartioides Wedd. (22)
Asteraceae          Flourensia fiebrigii S.F. Blake (15)
Bromeliaceae        Deuterocohnia strobilifera Mez (5)
Fabaceae            Cercidium andicola Griseb. (20)
Fabaceae            Prosopis ferox Griseb.(30)
Oxalidaceae         Oxalis cotagaitensis Griseb. (6)
Amaryllidaceae      Chlidanthus marginatus (R.E. Fr.) Ravenna (2)
Apocynaceae         Philibertia micrantha (Malme) Goyder (2)
Asteraceae          Chuquiraga acanthophylla Wedd. (8)
Asteraceae          Dasyphyllum hystrix (Wedd.) Cabrera (2)
Asteraceae          Gochnatia cardenasii S.F. Blake (15)
Asteraceae          Senecio jujuyensis Cabrera (7)
Bromeliaceae        Tillandsia camargoensis L. Hrom. (2)
Cactaceae           Cleistocactus tupizensis (Vaupel) Backeb. & F.M.
                      Knuth (3)
Cactaceae           Opuntia ferocior (Backeb.) G.D. Rowley (3)
Cactaceae           Oreocereus celsianus (Lem. ex Salm-Dyck)
                      Riccob. (7)
Cactaceae           Oreocereus trollii (Kupper) Backeb. (9)
Cactaceae           Parodia maassii (Heese) A. Berger (13)
Cactaceae           Tephrocactus chichensis Cardenas (3)
Cactaceae           Trichocereus tacaquirensis (Vaupel) Cardenas ex
                      Backeb. (5)
Cactaceae           Trichocereus werdermannianus Backeb. (7)
Fabaceae            Acacia feddeana Harms (14)
Iridaceae           Mastigostyla cabrerae R.C. Foster (3)
Ledocarpaceae       Balbisia integrifolia R. Knuth (3)
Malvaceae           Abutilon fuscicalyx Ulbr. (8)
Solanaceae          Iochroma cardenasianum Hunz. (8)
Zygophyllaceae      Bulnesia rivas-martinezii G. Navarro (6)
Acanthaceae         Justicia riojana Lindau (6)
Amaryllidaceae      Habranthus andalgalensis Ravenna (8)
Asteraceae          Chiliotrichiopsis ledifolia (Griseb.) Cabrera (3)
Asteraceae          Flourensia blakeana M.O. Dillon (4)
Asteraceae          Flourensia tortuosa Griseb. (12)
Asteraceae          Hysterionica pulchella Cabrera (6)
Asteraceae          Senecio toroanus Cabrera (3)
Bromeliaceae        Tillandsia tenebra L. Hrom. & W. Till (5)
Cactaceae           Gymnocalycium baldianum (Speg.) Speg. (6)
Cactaceae           Gymnocalycium pugionacanthum Backeb. ex H.
                      Till (7)
Cactaceae           Trichocereus andalgalensis (F.A.C. Weber)
                      Hosseus (10)
Cactaceae           Trichocereus cabrerae R. Kiesling (4)
Cactaceae           Trichocereus huascha (F.A.C. Weber) Britton &
                      Rose (7)
Fabaceae            Adesmia pseudoincana Burkart (2)
Fabaceae            Lupinus alivillosus C.P. Sm. (3)
Fabaceae            Lupinus tucumanensis C.P. Sm. (4)
Gentianaceae        Gentianella claytonioides (Gilg) T.N. Ho & S.W.
                      Liu (5)
Gentianaceae        Gentianella kurtzii (Gilg) Fabris (5)
Poaceae             Poa plicata Hack. (9)
Ranunculaceae       Ranunculus lancipetalus Griseb. (8)
Rosaceae            Lachemilla asplenifolia (Rothm.) Rothm. (2)
Solanaceae          Sclerophylax cynocrambe (Griseb.) Griseb. (10)
Verbenaceae         Aloysia catamarcensis Moldenke (11)
Violaceae           Viola triflabellata W. Becker (9)
Amaryllidaceae      Habranthus riojanus Ravenna (2)
Asteraceae          Aphyllocladus decussatus Hieron. (1)
Asteraceae          Baccharis famatinensis Ariza (1)
Asteraceae          Flourensia niederleinii S.F. Blake (5)
Asteraceae          Senecio cremnicola Cabrera (1)
Asteraceae          Senecio famatinensis Cabrera (1)
Asteraceae          Senecio krapovickasii Cabrera (1)
Asteraceae          Senecio lanosissimus Cabrera (3)
Asteraceae          Senecio niederleinii Cabrera (2)
Brassicaceae        Menonvillea famatinensis (Boelcke) Rollins (1)
Cactaceae           Gymnocalycium ritterianum Rausch (2)
Cactaceae           Trichocereus pseudocandicans Backeb. ex R.
                      Kiesling (3)
Cactaceae           Trichocereus vatteri R. Kiesling (2)
Fabaceae            Lupinus hieronymi C.P. Sm. (1)
Gentianaceae        Gentianella riojae (Gilg) Fabris ex J.S.
                      Pringle (2)
Malvaceae           Nototriche famatinensis A.W. Hill (1)
Malvaceae           Nototriche glabra Krapov. (1)
Malvaceae           Nototriche kurtzii Krapov. (1)
Malvaceae           Nototriche niederleinii A.W. Hill (4)
Malvaceae           Nototriche pulvilla A.W. Hill (2)
Verbenaceae         Acantholippia riojana Hieron. ex Moldenke (3)
Asteraceae          Isostigma molfinianum Sherff (2)
Cactaceae           Gymnocalycium albiareolatum Rausch (6)
Cactaceae           Gymnocalycium kieslingii O. Ferrari (3)
Cactaceae           Gymnocalycium mazanense (Backeb.) Backeb. (5)
Cactaceae           Gymnocalycium mucidum Oehme (3)
Cactaceae           Gymnocalycium uebelmannianum Rausch (2)
Anacampserotaceae   Anacampseros vulcanensis Anon (3)
Apocynaceae         Philibertia castillonii (Lillo ex T. Mey.)
                      Goyder (2)
Asteraceae          Eupatorium arachnoideum Legname (2)
Asteraceae          Eupatorium hickenii Cabrera & Vittet (2)
Asteraceae          Hieracium kieslingii Cabrera (1)
Asteraceae          Hieracium luteomontanum Cabrera (1)
Asteraceae          Laennecia altoandina (Cabrera) G.L. Nesom (2)
Asteraceae          Perezia volcanensis Cabrera (1)
Asteraceae          Senecio altoandinus Cabrera (1)
Asteraceae          Senecio keshua Cabrera (2)
Asteraceae          Senecio tilcarensis Cabrera (3)
Asteraceae          Senecio yalae Cabrera (2)
Asteraceae          Stevia crassicephala Cabrera (2)
Asteraceae          Stevia yalae Cabrera (2)
Bromeliaceae        Puya assurgens L.B. Sm. (2)
Cactaceae           Lobivia densispina (Werderm.) Buining (2)
Cactaceae           Lobivia gonjianii (R. Kiesling) R. Kiesling (2)
Cactaceae           Lobivia jajoiana Backeb. (1)
Cactaceae           Lobivia marsoneri (Werderm.) Backeb. (1)
Cactaceae           Parodia chrysacanthion (K. Schum.) Backeb. (3)
Cactaceae           Pyrrhocactus umadeave (Werderm.) Backeb. (3)
Cactaceae           Rebutia marsoneri Werderm. (2)
Cactaceae           Trichocereus fabrisii R. Kiesling (1)
Cactaceae           Tunilla tilcarensis (Backeb.) D.R. Hunt & J.
                      Iliff (1)
Caryophyllaceae     Pycnophyllum mucronulatum Mattf. (1)
Convolvulaceae      Cuscuta friesii Yunck. (2)
Crassulaceae        Sedum jujuyense Zardini (3)
Dioscoreaceae       Dioscorea castilloniana Hauman ex Castillon (3)
Fabaceae            Adesmia arenicola (R.E. Fr.) Burkart (1)
Malvaceae           Nototriche macrotuba Krapov. (1)
Malvaceae           Tarasa latearistata Krapov. (1)
Orchidaceae         Chloraea cogniauxii Hauman (3)
Plantaginaceae      Plantago jujuyensis Rahn (3)
Urticaceae          Pilea jujuyensis Soraru (2)
Velloziaceae        Barbaceniopsis humahuaquensis Noher (4)
Verbenaceae         Junellia ballsii (Moldenke) N. O'Leary & P.
                      Peralta (3)
Acanthaceae         Aphelandra lilacina C. Ezcurra (1)
Apiaceae            Mulinum axilliflorum Griseb. (9)
Apocynaceae         Macropharynx meyeri (C. Ezcurra) Xifreda (4)
Apocynaceae         Matelea schreiteri (T. Mey.) Pontiroli (7)
Apocynaceae         Philibertia affinis (Griseb.) Goyder (1)
Aristolochiaceae    Aristolochia oranensis Ahumada (2)
Asteraceae          Gamochaeta longipedicellata Cabrera (2)
Asteraceae          Mutisia saltensis Cabrera (2)
Asteraceae          Senecio tocomarensis Cabrera (1)
Asteraceae          Stevia centinelae Cabrera (1)
Asteraceae          Vernonia lipeoensis Cabrera (5)
Asteraceae          Vernonia novarae Cabrera (1)
Brassicaceae        Exhalimolobos burkartii (Romanczuk & Boelcke)
                      Al-Shehbaz & C.D. Bailey (2)
Brassicaceae        Lepidium jujuyanum Al-Shehbaz (1)
Brassicaceae        Petroravenia eseptata Al-Shehbaz (1)
Bromeliaceae        Pitcairnia saltensis L.B. Sm. (1)
Bromeliaceae        Puya micrantha Mez (4)
Cactaceae           Parodia nivosa Backeb. (1)
Celastraceae        Maytenus cuezzoi Legname (4)
Cucurbitaceae       Sicyos ignarus Mart. Crov. (1)
Euphorbiaceae       Acalypha friesii Pax & K. Hoffm. (1)
Fabaceae            Astragalus fabrisii Gomez-Sosa (2)
Fabaceae            Astragalus punae I.M. Johnst. (1)
Gentianaceae        Gentianella multiflora (Griseb.) Fabris (2)
Iridaceae           Cypella elegans Speg. (2)
Malvaceae           Nototriche cabrerae Krapov. (2)
Piperaceae          Peperomia aldrinii Villa (1)
Poaceae             Anatherostipa henrardiana (Parodi) Penailillo (2)
Poaceae             Aristida pedroensis Henrard (1)
Poaceae             Chusquea deficiens Parodi (1)
Poaceae             Jarava breviseta (Caro & E.A. Sanchez)
                      Penailillo (2)
Poaceae             Nassella novari Torres (11)
Rubiaceae           Psychotria argentinensis Bacigalupo (5)
Solanaceae          Solanum fabrisii Cabrera (1)
Solanaceae          Solanum zuloagae Cabrera (2)
Tropaeolaceae       Tropaeolum willinkii Sparre (2)
Amaryllidaceae      Zephyranthes andina (R.E. Fr.) Traub (4)
Apocynaceae         Metastelma microgynostegia Pontiroli (5)
Asteraceae          Flourensia suffrutescens (R.E. Fr.) S.F. Blake (8)
Asteraceae          Microliabum humile (Cabrera) Cabrera (3)
Asteraceae          Mikania jujuyensis Cabrera (3)
Asteraceae          Senecio infimus Cabrera (6)
Asteraceae          Senecio punae Cabrera (12)
Asteraceae          Stevia jujuyensis Cabrera (4)
Asteraceae          Trichocline macrorhiza Cabrera (6)
Begoniaceae         Begonia sleumeri L.B. Sm. & B.G. Schub. (3)
Brassicaceae        Dictyophragmus punensis (Romanczuk) Al-Shehbaz (5)
Brassicaceae        Mancoa venturii Al-Shehbaz (5)
Cactaceae           Lobivia einsteinii (Fric) Rausch (10)
Cactaceae           Lobivia nigricans Wessner (5)
Cactaceae           Maihueniopsis minuta (Backeb.) R. Kiesling (4)
Cactaceae           Parodia stuemeri (Werderm.) Backeb. (8)
Caryophyllaceae     Silene bersieri Bocquet (3)
Caryophyllaceae     Silene haumanii Bocquet (4)
Convolvulaceae      Ipomoea volcanensis O'Donell (9)
Fabaceae            Adesmia friesii Burkart ex Ulibarri (7)
Fabaceae            Lupinus jujuyensis C.P. Sm. (2)
Malvaceae           Nototriche castillonii B.L. Burtt & A.W. Hill (3)
Malvaceae           Nototriche friesii A.W. Hill (3)
Orobanchaceae       Bartsia jujuyensis Cabrera & Botta (9)
Poaceae             Anatherostipa brevis (Torres) Penailillo (4)
Poaceae             Aristida pubescens Caro & E.A. Sanchez (5)
Poaceae             Eragrostis andicola R.E. Fr. (21)
Solanaceae          Solanum calileguae Cabrera (5)
Valerianaceae       Valeriana altoandina Cabrera (5)
Amaryllidaceae      Hippeastrum aglaiae (A. Cast.) Hunz. & Cocucci (13)
Anthericaceae       Anthericum hickenianum Poelln. (8)
Apiaceae            Bowlesia hieronymusii H. Wolff (3)
Apocynaceae         Oxypetalum tucumanense (T. Mey.) Goyder &
                      Rapini (3)
Apocynaceae         Philibertia barbata (Malme) Goyder (14)
Apocynaceae         Philibertia cionophora (Griseb.) Goyder (6)
Asteraceae          Flourensia macroligulata Seeligm. (3)
Asteraceae          Senecio catamarcensis Cabrera (4)
Bromeliaceae        Puya lilloi A. Cast. (13)
Bromeliaceae        Puya smithii A. Cast. (5)
Bromeliaceae        Tillandsia friesii Mez (4)
Cactaceae           Rebutia minuscula K. Schum. (15)
Cactaceae           Trichocereus thelegonus (F.A.C. Weber) Britton &
                      Rose (3)
Euphorbiaceae       Acalypha schreiteri Lillo ex Lourteig &
                      O[acute accent]Donell (4)
Fabaceae            Astragalus burkartii I.M. Johnst. (7)
Gentianaceae        Gentianella cosmantha (Griseb.) J.S. Pringle (4)
Gentianaceae        Gentianella tubulosa (Gilg) Fabris (7)
Iridaceae           Cardenanthus venturi R.C. Foster (2)
Iridaceae           Ennealophus simplex (Ravenna) Roitman & A.
                      Castillo (3)
Loasaceae           Caiophora nivalis Lillo (9)
Malvaceae           Tarasa meyeri Krapov. (8)
Orchidaceae         Chloraea subpandurata Hauman (7)
Orchidaceae         Sacoila secundiflora (Lillo & Hauman) Garay (4)
Solanaceae          Solanum collectaneum C.V. Morton (10)
Solanaceae          Solanum delitescens C.V. Morton (18)
Solanaceae          Solanum venturii Hawkes & Hjert. (7)
Tropaeolaceae       Tropaeolum argentinum Buchenau (9)
Valerianaceae       Valeriana tucumana Borsini (6)
Acanthaceae         Dicliptera cabrerae C. Ezcurra (8)
Alstroemeriaceae    Bomarea macrocephala Pax (13)
Alliaceae           Schickendantziella trichosepala (Speg.) Speg. (4)
Amaranthaceae       Gomphrena radiata Pedersen (8)
Amaryllidaceae      Hieronymiella speciosa (R.E. Fr.) Hunz. (11)
Anacardiaceae       Schinus gracilipes I.M. Johnst. (11)
Anthericaceae       Anthericum argentinense (Hauman) Guagl. (9)
Apiaceae            Austropeucedanum oreopansil (Griseb.) Mathias &
                      Constance (6)
Apocynaceae         Philibertia nivea (Griseb.) Goyder (7)
Asteraceae          Flourensia riparia Griseb. (17)
Asteraceae          Gnaphalium yalaense Cabrera (1)
Asteraceae          Gutierrezia repens Griseb. (6)
Asteraceae          Hieracium niederleinii (Zahn) Sleumer (6)
Asteraceae          Hieracium tucumanicum (Zahn) Sleumer (8)
Asteraceae          Holocheilus fabrisii Cabrera (4)
Asteraceae          Hysterionica aberrans (Cabrera) Cabrera (4)
Asteraceae          Senecio cremeiflorus Mattf. (11)
Asteraceae          Senecio flagellifolius Cabrera (4)
Asteraceae          Trichocline exscapa Griseb. (17)
Berberidaceae       Berberis lilloana Job (11)
Brassicaceae        Draba burkartiana O.E. Schulz (4)
Brassicaceae        Parodiodoxa chionophila (Speg.) O.E. Schulz (8)
Brassicaceae        Polypsecadium tucumanense (O.E. Schulz)
                      Al-Shehbaz (2)
Bromeliaceae        Puya volcanensis Castillon (3)
Bromeliaceae        Tillandsia zecheri W. Till (2)
Cactaceae           Gymnocalycium spegazzinii Britton & Rose (28)
Cactaceae           Rebutia deminuta (F.A.C. Weber) A. Berger (3)
Caryophyllaceae     Stellaria cryptopetala Griseb. (22)
Convolvulaceae      Ipomoea lilloana O'Donell (6)
Cucurbitaceae       Pteropepon argentinense Mart. Crov. (7)
Cyperaceae          Carex pseudomacloviana G.A. Wheeler (1)
Fabaceae            Astragalus joergensenii I.M. Johnst. (9)
Fabaceae            Lupinus austrorientalis C.P. Sm. (5)
Fabaceae            Lupinus ultramontanus C.P. Sm. (3)
Gentianaceae        Gentianella bromifolia (Griseb.) T.N. Ho & S.W.
                      Liu (6)
Gentianaceae        Gentianella hieronymi (Gilg) Fabris (16)
Geraniaceae         Geranium leucanthum Griseb. (11)
Iridaceae           Ennealophus fimbriatus Ravenna (12)
Martyniaceae        Craniolaria argentina Speg. (7)
Montiaceae          Phemeranthus punae (R.E. Fr.) Eggli & Nyffeler (7)
Poaceae             Festuca superba Parodi ex Turpe (14)
Poaceae             Festuca uninodis Hack. (3)
Poaceae             Nassella arcaensis (Speg.) Torres (12)
Poaceae             Nassella parva Torres (8)
Poaceae             Poa ragonesei Nicora (2)
Poaceae             Tragus andicola Zapater & Sulekic (17)
Polygonaceae        Rumex lorentzianus Lindau (6)
Rosaceae            Tetraglochin paucijugatum (I.M. Johnst.) Rothm. (8)
Rubiaceae           Manettia jorgensenii Standl.(9)
Solanaceae          Solanum montigenum (C.V. Morton) Cabrera (6)
Solanaceae          Solanum vernei Bitter & Wittm. (6)
Urticaceae          Urtica lilloi (Hauman) Geltman (4)
Valerianaceae       Valeriana polybotrya (Griseb.) Hock (10)
Verbenaceae         Glandularia lilloana (Moldenke) Botta (17)
Verbenaceae         Lantana magnibracteata Tronc.(13)
Verbenaceae         Lantana tilcarensis Tronc. (11)
Violaceae           Viola castillonii (W. Becker) Xifreda & Sanso(9)
Amaryllidaceae      Zephyranthes diluta Ravenna (4)
Asteraceae          Baccharis kurtziana Ariza (6)
Asteraceae          Baccharis niederleinii Heering (4)
Asteraceae          Cabreraea andina (Cabrera) Bonifacino (5)
Asteraceae          Flourensia hirta S.F. Blake (9)
Asteraceae          Senecio diaguita Cabrera (8)
Asteraceae          Senecio sanagastae Cabrera (3)
Asteraceae          Tagetes riojana M. Ferraro (4)
Cactaceae           Lobivia famatimensis (Speg.) Britton & Rose (4)
Cactaceae           Pyrrhocactus kattermannii R. Kiesling (2)
Cactaceae           Tephrocactus alexanderi (Britton & Rose)
                      Backeb. (11)
Fabaceae            Adesmia hunzikeri Burkart (5)
Fabaceae            Adesmia nanolignea Burkart (8)
Fabaceae            Senna fabrisii (L. Bravo) H.S. Irwin & Barneby (7)
Sapindaceae         Guindilia cristata (Radlk.) Hunz. (7)
Solanaceae          Sclerophylax kurtzii Di Fulvio (19)
Solanaceae          Solanum kurtzianum Bitter & Wittm. (3)
Amaryllidaceae      Hieronymiella marginata (Pax) Hunz. (15)
Anacampserotaceae   Anacampseros kurtzii Bacigalupo
Apiaceae            Mulinum famatinense H. Wolff (6)
Asteraceae          Baccharis rupestris Heering (17)
Asteraceae          Conyza cordata Kuntze (9)
Asteraceae          Chersodoma argentina Cabrera (24)
Asteraceae          Chersodoma glabriuscula (Cabrera) M.O. Dillon &
                      Sagast. (8)
Asteraceae          Chuquiraga calchaquina Cabrera (6)
Asteraceae          Hieracium streptochaetum Zahn (8)
Asteraceae          Hyaloseris rubicunda Griseb. (21)
Asteraceae          Mutisia kurtzii R.E. Fr. (29)
Asteraceae          Senecio argophylloides Griseb. (12)
Asteraceae          Senecio cylindrocephalus Cabrera (5)
Asteraceae          Senecio friesii Cabrera (9)
Asteraceae          Senecio octolepis Griseb. (21)
Asteraceae          Senecio pseudotites Griseb. (8)
Asteraceae          Senecio schreiteri Cabrera (8)
Brassicaceae        Draba tucumanensis O.E. Schulz (6)
Brassicaceae        Lepidium argentinum Thell. (8)
Bromeliaceae        Deuterocohnia haumanii A. Cast. (7)
Cactaceae           Tephrocactus weberi (Speg.) Backeb. (11)
Cactaceae           Trichocereus terscheckii (Parm. ex Pfeiff.)
                      Britton & Rose (23)
Caryophyllaceae     Pycnophyllum convexum Griseb. (14)
Euphorbiaceae       Euphorbia marayensis Subils (3)
Ledocarpaceae       Balbisia calycina (Griseb.) Hunz. & Ariza (30)
Loasaceae           Caiophora mollis (Griseb.) Urb. & Gilg (6)
Nyctaginaceae       Mirabilis bracteosa (Griseb.) Heimerl (17)
Poaceae             Bromus flexuosus Planchuelo (12)
Poaceae             Jarava scabrifolia (Torres) Penailillo (8)
Poaceae             Nassella caespitosa Griseb. (29)
Poaceae             Panicum chloroleucum Griseb. (27)
Poaceae             Poa dolichophylla Hack. (19)
Poaceae             Poa hieronymi Hack. (9)
Poaceae             Poa nubensis Giussani, Fernandez Pepi &
                      Morrone (11)
Poaceae             Sporobolus maximus Hauman (7)
Polygalaceae        Polygala argentiniensis Chodat (10)
Polygalaceae        Polygala jujuyensis Grondona (10)
Solanaceae          Fabiana friesii Dammer (9)
Solanaceae          Jaborosa sativa (Miers) Hunz. & Barboza (10)
Solanaceae          Lycium schreiteri F.A. Barkley (8)
Solanaceae          Sclerophylax adnatifolia Di Fulvio (23)
Solanaceae          Sclerophylax caducifructus Di Fulvio (7)
Solanaceae          Sclerophylax tenuicaulis Di Fulvio (10)
Solanaceae          Solanum spegazzinii Bitter (8)
Verbenaceae         Aloysia castellanosii Moldenke (15)
Violaceae           Viola hieronymi W. Becker (2)
Zygophyllaceae      Bulnesia schickendantzii Hieron. ex Griseb. (40)
Zygophyllaceae      Plectrocarpa rougesii Descole, O[acute accent]Donell &
                      Lourteig (18)
Cactaceae           Gymnocalycium saglionis (Cels) Britton & Rose (35)
Fabaceae            Adesmia cytisoides Griseb. (14)
Fabaceae            Hoffmannseggia pumilio (Griseb.) B.B. Simpson (19)
Onagraceae          Oenothera lasiocarpa Griseb. (8)
Oxalidaceae         Oxalis famatinae R. Knuth (24)
Poaceae             Jarava media (Speg.) Penailillo (24)
Amaryllidaceae      Hieronymiella aurea Ravenna (5)
Apocynaceae         Philibertia subnivea (Malme) Goyder (1)
Asteraceae          Porophyllum cabrerae D.J.N. Hind (2)
Asteraceae          Verbesina saltensis Cabrera (1)
Boraginaceae        Ixorhea tschudiana Fenzl (10)
Bromeliaceae        Puya castellanosii L.B. Sm. (6)
Bromeliaceae        Puya weberiana E. Morren ex Mez (5)
Bromeliaceae        Tillandsia albertiana Verv. (1)
Cactaceae           Acanthocalycium thionanthum (Speg.) Backeb. (11)
Cactaceae           Gymnocalycium marsoneri Fric ex Y. Ito (2)
Cactaceae           Lobivia korethroides (Werderm.) Werderm. (2)
Cactaceae           Parodia aureicentra Backeb. (3)
Cactaceae           Parodia penicillata Fechser & Steeg (1)
Cactaceae           Tephrocactus molinensis (Speg.) Backeb. (3)
Cactaceae           Trichocereus angelesii R. Kiesling (2)
Fabaceae            Senna rigidicaulis (Burkart ex L. Bravo) H.S.
                      Irwin & Barneby (2)
Geraniaceae         Geranium tafiense Aedo & Munoz Garm. (2)
Malvaceae           Nototriche lorentzii A.W. Hill (1)
Solanaceae          Solanum incurvipilum Bitter (1)
Solanaceae          Solanum salamancae Hunz. & Barboza (3)
Asteraceae          Baccharis petrophila R.E. Fr. (2)
Bromeliaceae        Tillandsia brealitoensis L. Hrom. (1)
Cactaceae           Lobivia chrysantha (Werderm.) Backeb. (2)
Cactaceae           Lobivia walteri R. Kiesling (1)
Cactaceae           Trichocereus smrzianus (Backeb.) Backeb. (3)
Dioscoreaceae       Dioscorea stenopetala Hauman (6)
Euphorbiaceae       Euphorbia vervoorstii Subils (2)
Orchidaceae         Pelexia ovatifolia M.N. Correa (1)
Poaceae             Glyceria saltensis Sulekic & Rugolo (1)
Asteraceae          Senecio fabrisii Cabrera (2)
Asteraceae          Senecio vervoorstii Cabrera (2)
Cactaceae           Lobivia haematantha (Speg.) Britton & Rose (4)
Gentianaceae        Gentianella pulla (Griseb.) T.N. Ho & S.W. Liu (5)
Malvaceae           Tarasa trisecta (Griseb.) Krapov. (7)
Oxalidaceae         Oxalis sleumeri Lourteig (3)
Poaceae             Poa cabreriana Anton & Ariza (5)
Solanaceae          Sclerophylax cocuccii Di Fulvio (5)
Amaranthaceae       Alternanthera cana Suess. (2)
Amaranthaceae       Gomphrena cladotrichoides Suess. (8)
Asteraceae          Chuquiraga echegarayi Hieron. (4)
Asteraceae          Senecio calingastensis Tombesi (2)
Boraginaceae        Heliotropium ruiz-lealii I.M. Johnst. (3)
Brassicaceae        Descurainia brevifructa Boelcke ex
                      Mart.-Laborde (1)
Cactaceae           Pterocactus megliolii R. Kiesling (2)
Cactaceae           Pyrrhocactus sanjuanensis (Speg.) Backeb. (3)
Cactaceae           Tephrocactus halophilus (Speg.) Backeb. (1)
Fabaceae            Prosopis calingastana Burkart (2)
Violaceae           Viola roigii Rossow (1)
Asteraceae          Senecio belenensis Griseb. (1)
Asteraceae          Senecio delicatulus Cabrera & Zardini (2)
Asteraceae          Senecio lilloi Cabrera (1)
Cactaceae           Puna bonniae D.J. Ferguson & R. Kiesling (1)
Cactaceae           Tephrocactus geometricus (A. Cast.) Backeb. (4)
Calceolariaceae     Calceolaria lepidota Kraenzl. (1)
Caryophyllaceae     Silene margaritae Bocquet (1)
Malvaceae           Lecanophora jarae (Phil.) Krapov. (4)
Malvaceae           Nototriche chuculaensis Krapov. (1)
Malvaceae           Nototriche viridula Krapov. (1)
Polemoniaceae       Giliastrum castellanosii J.M. Porter (2)
Solanaceae          Jaborosa cabrerae Barboza (3)
Amaranthaceae       Alternanthera cinerella Suess. (3)
Apiaceae            Bowlesia venturii Mathias & Constance (2)
Apiaceae            Eryngium lorentzii H. Wolff (2)
Apocynaceae         Cynanchum samuelssonii Malme (1)
Apocynaceae         Jobinia glossostelma (Lillo ex T. Mey.) Liede &
                      Meve (2)
Apocynaceae         Petalostelma sarcostemma (Lillo) Liede & Meve (3)
Aristolochiaceae    Aristolochia melanoglossa Speg. (4)
Asteraceae          Baccharis cabrerae Ariza (1)
Asteraceae          Baccharis polygama Ariza (2)
Asteraceae          Baccharis rodriguezii Ariza (2)
Asteraceae          Hieracium cienegae Zahn (4)
Asteraceae          Leptostelma tucumanense (Cabrera) A. Teles (2)
Asteraceae          Lomanthus calchaquinus (Cabrera) B. Nord. &
                      Pelser (2)
Asteraceae          Microliabum eremophilum (Cabrera) H. Rob. (1)
Asteraceae          Mikania minima (Baker) B.L. Rob. (2)
Asteraceae          Mikania siambonensis Hieron. (1)
Asteraceae          Senecio asplenifolius Griseb. (3)
Asteraceae          Senecio cajonensis Cabrera (1)
Asteraceae          Senecio kunturinus Cabrera (2)
Asteraceae          Senecio maculatus Cabrera (2)
Asteraceae          Senecio roripifolius Cabrera (3)
Asteraceae          Senecio tucumanensis Cabrera (2)
Asteraceae          Tagetes rupestris Cabrera (4)
Begoniaceae         Begonia tafiensis Lillo (8)
Cactaceae           Acanthocalycium ferrarii Rausch (2)
Cactaceae           Gymnocalycium bayrianum H. Till (4)
Cactaceae           Lobivia bruchii Britton & Rose (4)
Cactaceae           Lobivia schreiteri A. Cast. (1)
Cactaceae           Trichocereus schickendantzii (F.A.C. Weber)
                      Britton & Rose (3)
Caryophyllaceae     Stellaria aphanantha Griseb. (1)
Cyperaceae          Carex tucumanensis G.A. Wheeler (1)
Dioscoreaceae       Dioscorea entomophila Hauman (2)
Fabaceae            Sophora rhynchocarpa Griseb. (3)
Hypoxidaceae        Hypoxis catamarcensis Brackett (1)
Iridaceae           Mastigostyla johnstoni R.C. Foster (1)
Iridaceae           Sisyrinchium tucumanum Ravenna (3)
Loasaceae           Caiophora aconquijae Sleumer (6)
Malvaceae           Nototriche caesia A. W. Hill (3)
Malvaceae           Nototriche cajonensis Krapov. (2)
Malvaceae           Nototriche calchaquensis Krapov. (2)
Malvaceae           Nototriche rohmederi Krapov. (2)
Malvaceae           Nototriche tucumana Krapov. (1)
Montiaceae          Schreiteria macrocarpa (Speg.) Carolin (1)
Onagraceae          Oenothera pedunculifolia W. Dietr. (4)
Orchidaceae         Chloraea castillonii Hauman (3)
Orchidaceae         Chloraea phoenicea Speg. (2)
Plantaginaceae      Plantago venturii Pilg. (3)
Poaceae             Nassella fabrisii Torres (6)
Poaceae             Nassella leptothera (Speg.) Torres (3)
Ranunculaceae       Ranunculus hilii Lourteig (4)
Rosaceae            Lachemilla grisebachiana (L.M. Perry) Rothm. (1)
Solanaceae          Cestrum kunthii Francey (8)
Solanaceae          Eriolarynx iochromoides (Hunz.) Hunz. (3)
Solanaceae          Eriolarynx lorentzii (Dammer) Hunz. (8)
Solanaceae          Jaborosa oxipetala Speg. (3)
Solanaceae          Solanum sanctae-rosae Hawkes (5)
Valerianaceae       Valeriana lasiocarpa Griseb. (2)
Violaceae           Viola calchaquiensis W. Becker (1)
Violaceae           Viola lilloana W. Becker (2)
Violaceae           Viola munozensis W. Becker (1)
Violaceae           Viola rodriguezii W. Becker (4)
Violaceae           Viola tucumanensis W. Becker (2)
Acanthaceae         Justicia hunzikeri Ariza (5)
Cactaceae           Gymnocalycium castellanosii Backeb. (15)
Fabaceae            Ramorinoa girolae Speg. (6)
Poaceae             Neobouteloua paucirracemosa M.G. Lopez &
                      Biurrun (4)
Amaryllidaceae      Chlidanthus yaviensis (Ravenna) Ravenna (1)
Asteraceae          Antennaria sleumeri Cabrera (1)
Asteraceae          Hieracium neofurcatum Sleumer (2)
Asteraceae          Stevia okadae Cabrera (1)
Asteraceae          Vernonia centauropsidea Hieron. (3)
Bromeliaceae        Puya yakespala A. Cast. (3)
Cactaceae           Lobivia chrysochete (Werderm.) Wessner (1)
Cactaceae           Lobivia sanguiniflora Backeb. (1)
Cactaceae           Rebutia margarethae Rausch (1)
Gentianaceae        Gentianella cabrerae (Fabris) Fabris (1)
Iridaceae           Mastigostyla brachiandra Ravenna (1)
Iridaceae           Mastigostyla implicata Ravenna (1)
Malvaceae           Nototriche sleumeri Krapov. (2)
Poaceae             Danthonia rugoloana Sulekic (1)
Poaceae             Nassella yaviensis Torres (1)
Solanaceae          Solanum neorossii Hawkes & Hjert. (2)
Tropaeolaceae       Tropaeolum atrocapillare Sparre (2)
Amaryllidaceae      Habranthus ruizlealii Ravenna (1)
Asteraceae          Chiliotrichiopsis keidelii Cabrera (39)
Asteraceae          Eupatorium saltense Hieron. (31)
Asteraceae          Eupatorium tucumanense Lillo & B.L. Rob. (11)
Asteraceae          Flourensia leptopoda S.F. Blake (4)
Asteraceae          Stevia minor Griseb. (21)
Brassicaceae        Descurainia adpressa Boelcke (6)
Cactaceae           Gymnocalycium ragonesei A. Cast. (1)
Cyperaceae          Carex humahuacaensis G.A. Wheeler (2)
Fabaceae            Adesmia crassicaulis Phil. (4)
Fabaceae            Astragalus crypticus I.M. Johnst. (8)
Gentianaceae        Gentianella punensis (Fabris) Fabris (1)
Poaceae             Festuca nemoralis Turpe (14)
Poaceae             Festuca parodiana (St.-Yves ex Parodi) Nicora (15)
Poaceae             Jarava hystricina (Speg.) Penailillo (10)
Poaceae             Nassella glabripoda Torres (11)
Poaceae             Nassella meyeri Torres (9)
Poaceae             Pappostipa hieronymusii (Pilg.) (4)
Solanaceae          Jaborosa lanigera (Phil.) Hunz. & Barboza (9)
Solanaceae          Solanum annuum C.V. Morton (10)
Violaceae           Viola evae Hieron. ex W. Becker (2)

                                 Cell Size
Familiy              0.2x0.2    0.5x0.5   1x0.5

Alstroemeriaceae                 0.85
Amaryllidaceae        0.92
Apocynaceae                      0.82
Asteraceae                       0.83
Asteraceae                       0.85
Asteraceae                         1
Asteraceae             0.9       0.75

Asteraceae            0.77       0.88
Bromeliaceae                     0.82
Cactaceae                        0.86
Cactaceae                        0.87      0.4
Cactaceae             0.21       0.71
Calyceraceae          0.77       0.88
Convolvulaceae                    0.7     0.78
Dioscoreaceae         0.92
Fabaceae                         0.67
Fabaceae              0.48       0.83

Iridaceae                        0.83
Poaceae               0.77       0.88
Poaceae                            1
Poaceae                          0.79
Poaceae                          0.61
Solanaceae            0.83       0.83
Solanaceae                         1
Verbenaceae           0.83
Violaceae             0.81
Asteraceae                       0.32     0.82
Asteraceae                       0.76      0.7
Asteraceae                       0.74     0.69
Boraginaceae                              0.75
Brassicaceae                      0.7     0.71
Fabaceae                                  0.77
Fabaceae                         0.69     0.63
Fabaceae                                  0.66
Fabaceae                         0.72     0.75
Juncaceae                         0.5     0.55
Malvaceae                        0.81     0.77
Poaceae                          0.67     0.69
Solanaceae                       0.75     0.77
Valerianaceae                              0.7
Violaceae                         0.7     0.83
Asteraceae                       0.72     0.54
Asteraceae                       0.82     0.76
Bromeliaceae                              0.79
Fabaceae                         0.63     0.54
Fabaceae                         0.79     0.47
Oxalidaceae                               0.89
Amaryllidaceae                   0.67
Apocynaceae                      0.79     0.71
Asteraceae                       0.76
Asteraceae            0.78       0.87
Asteraceae                       0.58
Asteraceae                       0.69      0.9
Bromeliaceae                     0.63
Cactaceae              0.8        0.8

Cactaceae             0.69       0.85
Cactaceae                        0.78

Cactaceae                        0.84     0.62
Cactaceae                        0.83     0.64
Cactaceae                        0.78
Cactaceae                        0.85

Cactaceae                        0.81
Fabaceae                          0.8
Iridaceae                        0.64     0.77
Ledocarpaceae                    0.78
Malvaceae                        0.58
Solanaceae                       0.73
Zygophyllaceae                   0.73
Acanthaceae                      0.68
Amaryllidaceae                   0.54     0.67
Asteraceae                       0.77
Asteraceae                       0.58
Asteraceae                       0.81     0.61
Asteraceae                       0.76      0.7
Asteraceae                                0.58
Bromeliaceae                              0.58
Cactaceae                                  0.5
Cactaceae                        0.77

Cactaceae                                  0.7

Cactaceae                        0.73      0.8
Cactaceae                                 0.72

Fabaceae                         0.64
Fabaceae              0.86       0.71
Fabaceae              0.59       0.62
Gentianaceae                     0.77

Gentianaceae                     0.39
Poaceae                          0.75     0.69
Ranunculaceae                    0.74
Rosaceae              0.83       0.74
Solanaceae                       0.86
Verbenaceae                      0.47     0.68
Violaceae                        0.78     0.73
Amaryllidaceae                   0.62
Asteraceae            0.63
Asteraceae            0.69
Asteraceae                        0.6     0.57
Asteraceae            0.86       0.88
Asteraceae            0.74       0.75
Asteraceae            0.86       0.88
Asteraceae                       0.73     0.73
Asteraceae            0.64       0.88
Brassicaceae          0.86       0.88
Cactaceae             0.91        0.8     0.83
Cactaceae             0.44       0.88

Cactaceae             0.85
Fabaceae              0.86       0.88
Gentianaceae                     0.92

Malvaceae             0.86       0.88
Malvaceae             0.86       0.88
Malvaceae             0.86       0.75
Malvaceae             0.87       0.88
Malvaceae             0.87       0.88
Verbenaceae                      0.76
Asteraceae                       0.46     0.84
Cactaceae                         0.7
Cactaceae                        0.91     0.81
Cactaceae                        0.76     0.71
Cactaceae                        0.81
Cactaceae                        0.77     0.79
Anacampserotaceae                0.89     0.38
Apocynaceae           0.84         1

Asteraceae            0.79       0.88
Asteraceae            0.05       0.89     0.57
Asteraceae            0.81         1
Asteraceae            0.81         1
Asteraceae            0.76       0.81     0.54
Asteraceae            0.86         1       0.5
Asteraceae             0.9                0.88
Asteraceae            0.83       0.93     0.26
Asteraceae            0.76       0.76
Asteraceae            0.64       0.88
Asteraceae            0.79         1
Asteraceae            0.62         1       0.5
Bromeliaceae          0.89         1       0.5
Cactaceae             0.85       0.92      0.5
Cactaceae              0.8       0.79
Cactaceae             0.77
Cactaceae             0.84                0.65
Cactaceae             0.69       0.92      0.5
Cactaceae                        0.66     0.74
Cactaceae             0.86         1       0.5
Cactaceae             0.86         1       0.5
Cactaceae             0.92       0.69

Caryophyllaceae       0.79       0.88
Convolvulaceae        0.87       0.78     0.51
Crassulaceae          0.65         1       0.5
Dioscoreaceae         0.75         1       0.5
Fabaceae              0.48                 0.8
Malvaceae             0.81                0.65
Malvaceae             0.84       0.88
Orchidaceae           0.71       0.96
Plantaginaceae        0.91       0.92      0.5
Urticaceae                       0.81
Velloziaceae          0.86         1       0.5
Verbenaceae           0.46       0.97      0.5

Acanthaceae           0.83
Apiaceae                                  0.81
Apocynaceae                      0.77
Apocynaceae                      0.55
Apocynaceae                                0.8
Aristolochiaceae                 0.71
Asteraceae            0.54       0.69     0.65
Asteraceae            0.09        0.7
Asteraceae            0.83       0.66     0.68
Asteraceae            0.27       0.66
Asteraceae             0.8       0.79
Asteraceae                        0.8
Brassicaceae                     0.81

Brassicaceae           0.7        0.9      0.5
Brassicaceae          0.92       0.66     0.68
Bromeliaceae                              0.88
Bromeliaceae                     0.76
Cactaceae             0.83
Celastraceae          0.88       0.79
Cucurbitaceae          0.8
Euphorbiaceae                    0.75
Fabaceae
Fabaceae
Gentianaceae                     0.68     0.75
Iridaceae                        0.73
Malvaceae             0.92       0.66     0.68
Piperaceae                        0.8
Poaceae                                   0.73
Poaceae                                   0.88
Poaceae                0.9        0.7
Poaceae                           0.7

Poaceae                          0.72     0.82
Rubiaceae             0.33       0.67
Solanaceae                       0.66
Solanaceae                       0.85
Tropaeolaceae                    0.68     0.77
Amaryllidaceae                   0.77
Apocynaceae                      0.77
Asteraceae                       0.69     0.87
Asteraceae                        0.9     0.83
Asteraceae                       0.57
Asteraceae            0.88       0.88      0.8
Asteraceae                       0.78     0.88
Asteraceae                        0.8
Asteraceae                       0.88     0.78
Begoniaceae            0.8       0.88
Brassicaceae          0.82       0.88     0.51
Brassicaceae                     0.78     0.78
Cactaceae                        0.86     0.76
Cactaceae                         0.6      0.8
Cactaceae                        0.83
Cactaceae                        0.74     0.76
Caryophyllaceae                  0.75
Caryophyllaceae                  0.75     0.74
Convolvulaceae                   0.74
Fabaceae                         0.83     0.85
Fabaceae              0.16        0.8     0.93
Malvaceae                        0.79     0.79
Malvaceae             0.57       0.77      0.8
Orobanchaceae                    0.77     0.63
Poaceae                0.8       0.94
Poaceae                           0.7     0.83
Poaceae                          0.85
Solanaceae                       0.66
Valerianaceae                    0.73     0.87
Amaryllidaceae                   0.73
Anthericaceae                    0.81     0.79
Apiaceae                         0.79     0.85
Apocynaceae           0.81       0.88      0.8

Apocynaceae                      0.85
Apocynaceae           0.82       0.85     0.78
Asteraceae                       0.76     0.78
Asteraceae                       0.89     0.79
Bromeliaceae                     0.76     0.79
Bromeliaceae                     0.77     0.82
Bromeliaceae                     0.82     0.74
Cactaceae                        0.79     0.88
Cactaceae                        0.72

Euphorbiaceae         0.83       0.83     0.71

Fabaceae                         0.73     0.46
Gentianaceae                     0.79     0.61
Gentianaceae                     0.87     0.77
Iridaceae                        0.89     0.73
Iridaceae                        0.81     0.87

Loasaceae                        0.75     0. 80
Malvaceae                        0.83     0.84
Orchidaceae                       0.9     0.83
Orchidaceae                      0.79
Solanaceae                       0.69     0.79
Solanaceae                       0.83     0.72
Solanaceae                       0.93     0.88
Tropaeolaceae                    0.76     0.62
Valerianaceae         0.46       0.97     0.86
Acanthaceae                       0.8     0.66
Alstroemeriaceae                 0.84     0.91
Alliaceae                        0.83     0.88
Amaranthaceae                    0.68     0.45
Amaryllidaceae                   0.77     0.93
Anacardiaceae                    0.66     0.68
Anthericaceae                    0.78     0.86
Apiaceae                         0.72      0.8

Apocynaceae                       0.7     0.43
Asteraceae                       0.49     0.53
Asteraceae            0.82         1       0.5
Asteraceae                       0.75      0.6
Asteraceae            0.77       0.73     0.88
Asteraceae                       0.77     0.91
Asteraceae                       0.76     0.66
Asteraceae                       0.85     0.73
Asteraceae                       0.77     0.78
Asteraceae                       0.73     0.76
Asteraceae                       0.65     0.75
Berberidaceae                     0.8     0.72
Brassicaceae                     0.79      0.8
Brassicaceae                     0.64     0.75
Brassicaceae          0.79       0.84

Bromeliaceae                     0.78
Bromeliaceae          0.44       0.74
Cactaceae                        0.57     0.71
Cactaceae                        0.72     0.71
Caryophyllaceae                  0.77     0.73
Convolvulaceae                   0.59      0.9
Cucurbitaceae                    0.81     0.81
Cyperaceae            0.86       0.73
Fabaceae                         0.75     0.88
Fabaceae                         0.73     0.71
Fabaceae                         0.73     0.78
Gentianaceae          0.76       0.83     0.89

Gentianaceae                     0.81     0.82
Geraniaceae                      0.78     0.85
Iridaceae                        0.76     0.79
Martyniaceae                     0.66     0.72
Montiaceae                       0.76     0.73
Poaceae                          0.83     0.83
Poaceae
Poaceae                           0.7     0.76
Poaceae                          0.76     0.75
Poaceae                          0.59
Poaceae                          0.78     0.88
Polygonaceae          0.78       0.85     0.89
Rosaceae                         0.86     0.51
Rubiaceae                        0.81      0.9
Solanaceae                       0.68     0.79
Solanaceae                       0.77     0.85
Urticaceae                       0.69
Valerianaceae                    0.75     0.64
Verbenaceae                      0.55     0.52
Verbenaceae                      0.46      0.9
Verbenaceae                      0.72     0.79
Violaceae                        0.79     0.76
Amaryllidaceae                             0.7
Asteraceae                                0.89
Asteraceae                                0.68
Asteraceae                                0.51
Asteraceae                                 0.7
Asteraceae                                0.73
Asteraceae                                0.74
Asteraceae                                0.84
Cactaceae                                  0.8
Cactaceae                                 0.71
Cactaceae                                 0.62

Fabaceae                                  0.71
Fabaceae                                  0.45
Fabaceae                                   0.8
Sapindaceae                               0.64
Solanaceae                                0.71
Solanaceae                                0.74
Amaryllidaceae                            0.77
Anacampserotaceae
Apiaceae                         0.61     0.66
Asteraceae
Asteraceae                       0.51     0.65
Asteraceae                       0.44     0.43
Asteraceae                                0.68

Asteraceae                                 0.7
Asteraceae                                0.58
Asteraceae                                0.78
Asteraceae                       0.69     0.69
Asteraceae                                0.75
Asteraceae                                0.72
Asteraceae
Asteraceae                                0.85
Asteraceae                                0.68
Asteraceae                                0.58
Brassicaceae                     0.68      0.7
Brassicaceae                     0.74     0.71
Bromeliaceae                              0.72
Cactaceae                                 0.54
Cactaceae                                 0.57

Caryophyllaceae                  0.61     0.68
Euphorbiaceae
Ledocarpaceae                             0.89
Loasaceae                                 0.59
Nyctaginaceae                    0.72     0.59
Poaceae                          0.68     0.62
Poaceae                                   0.65
Poaceae                          0.74     0.65
Poaceae                           0.8     0.82
Poaceae                                   0.63
Poaceae                           0.8     0.77
Poaceae

Poaceae                          0,72     0,76
Polygalaceae                     0,65     0,68
Polygalaceae                     0,79     0,81
Solanaceae                                0,78
Solanaceae                                0,47
Solanaceae                                0,76
Solanaceae                                0,81
Solanaceae                                0,77
Solanaceae                       0,74     0,84
Solanaceae                       0,62     0,82
Verbenaceae                               0,74
Violaceae
Zygophyllaceae                   0,61     0,74
Zygophyllaceae                   0,77     0,81

Cactaceae
Fabaceae                         0,84
Fabaceae                                  0,54
Onagraceae                       0,72
Oxalidaceae                      0,61
Poaceae                          0,71
Amaryllidaceae                   0,93     0,73
Apocynaceae           0,91       0,88
Asteraceae            0,44       0,90
Asteraceae            0,75       0,88
Boraginaceae                     0,85     0,86
Bromeliaceae          0,60       0,87
Bromeliaceae                     0,89
Bromeliaceae          0,85       1,00
Cactaceae                        0,93
Cactaceae                        0,82     0,88
Cactaceae                        0,79
Cactaceae                        0,80
Cactaceae             0,92       1,00
Cactaceae                        0,80
Cactaceae             0,75       0,82
Fabaceae              0,50       0,88

Geraniaceae                      0,80     0, 75
Malvaceae             0,90       0,90
Solanaceae            0,81       0,88
Solanaceae                       0,80
Asteraceae                       0,89     0,85
Bromeliaceae          0,86
Cactaceae             0,53       0,90
Cactaceae             1,00
Cactaceae             1,00
Dioscoreaceae                    0,74     0,79
Euphorbiaceae                    0,83     0,64
Orchidaceae                      0,75
Poaceae               1,00
Asteraceae                       0,61     0,68
Asteraceae                       0,70
Cactaceae                        0,74
Gentianaceae                     0,51
Malvaceae                        0,66     0,85
Oxalidaceae                      0,73     0,73
Poaceae                          0,46     0,53
Solanaceae                       0,86     0,74
Amaranthaceae                    0,74     0,82
Amaranthaceae                    0,61     0,91
Asteraceae                       0,76     0,73
Asteraceae                       0,64     0,86
Boraginaceae                     0,90     0,83
Brassicaceae                     0,69     0,72

Cactaceae                        0,69     0,75
Cactaceae                        0,82     0,62
Cactaceae
Fabaceae                         0,73     0,71
Violaceae                        0,69     0,75
Asteraceae            0,77       0,80
Asteraceae            0,93       1,00
Asteraceae            0,68       0,88
Cactaceae
Cactaceae                        0,80
Calceolariaceae       0,92       0,90
Caryophyllaceae       0,77       0,80
Malvaceae                        0,48
Malvaceae             0,75       0,88
Malvaceae                        0,75
Polemoniaceae         0,68       0,88
Solanaceae                       0,75
Amaranthaceae         0,65       0,85
Apiaceae              0,79       0,84
Apiaceae              0,75       0,88     1,00
Apocynaceae           0,76       1,00     1,00
Apocynaceae                      0,81

Apocynaceae                      0,54
Aristolochiaceae                 0,70
Asteraceae            0,92       0,88
Asteraceae            0,92
Asteraceae            0,92       0,88
Asteraceae            0.93       0.84
Asteraceae            0.88        0.8       1
Asteraceae            0.78        0.8

Asteraceae            0.92       0.88
Asteraceae            0.32       0.87
Asteraceae            0.85        0.8
Asteraceae            0.77       0.83
Asteraceae            0.81       0.69
Asteraceae             0.8       0.81
Asteraceae            0.85       0.82
Asteraceae            0.78       0.88
Asteraceae            0.81       0.92
Asteraceae            0.76       0.88
Begoniaceae           0.68       0.85
Cactaceae             0.77       0.92
Cactaceae                        0.69     0.79
Cactaceae              0.8       0.74     0.82
Cactaceae             0.74       0.72
Cactaceae             0.39       0.78

Caryophyllaceae       0.85
Cyperaceae            0.75       0.88
Dioscoreaceae                    0.66     0.86
Fabaceae                         0.94
Hypoxidaceae          0.81       0.71     0.88
Iridaceae             0.75       0.88
Iridaceae             0.69       0.78
Loasaceae             0.88       0.86
Malvaceae                        0.72
Malvaceae             0.85
Malvaceae             0.75       0.84
Malvaceae             0.91        0.8
Malvaceae             0.86
Montiaceae            0.86
Onagraceae                       0.89
Orchidaceae           0.77       0.92
Orchidaceae           0.68       0.88     0.86
Plantaginaceae        0.48       0.77
Poaceae                          0.72
Poaceae               0.52       0.79
Ranunculaceae         0.66       0.81
Rosaceae              0.83                0.72
Solanaceae                       0.76
Solanaceae            0.77       0.86
Solanaceae            0.42       0.77
Solanaceae                       0.84
Solanaceae            0.77       0.83
Valerianaceae         0.85       0.75
Violaceae             0.81
Violaceae             0.69       0.77
Violaceae             0.81       0.88
Violaceae             0.67       0.82
Violaceae             0.81       0.85
Acanthaceae
Cactaceae
Fabaceae
Poaceae

Amaryllidaceae        0.74       0.79
Asteraceae            0.57
Asteraceae            0.62       0.73
Asteraceae            0.92
Asteraceae            0.75       0.75
Bromeliaceae          0.92       0.73
Cactaceae             0.71       0.73
Cactaceae             0.81       0.73
Cactaceae             0.86       0.73
Gentianaceae          0.92
Iridaceae             0.86       0.73
Iridaceae             0.74       0.73
Malvaceae             0.82                0.83
Poaceae               0.71       0.79
Poaceae               0.74       0.73
Solanaceae            0.78       0.73
Tropaeolaceae          0.6       0.73
Amaryllidaceae
Asteraceae
Asteraceae
Asteraceae
Asteraceae
Asteraceae
Brassicaceae
Cactaceae
Cyperaceae
Fabaceae
Fabaceae
Gentianaceae
Poaceae
Poaceae
Poaceae
Poaceae
Poaceae
Poaceae
Solanaceae
Solanaceae
Violaceae

Familiy             Altitude range (mas)   Aridity range   Area

Alstroemeriaceae         1580-3000           5.6-15.6       1
Amaryllidaceae              1420               15.7         1
Apocynaceae              1420-3040           12.1-15.7      1
Asteraceae               2700-3720           7.3-18.1       1
Asteraceae               2700-3460           6.3-18.1       1
Asteraceae                  2825               15.9         1
Asteraceae                  3270               11.4         1

Asteraceae                  3460               18.1         1
Bromeliaceae             1690-3410           7.3-12.2       1
Cactaceae                1500-1900            5.6-9.0       1
Cactaceae                1045-1910           5.9-17.5       1
Cactaceae                   2170               14.3         1
Calyceraceae                3460               18.1         1
Convolvulaceae           1160-2930           5.9-15.6       1
Dioscoreaceae               1420               15.7         1
Fabaceae                 2960-3615           6.3-14.1       1
Fabaceae                    2240                6.6         1

Iridaceae                2700-3460           13.3-18.1      1
Poaceae                     3460               18.1         1
Poaceae                   480-520            12.8-12.9      1
Poaceae                     1675               18.1         1
Poaceae                  1670-1960           8.8-18.1       1
Solanaceae                  1710               14.1         1
Solanaceae               730 -1120           12.6-13.9      1
Verbenaceae                 2650                 9          1
Violaceae                   3750               12.5         1
Asteraceae               1000-2150            3.8-5.3       2
Asteraceae               3245-3945            7.3-8.2       2
Asteraceae               3520-4150           7.3-10.6       2
Boraginaceae                2405                 5          2
Brassicaceae             2900-4250            6.5-8.1       2
Fabaceae                 2375-3245            7.8-9.7       2
Fabaceae                 3570-4530           7.4-24.1       2
Fabaceae                 3320-4810           8.7-27.2       2
Fabaceae                    3530                7.5         2
Juncaceae                3550-4440           4.4-24.1       2
Malvaceae                4095-4305           7.6-17.9       2
Poaceae                  3195-4070           8.9-14.8       2
Solanaceae               1650-2590            4.4-5.7       2
Valerianaceae            2910-4090           9.9-19.9       2
Violaceae                3430-4070           12.8-14.8      2
Asteraceae               2030-3480           6.6-21.2       3
Asteraceae               2540-3975           8.4-18.1       3
Bromeliaceae             2540-3600           11.7-17.8      3
Fabaceae                 2160-4060           8.3-19.6       3
Fabaceae                 2411-4030           6.9-19.2       3
Oxalidaceae              2500-3500           9.9-17.0       3
Amaryllidaceae           3680-3750            8.8-8.9       4
Apocynaceae              3460-3705          10.5 -15.2      4
Asteraceae               3355-3810           6.7-18.6       4
Asteraceae               3355-3620           12.1-18.6      4
Asteraceae               2410-3880           9.7-20.4       4
Asteraceae               3360-4380           7.2-13.7       4
Bromeliaceae             2600-2980           12.4-17.8      4
Cactaceae                2540-3620           12.1-17.8      4

Cactaceae                3355-3930           12.1-18.6      4
Cactaceae                2990-3165           13.1-16.1      4

Cactaceae                3340-4090           8.8-18.6       4
Cactaceae                2710-3760           10.0-19.0      4
Cactaceae                3020-3930           13.1-17.8      4
Cactaceae                2500-3190           15.1-17.6      4

Cactaceae                3070-3290           11.4-18.2      4
Fabaceae                 2410-3165           12.3-18.3      4
Iridaceae                3830-4100           6.8-10.1       4
Ledocarpaceae            3110-3355           15.6-19.5      4
Malvaceae                1370-3670            13.8-24       4
Solanaceae               3050-3140           15.0-18.1      4
Zygophyllaceae           2870-3190           15.9-26.3      4
Acanthaceae              1810-3195           5.5-11.1       5
Amaryllidaceae            510-1670           7.6-14.5       5
Asteraceae               2490-3410            7.3-9.6       5
Asteraceae               1240-2860           7.0-10.8       5
Asteraceae               1110-3250           5.5-14.4       5

Asteraceae               2270-4375           7.5-18.1       5
Asteraceae               1030-2780           4.6-10.4       5
Bromeliaceae             1030-2070           5.9-14.7       5
Cactaceae                 750-1960           11.6-19.6      5
Cactaceae                 845-2180           5.7-12.5       5

Cactaceae                 780-2180           8.9-19.7       5

Cactaceae                 920-3690           8.1-12.9       5
Cactaceae                 190-2100           5.4-13.8       5

Fabaceae                  995-1710           10.7-14.1      5
Fabaceae                 3180-4070           12.2-14.8      5
Fabaceae                 1860-4070           11.0-14.8      5
Gentianaceae             2910-3690           9.5-18.1       5

Gentianaceae             3640-3840           4.8-14.8       5
Poaceae                  2700-4470           8.0-18.6       5
Ranunculaceae             670-4080           5.8-15.7       5
Rosaceae                 2560-2700           9.6-13.3       5
Solanaceae                930-2330            5.6-8.9       5
Verbenaceae               760-2100           6.1-15.2       5
Violaceae                2330-4375           8.7-18.1       5
Amaryllidaceae           2815-3030           6.7-10.7       6
Asteraceae                  2330                8.5         6
Asteraceae                  2490                8.8         6
Asteraceae               1550-2100           4.5-10.2       6
Asteraceae                  4070               14.8         6
Asteraceae                  3780                12          6
Asteraceae                  4070               14.8         6
Asteraceae               1770-3195           5.1-10.9       6
Asteraceae               3165-3690           9.8-12.9       6
Brassicaceae                4070               14.8         6
Cactaceae                1405-1860            8.1-8.4       6
Cactaceae                1380-2100            7.0-8.8       6

Cactaceae                1630-1860            7.9-8.4       6
Fabaceae                    2560                9.6         6
Gentianaceae             2250-4090           11.9-15.0      6

Malvaceae                   4070               14.8         6
Malvaceae                   4070               14.8         6
Malvaceae                   3970               13.8         6
Malvaceae                3950-4090           12.6-15.0      6
Malvaceae                3950-4090           12.6-15.0      6
Verbenaceae              1155-1670            5.3-7.6       6
Asteraceae                850-880            8.7-10.8       6 *
Cactaceae                1110-1550           10.1-10.2      6 *
Cactaceae                1260-1360           9.3-11.8       6 *
Cactaceae                 740-1270           7.4-11.4       6 *
Cactaceae                 740-1010           7.2-11.0       6 *
Cactaceae                1425-2240           9.8-11.2       6 *
Anacampserotaceae        2255-2270           13.8-17.0      7
Apocynaceae              1440-3100           11.2-27.7      7

Asteraceae               1220-1495           28.0-28.6      7
Asteraceae               1860-2190           16.4-18.3      7
Asteraceae                  2720               16.7         7
Asteraceae                  2720               16.7         7
Asteraceae               3605-4100           10.1-10.4      7
Asteraceae                  2270                17          7
Asteraceae                  3770                9.6         7
Asteraceae               3950-4270           10.8-14.5      7
Asteraceae               3480-4110           9.4-12.5       7
Asteraceae               2190-3450           13.3-18.3      7
Asteraceae               1920-2190           18.3-23.9      7
Asteraceae               2190-2670           14.2-18.3      7
Bromeliaceae             2190-2970           12.6-18.3      7
Cactaceae                2560-2640           13.8-14.6      7
Cactaceae                3360-3810           9.9-10.7       7
Cactaceae                   3337               10.5         7
Cactaceae                   4100               10.1         7
Cactaceae                2010-3055           11.8-20.7      7
Cactaceae                2750-3510           8.1-10.5       7
Cactaceae                1890-2270           17.0-17.9      7
Cactaceae                   3100               11.2         7
Cactaceae                   2930               10.2         7

Caryophyllaceae             3395               13.2         7
Convolvulaceae           3430-3750           10.0-10.9      7
Crassulaceae             1800-2270           17.0-23.9      7
Dioscoreaceae            2190-2640           14.6-18.3      7
Fabaceae                    3510                8.4         7
Malvaceae                   4750               12.6         7
Malvaceae                   3450                12          7
Orchidaceae              2640-3980           11.2-14.6      7
Plantaginaceae           2670-3900           11.2-14.2      7
Urticaceae               1170-1675           25.8-27.3      7
Velloziaceae             1740-2270           17.0-25.3      7
Verbenaceae              4000-4150           10.8-12.2      7

Acanthaceae                 1420               26.3         7 *
Apiaceae                 3500-4810           5.4-12.7       7 *
Apocynaceae               950-1640           25.7-27.5      7 *
Apocynaceae               480-1860           16.4-29.0      7 *
Apocynaceae                 970                21.5         7 *
Aristolochiaceae         1380-2210           20.3-27.9      7 *
Asteraceae               3520-4100           10.0-10.1      7 *
Asteraceae               1155-1620           25.4-26.8      7 *
Asteraceae                  4400                 7          7 *
Asteraceae                  1880               26.6         7 *
Asteraceae               1495-1630           25.5-28.0      7 *
Asteraceae                  360                33.7         7 *
Brassicaceae              380-1260           23.5-31.8      7 *

Brassicaceae                3750               10.4         7 *
Brassicaceae                4520                7.8         7 *
Bromeliaceae                1110               30.9         7 *
Bromeliaceae             1750-2000           20.6-27.0      7 *
Cactaceae                   2620               11.4         7 *
Celastraceae             1495-1660           25.1-28.0      7 *
Cucurbitaceae               1710               25.5         7 *
Euphorbiaceae               570                25.5         7 *
Fabaceae                 4140-4940              6.8         7 *
Fabaceae                    4070                5.2         7 *
Gentianaceae             2720-3280           13.2-16.6      7 *
Iridaceae                 470-2450           17.7-23.6      7 *
Malvaceae                   4520                7.8         7 *
Piperaceae                  360                33.7         7 *
Poaceae                  4505-4810            5.4-9.0       7 *
Poaceae                     570                20.5         7 *
Poaceae                     1640               26.5         7 *
Poaceae                  3230-3430           9.2-10.9       7 *

Poaceae                  2530-4010           7.9-13.7       7 *
Rubiaceae                1290-1770           24.1-29.1      7 *
Solanaceae                  1480                28          7 *
Solanaceae               1355-1750           21.2-27.0      7 *
Tropaeolaceae             350-460            24.2-27.9      7 *
Amaryllidaceae           2075-3230           9.2-21.6      7 **
Apocynaceae               785-1390           22.7-31.6     7 **
Asteraceae               2650-3750           8.0-15.4      7 **
Asteraceae               2750-3900           12.0-17.3     7 **
Asteraceae               1495-2190          18.3-28. 0     7 **
Asteraceae               2670-4250           11.2-17.3     7 **
Asteraceae               3460-4350           6.2-15.2      7 **
Asteraceae               1155-1470           26.7-30.3     7 **
Asteraceae               2600-4330           8.1-18.9      7 **
Begoniaceae              2600-3280           11.9-18.9     7 **
Brassicaceae             2530-4100          10.1-13. 7     7 **
Brassicaceae             2270-4140           6.8-17.0      7 **
Cactaceae                2940-4160           7.1-12.9      7 **
Cactaceae                2610-2930           8.3-13.5      7 **
Cactaceae                2640-3890           10.5-13.7     7 **
Cactaceae                2255-3030           9.0-20.1      7 **
Caryophyllaceae          2670-4150           11.8-16.6     7 **
Caryophyllaceae          3770-4110           6.6-13.7      7 **
Convolvulaceae           1590-2460           17.0-26.7     7 **
Fabaceae                 2270-4110           8.4-17.0      7 **
Fabaceae                 3500-4070           9.0-10.5      7 **
Malvaceae                4000-4410           10.1-16.5     7 **
Malvaceae                4010-5080           14.6-16.5     7 **
Orobanchaceae            1605-4160           10.7-27.7     7 **
Poaceae                  3100-3680           11.2-15.6     7 **
Poaceae                  3260-3830           7.0-11.8      7 **
Poaceae                  2270-4000           8.4-17.0      7 **
Solanaceae               1390-2680           16.5-30.3     7 **
Valerianaceae            4080-4810           8.4-16.6      7 **
Amaryllidaceae           1200-2860           11.8-34.7      8
Anthericaceae            1540-2270           15.2-28.0      8
Apiaceae                 2385-4150           11.8-16.9      8
Apocynaceae              1150-1500           17.9-40.4      8

Apocynaceae              1330-3100           11.4-26.8      8
Apocynaceae              1150-1920           21.3-40.3      8
Asteraceae               2110-2640           6.9-17.0       8
Asteraceae               2270-3980           11.4-17.0      8
Bromeliaceae              830-2730           13.8-34.4      8
Bromeliaceae              710-3180           12.2-24.8      8
Bromeliaceae             2110-3290           6.9-11.2       8
Cactaceae                1070-2710           10.3-28.0      8
Cactaceae                 460-1465           11.2-33.3      8

Euphorbiaceae             820-2070           17.1-27.4      8

Fabaceae                 1705-4670           8.5-20.1       8
Gentianaceae             1760-2190           9.7-18.3       8
Gentianaceae             3100-4250           9.5-14.8       8
Iridaceae                3950-4110           13.6-16.2      8
Iridaceae                2700-3040           12.2-13.3      8

Loasaceae                2830-4610           7.5-19.1       8
Malvaceae                2850-3890           8.9-13.4       8
Orchidaceae              1820-2970           12.6-22.0      8
Orchidaceae               460-1590           24.5-32.5      8
Solanaceae                390-1495           23.4-33.6      8
Solanaceae                710-1920           22.8-36.6      8
Solanaceae               2140-3040           10.7-20.3      8
Tropaeolaceae             790-2270           10.8-29.0      8
Valerianaceae            1150-3040           12.2-40.3      8
Acanthaceae              2110-2170            12.6-17       8 *
Alstroemeriaceae         1580-3250           8.0-22.0       8 *
Alliaceae                2640-3100           7.2-18.0       8 *
Amaranthaceae            2650-3660           5.2-12.9       8 *
Amaryllidaceae           1160-4035           6.8-15.2       8 *
Anacardiaceae             690-3460           8.75-28.2      8 *
Anthericaceae            1110-2640           9.1-22.7       8 *
Apiaceae                  830-3395           12.2-34.5      8 *

Apocynaceae              1420-3070           9.2-30.4       8 *
Asteraceae                770-3285           5.2-28. 0      8 *
Asteraceae                  2190                19          8 *
Asteraceae               2880-4540           9.7-18.1       8 *
Asteraceae               2400-3040           10.7-18.2      8 *
Asteraceae               1880-3100           9.1-17.1       8 *
Asteraceae               1605-2140           16.7-27.7      8 *
Asteraceae                980-4110           15.6-23.8      8 *
Asteraceae                650-2600           12.8-34.4      8 *
Asteraceae               1980-3270           10.4-32.5      8 *
Asteraceae               2325-4470            5.3-8.1       8 *
Berberidaceae            1400-2570           9.1-30.7       8 *
Brassicaceae             3350-4400           11.8-16.6      8 *
Brassicaceae             3460-5000           9.6-18.6       8 *
Brassicaceae             3040-3615           12.2-14.1      8 *

Bromeliaceae             2670-3720           11.0-14.8      8 *
Bromeliaceae             1985-2740            6.1-6.2       8 *
Cactaceae                1290-3160           7.5-17.2       8 *
Cactaceae                2000-3000         11.48-20.72      8 *
Caryophyllaceae          1860-4350           6.6-16.6       8 *
Convolvulaceae            750-2320           10.3-31.7      8 *
Cucurbitaceae             350-2610           11.5-28.1      8 *
Cyperaceae                  3280               16.6         8 *
Fabaceae                 1670-3220           8.5-17.0       8 *
Fabaceae                 3110-4000           5.9-16.6       8 *
Fabaceae                 2030-4090           13.6-17.4      8 *
Gentianaceae             2600-3940           11.0-18.1      8 *

Gentianaceae             3395-4580           10.4-17.8      8 *
Geraniaceae              1660-3180           12.2-26.4      8 *
Iridaceae                 820-3240           9.1-29.0       8 *
Martyniaceae              400-2130           6.3-28.0       8 *
Montiaceae               3260-3860           7.0-14.1       8 *
Poaceae                  1800-3940           12.2-25.9      8 *
Poaceae                  3400-4160           8.9-13.3       8 *
Poaceae                  2270-4140           5.3-19.6       8 *
Poaceae                  2270-3460           10.7-18.1      8 *
Poaceae                  1710-1840           15.2-25.5      8 *
Poaceae                  1900-3690           6.7-15.5       8 *
Polygonaceae             2300-3460           11.0-18.1      8 *
Rosaceae                 2460-3400           10.1-15.9      8 *
Rubiaceae                1490-2430           12.8-19.6      8 *
Solanaceae                470-3460           9.2-24.7       8 *
Solanaceae               2270-3100           9.7-17.0       8 *
Urticaceae               1130-1840           15.2-28.4      8 *
Valerianaceae             730-3410           7.3-19.2       8 *
Verbenaceae               295-3100           6.8-27.9       8 *
Verbenaceae               330-2270           9.9-26.6       8 *
Verbenaceae              1110-2930           7.5-18.7       8 *
Violaceae                2490-3510           11.0-17.0      8 *
Amaryllidaceae            770-4810           5.8-27.1       9
Asteraceae               1815-3195           6.2-11.9       9
Asteraceae               1410-2230           6.4-10.2       9
Asteraceae               2240-3830           6.2-12.9       9
Asteraceae               1660-3680           4.8-11.0       9
Asteraceae                840-2665           5.1-12.9       9
Asteraceae               1030-2780           4.6-10.5       9
Asteraceae               1110-2100           8.8-12.0       9
Cactaceae                1230-2770           4.1-16.7       9
Cactaceae                2100-3000          8.81-18.58      9
Cactaceae                 620-1960           4.4-10.9       9

Fabaceae                 2180-3510            5.8-8.5       9
Fabaceae                 3590-4360           5.6-10.1       9
Fabaceae                 1070-2230            4.2-8.1       9
Sapindaceae              1460-2650            3.1-9.3       9
Solanaceae                510-2765           3.8-14.8       9
Solanaceae               1030-2780           4.6-10.5       9
Amaryllidaceae           1740-3620           8.5-25.2       10
Anacampserotaceae        3200-3800          6.66-13.55      10
Apiaceae                 3180-4810           6.2-14.8       10
Asteraceae               1310-3940           9.1-24.2       10
Asteraceae               1470-3200           11.9-28.1      10
Asteraceae               2830-4680           5.0-18.1       10
Asteraceae               2340-4320           7.3-18.1       10

Asteraceae               2420-3450           6.5-15.6       10
Asteraceae               1500-4150           9.1-26.1       10
Asteraceae                567-2530           4.2-14.6       10
Asteraceae               2750-4110           6.9-18.5       10
Asteraceae               2770-4150           5.1-14.8       10
Asteraceae               1250-3265           7.9-12.2       10
Asteraceae               2505-3830           8.4-13.0       10
Asteraceae               1190-3510           7.4-19.0       10
Asteraceae               2250-3460           9.8-18.1       10
Asteraceae                810-2550            5.5-9.1       10
Brassicaceae             3000-4470           7.3-18.6       10
Brassicaceae             1140-3510           7.1-16.1       10
Bromeliaceae             1170-2010           5.0-22.0       10
Cactaceae                 560-2050            4.1-9.2       10
Cactaceae                 600-2720           5.9-28.0       10

Caryophyllaceae          3105-4610           5.1-19.1       10
Euphorbiaceae             600-3500          8.25-13.13      10
Ledocarpaceae            1530-4375           4.8-21.2       10
Loasaceae                2245-3960           6.3-16.5       10
Nyctaginaceae            1580-4390           6.7-22-6       10
Poaceae                  1850-4350           6.9-22.1       10
Poaceae                  1070-3680           7.0-20.6       10
Poaceae                  2270-4640           7.5-18.1       10
Poaceae                  1065-3500           5.2-11.3       10
Poaceae                   910-3180           3.1-29.2       10
Poaceae                  1220-2190           8.3-28.6       10
Poaceae                  3500-4600           6.7-16.9       10

Poaceae                  1780-2950            5,3-8,6       10
Polygalaceae             1410-3195           5,5-23,4       10
Polygalaceae             1920-4240           5,1-15,4       10
Solanaceae               1730-4310           5,6-15,9       10
Solanaceae                940-3520           3,3 -23,7      10
Solanaceae               1140-3200           5,3-12,9       10
Solanaceae               1150-2900           4,3-16,7       10
Solanaceae               1620-2300            4,3-6,8       10
Solanaceae                910-3290           4,3-15,3       10
Solanaceae               1840-4390           6,5-11,8       10
Verbenaceae               670-2460           4,0-10,6       10
Violaceae                3040-3195           10,9-15,1      10
Zygophyllaceae           1060-3265           4,1-19,9       10
Zygophyllaceae            970-2710            4,0-9,6       10

Cactaceae                 610-2600            6,3-29        10
Fabaceae                 1530-4100           4,7-17,0      10 *
Fabaceae                 1750-3500           4,9-11.9      10 *
Onagraceae               2580-4070           9,2-17,4      10 *
Oxalidaceae              1670-4610           4,2-19,1      10 *
Poaceae                  2102-4170           5,1-17,0      10 *
Amaryllidaceae           2260-3230            5,9-9,2       11
Apocynaceae                 1880                9,1         11
Asteraceae               2900-3015            8,2-9,0       11
Asteraceae                  1720               10,6         11
Boraginaceae             1540-1965            6,3-9,9       11
Bromeliaceae             2110-2930            6,2-8,4       11
Bromeliaceae             1650-3510           5,9-10,5       11
Bromeliaceae                1160               14,1         11
Cactaceae                1640-3440            5,4-9,0       11
Cactaceae                 810-1650           19,0-19,9      11
Cactaceae                3000-4200           6,1-11,3       11
Cactaceae                2610-3230            5,8-9,2       11
Cactaceae                   1650                7,0         11
Cactaceae                1480-2190            6,7-9,3       11
Cactaceae                1640-1740            7,8-8,3       11
Fabaceae                 2220-2480            6,8-8,4       11

Geraniaceae              2730-2820           9,7-12,9       11
Malvaceae                   4150               11,8         11
Solanaceae                  1705                9,6         11
Solanaceae               1590-2820           6,9-10,2       11
Asteraceae               3260-3565              8,4        11 *
Bromeliaceae                2480                6,4        11 *
Cactaceae                3260-3950            8,1-8,7      11 *
Cactaceae                   2240               11,5        11 *
Cactaceae                3230-3540            9,2-9,5      11 *
Dioscoreaceae            1000-1820           18,7-36,6     11 *
Euphorbiaceae             740-1290           12,5-21,7     11 *
Orchidaceae                 800                22,1        11 *
Poaceae                     3540                9,5        11 *
Asteraceae               1720-3130            5,8-9,7       12
Asteraceae               2300-3370            4,3-8,4       12
Cactaceae                1900-3540            5,7-9,0       12
Gentianaceae             2580-3230           9,2-15,6       12
Malvaceae                1640-2900           5,5-10,8       12
Oxalidaceae              4370-5150           11,4-19,1      12
Poaceae                  1810-4645           5,5-18,1       12
Solanaceae               1510-2900           4,3-11,4       12
Amaranthaceae            2160-2330            5,1-6,6       13
Amaranthaceae             580-1590            3,6-5,6       13
Asteraceae               2510-3250           9,7-15,6       13
Asteraceae               2890-3430           11,7-12,2      13
Boraginaceae              820-1070            4,3-4,7       13
Brassicaceae                2940               15,1         13

Cactaceae                 660-920             3,7-4,4       13
Cactaceae                 600-1330            3,8-9,0       13
Cactaceae                   600                 4,5         13
Fabaceae                 2180-2590            8,4-8,5       13
Violaceae                   2960               15,6         13
Asteraceae                  3410                7,3         14
Asteraceae               4090-4240            5,5-5,7       14
Asteraceae                  2600                3,1         14
Cactaceae                   2150                4,3         14
Cactaceae                1645-3070            4,1-5,3       14
Calceolariaceae             1810                5,5         14
Caryophyllaceae             3410                7,3         14
Malvaceae                3140-3690            3,3-5,5       14
Malvaceae                   4240                5,5         14
Malvaceae                   3740                7,2         14
Polemoniaceae            2600-2900            3,1-3,3       14
Solanaceae               2730-3420            5,6-7,0       14
Amaranthaceae            1790-1800            7,9-9,0       15
Apiaceae                 3040-3615           12,2-14,1      15
Apiaceae                    3040               12,2         15
Apocynaceae                 1955                9,6         15
Apocynaceae              1420-1820           15,7-22,0      15

Apocynaceae               660-975            22,1-28,8      15
Aristolochiaceae         1610-2130           6,3-18,6       15
Asteraceae                  2860                7,6         15
Asteraceae                  2885               10,7         15
Asteraceae                  3250                8,3         15
Asteraceae               2570-3040           11.4-13.2      15
Asteraceae               1740-3040           12.2-15.0      15
Asteraceae               2930-3800           11.4-14.7      15

Asteraceae                  3250                8.3         15
Asteraceae                810-1070           15.7-19.9      15
Asteraceae                  1300               34.4         15
Asteraceae               3720-4375           12.3-17.8      15
Asteraceae                  4560               13.1         15
Asteraceae               4460-4470           12.9-18.6      15
Asteraceae               1650-4375           12.4-17.8      15
Asteraceae               3360-3710           8.4-14.9       15
Asteraceae               3510-4000           9.3-14.6       15
Asteraceae               1820-3810           12.2-24.5      15
Begoniaceae              2100-3540           11.0-15.8      15
Cactaceae                2320-3110            8.3-8.6       15
Cactaceae                 890-1490           15.2-28.4      15
Cactaceae                2100-3180           8.6-15.8       15
Cactaceae                   1970               18.2         15
Cactaceae                1860-2170           9.3-19.3       15

Caryophyllaceae             3210               10.1         15
Cyperaceae                  3040               12.2         15
Dioscoreaceae             530-1740           24.4-31.8      15
Fabaceae                  930-2030           10.5-19.6      15
Hypoxidaceae                1740                15          15
Iridaceae                   3040               12.2         15
Iridaceae                 560-2560           12.4-34.4      15
Loasaceae                1710-3040           12.2-15.7      15
Malvaceae                3460-4440           12.4-18.1      15
Malvaceae                4115-4460           10.7-12.9      15
Malvaceae                4470-4760           18.6-21.3      15
Malvaceae                4420-4760           17.8-21.3      15
Malvaceae                   4375               17.8         15
Montiaceae                  2445               10.4         15
Onagraceae               830 -2500           9.7-34.5       15
Orchidaceae              2110-3045           6.9-12.2       15
Orchidaceae              3230-3640           9.1-11.2       15
Plantaginaceae           1260-3040           12.2-19.7      15
Poaceae                   990-3180           12.2-32.4      15
Poaceae                  1880-2940           9.1-11.3       15
Ranunculaceae            1660-1840           15.0-26.4      15
Rosaceae                    2700               13.3         15
Solanaceae                940-1960           9.6-34.0       15
Solanaceae               1510-1600           15.7-29.2      15
Solanaceae               1280-1940           14.2-34.4      15
Solanaceae                690-1960           15.4-25.3      15
Solanaceae               2840-3220           9.1-12.5       15
Valerianaceae            3210-3720           10.1-14.7      15
Violaceae                   4375               17.8         15
Violaceae                4190-4515           11.8-12.7      15
Violaceae                   4300               17.3         15
Violaceae                3000-4560           10.5-17.8      15
Violaceae                3400-3480           12.2-13.3      15
Acanthaceae               560-1080           9.0-16.0       16
Cactaceae                 400-1100           8.9-15.9       16
Fabaceae                  800-1275           6.4-13.7       16
Poaceae                   310-750            12.6-15.7      16

Amaryllidaceae              3460               15.7         17
Asteraceae                  1800               24.3         17
Asteraceae               3220-3940           16.8-17.9      17
Asteraceae                  3685               16.5         17
Asteraceae               1660-1850           23.4-25.1      17
Bromeliaceae             3280-3940           16.6-16.8      17
Cactaceae                   3480               17.4         17
Cactaceae                   3610               15.8         17
Cactaceae                   3940               16.8         17
Gentianaceae                2990               18.1         17
Iridaceae                   3280               16.6         17
Iridaceae                   3540               15.5         17
Malvaceae                4150-4410           15.9-16.5      17
Poaceae                     3680               15.6         17
Poaceae                     3540               15.5         17
Solanaceae                  3520                17          17
Tropaeolaceae            2610-3030           17.1-18.4      17
Amaryllidaceae              280                14.5         ?
Asteraceae               2490-4620           6.8-17.4       ?
Asteraceae               1310-2590           16.0-35.0      ?
Asteraceae                560-1720           22.4-31.7      ?
Asteraceae               410 -2420           6.5-15.0       ?
Asteraceae               2940-4250           5.2-18.6       ?
Brassicaceae             3480-4820           7.6-15.9       ?
Cactaceae                   180                15.3         ?
Cyperaceae               3705-4030           4.4-10.6       ?
Fabaceae                 3040-4375           3.4-21.3       ?
Fabaceae                 2960-4375           7.8-17.8       ?
Gentianaceae                4250                4.3         ?
Poaceae                   830-4090           12.2-34.5      ?
Poaceae                  1000-2870           6.1-36.6       ?
Poaceae                  2815-4210           5.1-12.9       ?
Poaceae                  1050-4100           6.9-19.0       ?
Poaceae                  2410-3705           6.2-12.8       ?
Poaceae                  2470-4480            2.8-8.9       ?
Solanaceae               3510-4680           5.7-10.4       ?
Solanaceae               2310-3685           7.7-16.5       ?
Violaceae                4070-4170           3.9-14.8       ?

Table 2b. Code of area names used in Table 2a.

Area                             Area                            Fig.
number

1        Ambato                                                   4M
2        Andes La Rioja-San Juan                                  4F
3        South Bolivia-north NOA                                  4I
4        Bolivian Prepuna                                         4J
5        Catamarca and La Rioja, summits and dry inner valleys    4P
6        Famatina                                                 4H
6 *      Famatina, border                                         4H
7        Jujuy, core                                              4C
7 *      Jujuy, border                                            4C
7 **     Jujuy, core and border                                   4C
8        Jujuy-Tucuman, core                                      4E
8 *      Jujuy-Tucuman                                            4E
9        La Rioja and San Juan, summits and dry inner valleys     4Q
10       NOA                                                      4B
10 *     NOA (San Juan)                                           4B
11       Salta, core                                              4L
11 *     Salta                                                    4L
12       Salta-Catamarca, summits and dry inner valleys           4O
13       San Juan                                                 4G
14       Tinogasta--Belen                                         4N
15       Tucuman                                                  4D
16       Valle Fertil                                             4R
17       Yavi-Santa Victoria                                      4K

Table 3. Areas separating from main consensus in Fig. 4, ordered
according to max. area score (grid size 0.5[degrees] x 0.5[degrees]).
The areas were obtained by gradually increasing consensus criterion
from 5% to 50% (ae and aa refer to different consensus rules).

Area                             Obtained        Nr. Of        Area
                                  under        sub-sets in     score
                                                consensus
                               0.2[degrees]
                              x 0.2[degrees]

Jujuy-core and border              yes             101       2.1-26.3
Tucuman                            yes             17        7.3-23.3
Salta                              yes             36        2.7-13.3
Famatina                           yes              2        10.9-13.1
Famatina border                                     1         3.9-4.2
Yavi-Santa Victoira                yes              2        10.6-10.8
Jujuy-Tucuman                      yes             370       2.0-10.4
Ambato                             yes             24        2.4-10.3
Bolivian Prepuna                   yes             19         2.2-6.5
Tinogasta-Belen                    yes              9         2.6-4.5
San Juan                            no              8         2.0-3.8
Catamarca-La Rioja summits          no             19         2.0-3.8
  and dry inner valleys
NOA                                 no             14         2.0-3.5
NOA-San Juan                                        2         2.1-2.4
South Bolivia-north NOA             no              1         3.0-3.2
Andes La Rioja-San Juan             no              6         2.1-2.8
Salta-Catamarca                     no              4         2.1-2.8
  summits and dry inner
  valleys

Area                            Nr. of     Altitude     Aridity
                              Families/      range       range
                               Genera/
                               Species

Jujuy-core and border          31/66/87    350-5100    6.2-33.7
Tucuman                        22/38/55    530-4800    6.3-34.5
Salta                          14/23/27    740-4200    5.4-36.6
Famatina                        43687      1150-4100   4.5-15.0
Famatina border                 38750      750-2200    7.2-11.8
Yavi-Santa Victoira             41498      1700-4400   15.5-25.1
Jujuy-Tucuman                  38/59/85    350-5000    5.2-40.4
Ambato                          44879      1000-3700   6.6-18.1
Bolivian Prepuna                44516      1400-4400   6.7-26.3
Tinogasta-Belen                 40732      1600-4300    3.1-7.3
San Juan                        40369      850-3400    3.6-15.6
Catamarca-La Rioja summits      43813      500-4500    4.8-18.6
  and dry inner valleys
NOA                             44123      560-4800    3.1-29.2
NOA-San Juan                    38081      150-4600    4.7-19.1
South Bolivia-north NOA         37350      2000-4000   6.6-19.6
Andes La Rioja-San Juan         40765      1000-4500   3.8-24.1
Salta-Catamarca                 39636      1500-5100   4.3-19.1
  summits and dry inner
  valleys

Area                          Fig.         Separates
                                           from main
                                             area

Jujuy-core and border          4C           40% aa
Tucuman                        4D           45% aa
Salta                          4L           50% aa
Famatina                       4H           25% aa
Famatina border                             25% aa
Yavi-Santa Victoira            4K           25% aa
Jujuy-Tucuman                  4E           50% aa
Ambato                         4M           45% aa
Bolivian Prepuna               4J           25% aa
Tinogasta-Belen                4N           45% aa
San Juan                       4G    not part of main area
Catamarca-La Rioja summits     4P           30% aa
  and dry inner valleys
NOA                            4B    not part of main area
NOA-San Juan                         not part of main area
South Bolivia-north NOA        4I    not part of main area
Andes La Rioja-San Juan        4F    not part of main area
Salta-Catamarca                4O           45% aa
  summits and dry inner
  valleys

Table 4. Main radiations within the study region. Only families with
10 or more endemic species, and genera with five or more species have
been included. For the families, the percentage of the total number of
endemic species within the region is shown.

Main families: nr. of    Main genera: nr. of
   endemic species         endemic species

Asteraceae: 126 (23%)    Senecio: 39
                         Hieracium: 10
                         Flourensia: 9
                         Baccharis: 8
                         Stevia: 6

Cactaceae: 75 (14%)      Lobivia: 16
                         Gymnocalycium: 15
                         Trichocereus: 13
                         Parodia: 7
                         Tephrocactus: 6

Poaceae: 46 (8%)         Nassella: 12
                         Poa: 6

Fabaceae: 34 (6%)        Lupinus: 10
                         Adesmia: 8
                         Astragalus: 8

Solanaceae: 29 (5%)      Solanum: 18
                         Sclerophylax: 6

Malvaceae: 25 (5%)       Nototriche:19
Bromeliaceae: 17 (3%)    Puya: 9
Apocynaceae: 15 (3%)     Philibertia: 8
Violaceae: 12 (2%)       Viola: 12
Gentianaceae: 11 (2%)    Gentianella: 11
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Title Annotation:ecologia y fitogeografia; texto en ingles
Author:Aagesen, Lone; Bena, Maria J.; Nomdedeu, Soledad; Panizza, Adela; Lopez, Ramiro P.; Zuloaga, Fernand
Publication:Darwiniana
Date:Dec 1, 2012
Words:21825
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