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Biodiversity and Conservation - Supplementary material

A framework for the classification Chilean terrestrial

ecosystems as a tool for achieving global conservation targets

Karina Martínez-Tilleria1, 3, Mariela Núñez-Ávila2, *, Carolina A. León2, 5, Patricio

Pliscoff 3, Francisco A. Squeo1, 4, Juan J. Armesto2, 3, 6

1 Departamento de Biología, Facultad de Ciencias, Universidad de La Serena,

Raúl Bitrán 1305, La Serena, Chile.

2 Instituto Milenio de Ecología y Biodiversidad (IEB), Universidad de Chile,

Santiago de Chile.

3 Departamento de Ecología, Facultad de Ciencias Biológicas, Pontificia

Universidad Católica de Chile, Casilla 114-D, Santiago, Chile.

4 Centro de Estudios Avanzados en Zonas Áridas (CEAZA), Raúl Bitrán 1305,

La Serena, Chile.

5 Universidad Bernardo O Higgins, Centro de Investigación en Recursos

Naturales y Sustentabilidad, Fábrica 1990, segundo piso, Santiago, Chile.

6 Cary Institute of Ecosystem Studies, Millbrook, New York, U. S. A.

Author for correspondence*: [email protected], phone: +56996566047.

Supplementary material 1


Ecosystem descriptions

1. HIGH-ANDEAN STEPPE: Extensive steppes dominated by tussock grasses and cespitose herbs, with sporadic presence of shrubs, are found above 3000 m in the Andean Highlands of the two northernmost regions of the country (Fig. 1). Biota: Among the dominant tussock grasses the following species are frequent: Festuca orthophylla, Deschampsia cespitosa, Deyeuxia breviaristata, Deyeuxia antoniana, Deyeuxia curvula (Arroyo et al. 1992). In areas with saline soils, species such as Distichlis humilis, Distichlis spicata, Stipa chrysophylla become predominant. Characteristic shrub species in this ecosystem is Parastrephia lucida. Rocky outcrops are often occupied by long-lived cushion plants, primarily Azorella compacta, and provide suitable habitats for populations of the herbivorous southern viscacha (Lagidium viscacia). Important herbivores of the high-Andean steppe are three species of camelids. A threated species from the high-Andean steppe is the rare Andean cat, Leopardus jacobita (CONAMA 2008). Abiotic parameters: The altitudinal distribution of high Andean steppe has been affected by millennial precipitation cycles related to Glacial Cycles (Latorre et al. 2003). In wetter periods, the steppe has expanded downslope into the drier desert. Moisture availability is strongly dependent on storm activity associated with the Inter-Tropical Convergence Zone (ITCZ), which during the summer extends its influence to subtropical latitudes. The resulting climate pattern is characterized by warm summer rains (December-March, “invierno boliviano”) followed by cold, drier austral winters. At 18o S, average precipitation above 4,000 m elevation is close to 400 mm per year (Arroyo et al. 1988). Despite the limited amount of rainfall, the high-Andean steppe commonly exhibits two flowering seasons, October-December and February-April, when mean temperatures are between 10 and 11 C. At 4500 m, flowering occurs with mean temperatures of 4 to 6 C (Arroyo et al. 1987). Functional ecosystem model: The steppe ecosystem supports a rigorous climate, with low partial pressure of oxygen and carbon dioxide, high solar radiation and extreme seasonal and diurnal temperature fluctuations (Marquet et al. 1998). Many animal species exhibit hibernation periods. For the plants, productivity is limited by periodical droughts and soils with low nitrogen supply, and intermediate to high phosphorous and potassium availabilities. The grazing pressure of camelids (domestic and wild) and other mammalian herbivores can be very high, which has resulted in vegetation dominated by shrubs and grasses with tough, resistant foliage. Current threats: A striking singularity within the realm of the steppe in the Andean highlands is the presence of a small tree (5-10 m tall) Polylepis tarapacana (queñoa), forming patchy woodlands at elevations above 4,000 meters. These woodlands, which are shared with neighboring high-Andean zones of Bolivia and Peru, are considered one of the most threatened ecosystems in the world because of their narrow range, topographic location, and past extraction of firewood (Renison et al.



2004). Sparse Polylepis woodlands are unique components of the predominantly treeless high-Andean steppe. Temperatures below freezing during the nights and diurnal fluctuations of 20-30 C are conditions often restrictive for the development of trees (Kessler 2006). According to Jaksic et al. (1997), Polylepis woodlands harbor a wide diversity of Andean birds, many of them endemic. The extreme climate of the high Andean steppe and the poor soils, together with the susceptibility to long-lasting dry and wet cycles and high biomass loads of grazing animals, make this ecosystem extremely fragile in the face of natural or human disturbances (Jaksic et al. 1997). The extraction of firewood from Polylepis and longevous cushion plants (genus Azorella) for mining operations in the past produced long-lasting impacts in the distribution of species, due to their low regeneration potential and slow growth. Distribution: They are exclusively found in the highlands of the northern Andes, from 15 to approximately 26o S, with an extensive area of 21,038 km2, equivalent to 3% of the country.

2. CENTRAL-CHILEAN ANDEAN STEPPE: This is the mountain steppe of the Andes of central Chile, with a narrower distribution, under a Mediterranean-climate (30-34o S). It has a mixture of shrubby and herbaceous cover, without trees; the steppe ecosystem of central Chile has precipitation concentrated in the austral winter months (June to August) in the form of snow, with a generally long dry summer, interrupted by occasional summer storms (Arroyo et al. 1981). Biota: Among the dominant shrubs of the central Chilean steppe are Chuquiraga oppositofolia, Anarthrophyllum cummingii, Berberis empetrifolia; the most conspicuous grasses are Stipa frigida and Stipa rupestris, and the cushion plants that occur above 2700 m are Azorella monantha and Laretia acaulis. The rich herbaceous flora of this ecosystem includes annuals (e.g., the genus Viola) and perennials (e.g., Adesmia, Astragalus, Nassauvia), and some endemic geophytes (e.g., the genus Placea). Abiotic parameters: The major difference with the Andean highlands steppes is that the rain here is determined by storm fronts from the west, progressing inland and northward during the austral winter period. Continuous snow cover lasts for 3-5 months while the summers are extremely dry (Arroyo et al. 1981). Winter snow restricts the length of the growing season in the central Andean steppe; consequently, the flowering season begins as soon as snow melts, with a peak in the warmest months (December-January). Mean temperature during the peak of the growing season can be as high as 18o C, wile the winter months have a mean of 5o C (Arroyo et al. 1981). Functional ecosystem model: Cavieres et al. (2000) documented the pattern of altitudinal zonation in vegetation structure and composition through the steppe ecosystem, showing that woody plants and annuals are restricted to hillsides at lower elevations (2300-2500 m), while cushion plants and grasses tend to predominate above 2500 m. Mean annual temperature and low soil nitrogen (N) contents appear to be determining the altitudinal zones in the shrubland-steppe transition. Cushion plants have been shown to facilitate the



establishment of various herbaceous species in the upper elevations (Cavieres et al. 2002). Arroyo et al. (1983) described the pollination systems in the steppe-shrubland transition and the high Andean steppe, showing that anemophily becomes increasingly important in the subtropical highland steppe, but it declines in favor of insect pollination (melittophily and psychophily) in the high Andes of central Chile, to increase again in the Patagonian steppe. Current threats: Present impacts derive primarily from cattle and horse grazing, especially in wet areas (“vegas”), extraction of firewood, and the expansion of housing developments and hotels around ski fields. The ski fields themselves are subjected to severe disturbance by the use of heavy machinery to maintain areas devoid of woody vegetation. Access roads have negative effects on the native bird, reptile, and mammal fauna, promote slope erosion, landslides, and provide opportunities for exotic species invasion. Such activities may be particularly disruptive for populations of endemic geophytes. Distribution: Andean steppe ecosystems are found above the treeline, from 2500 m up to the limit of vegetation, between the latitudes 30 and 33o S (Arroyo et al. 1987). They cover 3986 km2, 0.55% of the country.

3. PATAGONIAN STEPPE: These are ecosystems dominated by a sparse cover of shrubs, and abundant perennial herbs and tussock grasses. Its most distinctive climatic feature is the low precipitation, distributed all-year long. They are often considered part of the arid diagonal of southern South America and have floristic affinities with northern semiarid ecosystems (Armesto et al. 2007). Biota: Among the most frequent perennials are Festuca gracillima, Mulinum spinosum, Poa scaberula. An abundant herbivorous fauna is again present, including populations of guanaco (Lama guanicoe) that are the prey of large predators, such as the puma. The native herbivore fauna has been enriched by the introduction of rabbits and hares, all of which have impacts on post-disturbance regeneration. Abiotic parameters: Precipitation in the Patagonian steppe is generally lower than 300 mm per year, with constant and strong winds, causing high evaporation from soil and plant surfaces. Conditions are harsh for the biota, indicated by the short flowering period (December-February) restricted to the warmer months (Arroyo et al. 1987). Functional ecosystem model: Moisture availability is more limited than in the other steppe ecosystems, affecting productivity. The onset of the growing season is primarily related to warming temperatures, while the end is related to lower soil moisture availability (Aguiar et al. 1996). Dominant growth forms in the steppe are tussock grasses, cushion plants and perennial herbs with variable abundances (Soriano and Sala 1983). Productivity is mostly due to grasses and shrubs, despite being less abundant than herbs. Grasses and shrubs have different strategies to face limited water availability; while the former use water near the soil surface, shrubs concentrate their root systems in the deeper soil levels (Aguiar et al. 1996). Current threats: The main threat to the Patagonian steppe ecosystem is the expansion of exotic grasses sown to create pastures for



sheep. Recently, increasing drought frequency, combined with human-set fire, have favored the abundance of exotic grasses, excluding shrub cover (León & Aguiar 1985; Aguiar et al. 1996). Such changes modify productivity and carbon storage patterns, also altering soil nutrient and water storage capabilities (Aguiar et al. 1996). It seems quite likely that during the long history of sheep ranching, the original structure and functions of Patagonian steppe ecosystems have been changed, leading to the replacement of native grasses by exotic pasture species. Distribution: Between Aysen and Magellanes Regions (Fig. 1), Patagonian steppe covers an area of 19,749 km2, equivalent to 2.75% of mainland Chile.

4. ECOSYSTEMS DOMINATED BY SUCCULENTS: This ecosystem is represented cartographically by the unit ‘formation with succulent cover’ distinguished in the Chilean terrestrial vegetation survey (CONAF et al.1999). Such ecosystems are characterized by the dominance of succulent plants (particularly with CAM photosynthesis) and leaves replaced by spines, particularly columnar and spherical cactuses and large terrestrial bromeliads (e.g., genus Puya), with a minor biomass component of shrubs and seasonal herbaceous plants. Biota: The main cactus species in these particular systems are Browningia candelaris, Copiapoa cinerascens, Copiapoa cinerea, Copiapoa dealbata, Copiapoa eremophila, Copiapoa gigantea, Corryocactus brevistylus, Echinopsis chiloensis, Eulychnia aricensis, Eulychnia iquiquensis, Eulychnia morromorenoensi, Eulychnia saint-pieana. On rocky substrates dominance can be shared with various species of Puya (Bromeliaceae). Copiapoa is an important endemic genus with about 30 species along the coast of northern Chile, in the regions of Antofagasta and Atacama (Fig. 1). Abiotic parameters: This type of ecosystem occupies arid and semiarid zones, on coastal or Andean foothills, where shrubs are sparse or rare due the extremely low rainfall (50 mm or less per year), and only annuals with persistent seed banks are able to survive. Soils are permanently exposed to wind erosion and organic matter is lacking. Functional ecosystem model: These ecosystems harbor a significant number of endemic animal species; particularly many insects feed on the pollen and nectar of large cactus flowers and help support a rich food chain (Jaksic and Fuentes 1980; Saiz and Campalans 1984; Solervicens et al. 2004). Spines protect water storage organs from herbivory, and CAM photosynthetic pathway allows plants to survive under extreme drought in the lowland Atacama Desert. On some coastal locations (e.g., Paposo), columnar Cactaceae appear associated with oceanic fog zones above 400 m elevation (Rundel and Mahu 1976). Spines and epiphytes (mainly fruticose lichens) associated with cactus spines ‘harvest’ water from fog (Stanton et al. 2014) and moderate temperature fluctuations and daytime water pressure deficit for the host plants. Populations of thousands of Cactaceae in extremely arid environments of the Atacama Desert reproduce only sporadically and may represent relicts of past wetter periods. Current threats: Many succulent plants have medicinal properties,



produce fruits or materials useful to people, or have commercial values (Belmonte et al., 1998; Pardo 2002). For this reason some populations have been greatly reduced in recent times. The loss of a large columnar cactus individual from one location in these ecosystems can be a serious problem for many years because of the low recruitment and slow growth. Distribution: Ecosystems dominated by succulent plants are primarily abundant between the regions of Tarapacá and Coquimbo (Fig. 1), from 18 to 30o S. Their areal extent is about 1810 km2, i.e. 0.25% of mainland Chile.

5. EVERGREEN SCRUB WITH MICROPHYLLOUS LEAVES: Dominant shrub species have characteristically small (<2 cm long) evergreen leaves that remain on the plant during unfavorable drought periods. In some instances, the leaves are reduced to scales on a photosynthetic stem. Biota: Important genera, with many species in this ecosystem, are Adesmia, Baccharis, Berberis, Ephedra, Fabiana, Haplopappus, Heliotropium, Nolana, Parastrephia, Tetragonia. Frequently, shrub cover is associated with a variable carpet of annuals, often appearing after the rains. Seed and herb eating rodents, such as the endemic and fossorial Spalacopus cyanus, are frequently found here, especially on sandy soils (Díaz I. et al. 2002). Abiotic parameters: This type of scrub occurs on relatively nutrient-poor soils due to the slow nutrient cycling associated with semiarid climate. Shrubs are distributed at different elevations in a broad latitudinal range, where annual precipitation is lower than 300 mm. Sandy soils are common and subjected to loss of nutrients due to surface runoff after large rain events. Functional ecosystem model: Areas dominated by microphyllous (small-leaved) shrubs represent a significant portion of what has been previously classified as desert scrub at lower latitudes, along with the Andean-Patagonian scrub in the drier slopes at higher latitudes. In both cases, limiting environmental conditions for plants and animals are related to permanent or frequent drought. Reduced leaf area regulates water loss by transpiration when soils are dry. Shrub cover (30-50%) seem to provide nursing conditions for annual and perennial herbs (Gutiérrez et al. 1993; Aguilera et al., 1999; Gutiérrez and Squeo 2004), increasing soil moisture through hydraulic lift from deep roots (Squeo et al. 1999; León and Squeo 2004; León et al. 2011). This physiological process has positive consequences at the ecosystem level by enhancing carbon assimilation, fine root growth and nutrient absorption by plants and microorganisms. Shrub cover also contributes to reduce water runoff and reduce the potential for soil and nutrient erosion (Gutiérrez and Squeo 2004). Current threats: Historically, these ecosystems have severely changed by human impact, because of the use of shrubs, such as Fabiana imbricata or Adesmia hystrix, for firewood and charcoal production (Estevez et al. 2010). Two centuries of widespread mining operations led to the reduction of woody species cover, firewood extraction for use in hundreds of smelting plants (Armesto et al. 2010). Such impacts, added to the permanent grazing pressure of cattle and goats, in an environment subjected to intense periodic droughts,



caused grave losses biodiversity and reduced ecosystem resilience (Holmgren et al. 2001). Distribution: The microphyllous scrub ecosystem is found in areas subjected to strong seasonal drought, particularly above 3000 m in the northern Andes, but at lower elevations in the semiarid region, at the southern boundary of the Atacama Desert, and along the margins of the Patagonian steppe. These are highly heterogeneous ecosystems in terms of floristic composition and ground cover distribution and much work is needed to provide cartographic units with finer resolution. Their land cover is 88,912 km2, equivalent to 12.4% of the Chilean mainland.

6. EVERGREEN SCRUB WITH MACROPHYLLOUS LEAVES: Dominant species in this ecosystem, also known as Chilean matorral, are shrubs with persistent foliage and leaves > 2 cm long (macrophyllous), characterized by leathery texture and structurally tough to decompose (sclerophyllous). Because of the presence of structural tissue, leaves typically have high leaf mass per area (high LMA) compared to other ecosystems. Dominant growth forms are tall shrubs (1-5 m) occurring on top of a dense cover of annuals and perennial herbs in the rainy season. Biota: A high diversity of shrub species includes the following: Azara dentata, Azara microphylla, Cestrum parqui, Colliguaja dombeyana, Colliguaja odorifera, Colliguaja salicifolia, Escallonia alpina, Escallonia angustifolia, Escallonia revoluta, Kageneckia oblonga, Kageneckia angustifolia, Lithrea caustica, Schinus latifolius, Schinus montanus, Schinus polygamus. In the southern limit of the sclerophyllous scrub, some small trees such as Lomatia hirsuta and L. dentata are also present. For a complete floristic list, see Villagrán et al. (2007). Comparative studies of the flora and fauna of Chilean sclerophyllous scrub, in Chile, California and other Mediterranean-climate regions, can be found in the literature (e.g., Mooney 1977; Rundel 1981; Jaksic 1997; Arroyo et al. 1995). Abiotic parameters: This ecosystem occupies the Mediterranean-climate region of southern South America, with rains restricted to the austral winter period (May-August), but varying greatly with orography (elevation and distance from the ocean) as well as latitude, ranging between 300 and 800 mm per year. The Chilean matorral develops on the coastal and Andean ranges, and in sectors of the central valley where agriculture is absent. Soils vary from highly weathered metamorphic substrates on coastal mountains to more recently derived igneous rocks and rocky conglomerates in the Andean foothills. Consequently, a strong biotic heterogeneity has been described for the sclerophyllous scrub ecosystem across its geographic range (Rundel 1981; Armesto et al. 2007). The distribution and amount of rain varies greatly among years, modulated by the effects of ENSO (Aceituno 1988; Holmgren et al. 2001). Ecosystem production and animal activity are strongly dependent on this climatic variability. Functional ecosystem model: The mosaic of communities represented in this ecosystem varies depending on slope aspect, altitude, soils, and history of human impacts (Armesto and Gutiérrez 1978a, b). An empirical model of vegetation recovery or



succession after fire, logging, and grazing disturbance has been proposed for this ecosystem (Fuentes et al. 1984, Armesto and Pickett 1985, Armesto et al. 1995a), based on mechanisms driving the coalescence of shrub patches generated by past disturbance. Biotic processes such as seed dispersal by animals (Reid & Armesto 2011 a, b) and control of grazers and fire are important drivers of succession. In the upper elevations of the Andes the sclerophyllous scrub gives way to the Andean steppe of central Chile. Current threats: Major impacts on this Mediterranean-climate ecosystem are related to the long history of logging and anthropogenic fire (Fuentes et al. 1984; Fuentes 1990; Armesto et al., 1995; Figueroa et al., 2004; Figueroa and Jaksic, 2004; Montenegro et al., 2004) and more recently to increasing land clearing for agriculture, forestry, and urban expansion (Armesto et al. 2010). Extensive pastures dominated by exotic weeds (e.g., Medicago sativa) with scattered Acacia caven shrubs, as well as massive plantations of exotic pines and eucalypts, have replaced areas originally covered by native woodlands, with consequent losses of biodiversity (Armesto et al. 2007). Pavéz et al. (2010) reported significant reductions in raptor populations (hawks and eagles), attributed to the decline of native small mammal populations (e.g., Octodon degus) due to the loss of native shrub cover and urban expansion. Distribution: The sclerophyllous matorral ecosystem is widely distributed between Valparaíso and Bio-Bio (32-38o S), occupying a surface of 10,191 km2, equivalent to 1.42% of Chile’s mainland.

7. DROUGHT-DECIDUOUS SCRUB: These ecosystems are characterized by short-lived foliage, absent during the long dry season (austral summer, December-March). Drought-deciduous shrubs occupy the dry end of the aridity gradient in the Mediterranean-climate region. Vegetation is often thorny, with some succulents, and a rich herbaceous cover in the understory. Biota: Among the main drought-deciduous shrubs are Acacia caven, Aristeguietia salvia, Flourensia thurifera, Fuchsia lycioides, Oxalis gigantea, Proustia cinerea, Proustia cuneifolia, Proustia illicifolia, Senna birostris var. arequipensis, Senna cumingii, Talguenea quinquinervia, Trevoa trinervis. Presence of aphyllous shrubs Colletia spinossa and Colletia spinossisima is also common. Abiotic parameters: They predominate in semiarid environments or in equatorial (north)-facing slopes of the Mediterranean-climate region of Chile. Rains concentrate in a short period of the year (austral winter) and plants are exposed to high temperatures and high rates of evapotranspiration, which are unfavorable for evergreen shrubs (Olivares and Squeo 1999). Functional ecosystem model: Species with intermittent foliage tend to start their vegetative activity synchronously with the first winter rains (Olivares and Squeo 1999), and their productivity is greatly enhanced during rainy years. Their root systems are often shallow, with primary access to shallow soil moisture. After the onset of summer drought, most plants in this ecosystem loose all of their foliage or die, entering a dormant stage as rootstocks or seeds. High photosynthetic rates



characterize the wet period. Current threats: A conspicuous component of the drought-deciduous scrub is the ‘espinal’ or Acacia caven woodland. This ecosystem has been promoted by a variety of human land uses, such as forage and firewood (Ovalle and Avendaño 1984; Ovalle et al.1990). These uses have led to rapid degradation; soil carbon content of degraded espinal is decreased by as much as 50% compared to better-conserved systems. Chronic disturbance by fire and grazing is the main factor maintaining the espinal and preventing shrub invasion (Armesto and Pickett 1985). Together with the action of introduced rabbits and native herbivores are factors that limit the colonization of sclerophyllous species (Holmgren et al. 2006). This suggests that some areas dominated today by drought-deciduous and thorny scrub, especially in the central valley, could have been originally covered by sclerophyllous scrub, representing an ecosystem transition mediated by humans. Distribution: It is distributed in the semiarid and Mediterranean-climate region of central Chile, primarily in the central valley and north-facing slopes, at low and mid elevations, occupying an area of 21,376 km2, or 3% of the country’s land area.

8. BROAD-LEAVED SCLEROPHYLLOUS FOREST: This ecosystem represents a native forest type recognized in the survey of Chile’s terrestrial vegetation (CONAF et al.1999). The forest canopy (15-20 m tall) is dominated by broad-leaved trees with perennial and sclerophyllous (tough) foliage, with occasional presence of a diverse set of lianas and vine species. Sclerophyllous forests are restricted to deep, humid ravines in the coast and the Andes and to south-facing slopes of coastal hills, where maritime fogs provide an additional source of moisture (Villagrán et al. 2007). Biota: In the more humid sites, taller trees with a rather restricted distribution, such as Beilshmiedia miersii, Drimys winteri, Crinodendron patagua, Persea lingue, Citronella mucronata y Myrceugenia exsucca, are found (Armesto and Martínez 1978). Less humid sites are dominated by widespread sclerophyllous trees, including Cryptocarya alba, Dasyphyllum excelsum, Escallonia pulverulenta, Myrceugenia correifolia, Peumus boldus, Sophora macrocarpa and Lithrea caustica. Common trees in Andean sites are Kageneckia oblonga, Lomatia hirsuta, Quillaja saponaria and Peumus boldus. Lianas and vines are often endemic and particularly threatened because of their narrow distribution range and continued loss from degraded forests; among them are Lardizabala biternata, Boquila triofoliolata, and Bomarea salsilla (Villagrán et al. 2007). In restricted zones, such as the Ocoa sector of La Campana National Park, sclerophyllous forests integrate the endemic Chilean palm (Jubaea chilensis). Abiotic parameters: Because these ecosystems are greatly restricted to shaded ravines and deep creeks, they are not normally subjected to the long, summer droughts of the Mediterranean climate. Their distribution on oceanic coastal hills exposes these ecosystems to greater rainfall (>500 mm per year) than sclerophyllous scrub in the central valley. Humidity is reinforced by the influence of maritime fog on costal mountaintops (above 400 m). Functional ecosystem model: Because the



leathery litter (high LMA) is the main source of recycled nutrients transferred from plants to soil, studies of litter decomposition in this ecosystem are highly relevant. Lusk et al. (2001) showed that litter of Peumus boldus and Cryptocarya alba (two sclerophyllous trees) decomposes more slowly than litter of the deciduous tree Nothofagus obliqua under similar physical conditions. Trees in this ecosystem are therefore slow growing and have low productivity, yet they store large amounts of carbon over time, as shown by the few old-growth stands that remain (Armesto et al., unpublished data). Tree regeneration is largely dependent on the dispersal of seeds to suitable sites by avian fruit-eaters (Hoffmann and Armesto 1995; Vergara et al. 2010) and the seeds ability to germinate immediately after dispersal, as they do not form persistent seed banks. Fires have destructive impacts in this ecosystem as they kill seedlings and seeds, and alter biotic interactions in the soil, impeding regeneration. In addition, the decline of tree cover has negative consequences for the interception of oceanic fog, accentuating the impacts of summer drought, thereby leading to further ecosystem degradation. Current threats: The center of distribution of this ecosystem is the most populated zone of the country (central Chile), in an area strongly influenced by house developments and growing water demands from agriculture. At the same time, this ecosystem is one of 24 global biodiversity hotspots, because of its high endemism and multiple threats (Myers et al. 2000). Presently, the main threats are anthropogenic fire, increasing water extraction from coastal ravines, and the unregulated growth of housing developments on coastal areas. More circumscribed impacts can be attributed to the invasion of the forest understory by exotic species, such as Pittosporum. Distribution: Sclerophyllous forests occupy an area of 12,821 km2 in the Mediterranean-climate region of Chile, which is equivalent to 1.78% of Chile’s mainland.

9. BROAD-LEAVED WARM-TEMPERATE FOREST: This type of forest ecosystem is considered part of the native temperate forest of Chile (Arroyo et al. 1993). Dominant canopy species (25-30 m tall) are trees with broad, perennial foliage. Another name previously used to designate this ecosystem is Valdivian rain forest (Donoso et al., 1985; Cavieres et al., 2005; Luebert and Pliscoff 2005; Carmona et al. 2010; Lara et al., 2012). Biota: This is the center of diversity of temperate tree species, including Nothofagus dombeyi, Eucryphia cordifolia, Aextoxicon punctatum, Myrceugenia planipes, Amomyrtus luma, and Laurelia sempervirens. Occasionally present in the canopy are Drimys winteri, Lomatia hirsuta and some broad-leaved Podocarpaceae (Podocarpus saligna). Climbers and epiphytes are important components of this ecosystem, including Elytropus chilensis, Hydrangea serratifolia, Ercilla syncarpellata, Campsidium valdivianum, Sarmienta repens, and Mitraria coccinea. The stratification is complex and epiphytes contribute significant biomass loads and canopy species richness, serving as substrate for a large number of invertebrates that provide food for canopy birds, thus establishing a diverse “aerial” food web (Díaz et al.



2010, 2012). Abiotic parameters: This ecosystem is restricted to wet-temperate climate, with strong oceanic influence, which is found along the western margin of southern South America (Arroyo et al. 1996). Its best development occurs below 400 m elevation, especially on coastal hills. Annual rainfall is regularly in excess of 1200 mm (up to 2500 mm), from which a minimum of 10% falls during the warm austral summer (Alaback 1991). The ecosystem develops over a variety of soils, but primarily on deep volcanic substrates in the Andes and on shallower clay-dominated mineral soils on the coastal range. Functional ecosystem model: This is the biologically richest forest ecosystem in Chile, where a broad variety of growth habits including lianoid species and vascular and non-vascular epiphytes are represented (Armesto et al. 1996). Multiple vertical strata include understory tree species and canopy emergent trees (e.g., Eucryphia cordifolia) reaching up to 40 m height. The dynamics of warm-temperate forest ecosystems is associated with moderate disturbances due the fall of single trees or small groups of trees. Most trees have broad shade tolerances, regenerating continuously in small tree-fall gaps or under the canopy (Veblen et al.1981; Armesto and Figueroa 1987; Gutiérrez et al. 2008). This ecosystem has one of the greatest accumulations of biomass (over 1,000 tons per hectare) and carbon among Chilean forests (Armesto et al. 1996). Productivity depends primarily on internal nutrient cycling, which is modulated by the rate of organic matter decomposition. Hydrologic nutrient losses are generally limited in old-growth forests (Perakis and Hedin 2002), but logging and fire can enhance leaching and gaseous losses of soil nutrients (Pérez et al. 2010). Current threats: Warm-temperate forests are presently subjected to rapid rates of deforestation, mainly due to the growth of eucalyptus plantations over areas of secondary and juvenile stands. Activities such as agriculture, transportation, and industry (e.g., paper mills) can substantially interfere with the unpolluted atmospheric chemistry of the region (Hedin et al. 1995). Deforestation and forest fragmentation in rural areas (Echeverria et al. 2008) are still important threats to regional biodiversity (Jaña-Prado et al. 2007). Distribution: From 37 to 42o S (Chiloé Island) these wet forests occupy an important fraction of the landscape corresponding to 23,560 km2, or 3.3% of the Chilean mainland.

10. BROAD-LEAVED, COLD-TEMPERATE FOREST: This evergreen forest corresponds to the North-Patagonian rain forest as defined by Veblen and Schlegel (1982) and Villagrán (1985). Dominance is often shared by evergreen broad-leaved angiosperms and conifers (podocarps), and it is distributed at higher latitudes and altitudes (above 500 m) than the previous ecosystem (category A3.4.1.2), particularly in southern Chiloé Island and Aysén. The mean canopy height is 25-30 m, with some longevous Nothofagus nitida emerging above the canopy (up to 35 m). Biota: The characteristic angiosperm dominants are Nothofagus nitida and Drimys winteri, and the main podocarp species are Saxegothaea conspicua and Podocarpus nubigena. Other canopy



trees include Amomyrtus luma, A. meli and Caldcluvia paniculata. The understory can be made of dense patches of Tepualia stipularis, especially in poorly drained sites, with many prostrate horizontal stems. Canopy gaps are rapidly filled up with dense thickets of the native bamboo Chusquea tenuiflora. Tree trunks are typically covered with a green carpet of mosses, lichens and filmy ferns (Hymenophyllaceae) in addition to some vines and vascular epiphytes. Animal inhabitants of this dense and dark understory include Darwin’s fox, the kodkod cat (Leopardus guigna), and particularly the striking and non-volant rynocryptid birds (e.g., Scelosrchilus rubecula and Pterptochos tarnii). These species are abundant but not exclusive of North Patagonian forests. Detailed descriptions of the vascular flora can be found in (Arroyo et al., 1996; Daniels and Veblen, 2003; Amigo et al. 2004; Quintanilla 2005a, b; Álvarez et al. 2010; Bizama et al. 2011). Functional ecosystem model: Cold-temperate rainforests regenerate after intermediate disturbances when groups of canopy trees fall because of windstorms (Gutiérrez et al. 2004). Colonizing tree species are fast-growing Drymis winteri and Nothofagus nitida, which can survive for long periods in the canopy and eventually share dominance with slower growing and shade-tolerant podocarps in old stands (Aravena et al. 2002). Soils are thin (50-60 cm) and rich in organic matter, characterized by high nutrient retention and slow litter turnover (Pérez et al. 1998). In advanced stages of succession, tree growth can be limited nitrogen and phosphorous availability (Aravena et al. 2002). Current threats: These ecosystems were subjected to large destructive fires in 1936 and 1956, when settlers became established in Chiloé and Aysen. Forest cover was reduced by about 50% in the Region of Aysen (Quintanilla 2005b), south of Chiloé. Disturbances caused substantial forest fragmentation nd loss of habitat connectivity (Bizama et al. 2011). Recovery of forest connectivity is an important challenge today. After half a century, forests have been able to regenerate over most of their range, except in areas where the loss of the forest canopy altered the hydrologic cycle producing flooded conditions (Díaz et al. 2007). In large disturbed areas, tree regeneration is extremely slow due to the high water table and restrictive habitat conditions, which favor the colonization by Sphagnum mosses (Díaz and Armesto 2007). Distribution: This ecosystem is distributed from 39 to 47o S, occupying an area of 36,213 km2, equivalent to 5.45% of mainland Chile.

11. BROAD-LEAVED SUBANTARCTIC FOREST: This is the Chilean evergreen forest with lowest number of tree species. Forests are found under colder conditions than the previous evergreen forest types (Arroyo et al. 1993), occupying the margins of the myriad of islands forming the Chonos and sub-Antarctic Archipelagoes, south of Chiloé Island, all the way to Cape Horn (56o S), thus becoming the southernmost forests in the world (Llancabure 2011 Promis et al. 2008). Biota: The main canopy tree is the evergreen Nothofagus betuloides, the longest-lived among all the species of Nothofagus, with a record of 623 years (Gutiérrez et al. 1991). Because of the strong sub-Antarctic winds



the canopy is often 10-15 m in height, except in protected ravines. Other tree species occasionally present in the canopy of these forests are Drimys winteri, Embothrium coccineum, and in some sites the endemic conifer Pilgerodendron uviferum. Shrubs often produce showy red flowers that attract hummingbirds to these southern forests, in particular the conspicuous flowers of coicopihue (Philesia magellanica). Abiotic parameters: This ecosystem is associated with the oceanic climate and wind-exposed conditions of high-latitude fjords and archipelagoes, developed often over thin postglacial soils on rocky substrates, in areas where precipitation can be as high as 5 m per year. Functional ecosystem model: Because of the long-lived Nothofagus trees, dendrochronology studies in these forests have allowed the reconstruction of past climate trends at high latitudes in the southern hemisphere (Villalba et al. 2012), where no other trees grow. Studies indicate that the growth of N. betuloides shows positive correlations with early summer temperatures and negative correlations with summer precipitation (Llancabure 2011). Regeneration must take place in canopy gaps formed by multiple tree falls. Underdeveloped soils and strong winds are often limiting of tree growth. Current threats: In some zones, due to inaccessibility of the islands and steep slopes, this ecosystem remains largely pristine. However, in human sttled areas of Tierra del Fuego and on the mainland N. betuloides forests have been devastated by fire, used as a tool to open land for grazing. Distribution: this forest ecosystem occurs from 40o S (where it is restricted to high elevations), all the way to the southern tip of South America (56o S). The surface occupied by this ecosystem is 24,747 km2, equivalent to 3.44% of Chile’s mainland.

12. CONIFER-DOMINATED FOREST ECOSYSTEMS: Some evergreen forests are dominated by conifers, usually with small imbricated or sessile leaves, shaped as scales (Cupressaceae), and broad (Araucariaceae) or narrow leaves (Podocarpaceae). Angiosperm trees can be present or absent from the canopy of these conifer-dominated ecosystems, but they generally represent a less important component in terms of biomass or numbers. Biota: Conifer forests are dominated by a diverse set of gymnosperms in different parts of their geographic range, constituting three different forest types: Araucaria araucana, Fitzroya cupressoides, and Pilgeordendron uviferum. Some conifers may constitute mono-specific stands, occasionally with a limited number of accompanying species. Non-vascular epiphyte loads (lichens and mosses) can be high and contribute to moisture retention and nutrient cycling in Fitzroya cupressoides forests, forming thick carpets over tree trunks. Abiotic parameters: Some conifer-dominated forests are associated with high temperature contrasts between summer and winter, characteristic of continental environments. Both Austrocedrus chilensis and Araucaria araucana often become established on volcanic ash. Fitzroya forests in the Andes occur generally on steep slopes affected by landslides. Pilgerodendron uviferum-dominated forests on the western side of the Andes are often found on poorly drained sites. Thus,



conifers appear to dominate forests under conditions that are restrictive to angiosperm tree growth (Armesto et al. 1995b). Functional ecosystem model: Dominant conifers in these ecosystems are among the longest-lived in the world: F. cupresoides trees can live for 3600 years (Lara and Villaba 1993), while Araucaria and Austrocedrus live for over 1,000 years. Because of their longevity, these conifers maintain a unique millennial record of climate variation at mid and high latitudes in southern South America. The disturbance regimes that drive the dynamics of these conifer-dominated forests is associated with low frequency anthropogenic or natural fire (Burns 1993; Donoso et al. 1993a; Lara et al. 1999; Battles et al. 2002; Paulino et al. 2009). Current threats: These important ecosystems are threatened mainly by their reduction in area due to past exploitation and increasing fire frequencies (Veblen et al. 2011). Pilgerodendron uviferum stands disturbed by fire in northern Chiloé Island are being replaced by eucalyptus plantations after draining the soils (Laclau 2003). Studies by this author show that site preparation for planting eucalyptus lead to extremely high losses of soil carbon due to organic matter decay, which are not compensated by carbon uptake by growing eucalyptus trees. In the case of F. cupresoides forests, slow litter decomposition implies more efficient carbon retention in the soils than in forests dominated by angiosperms (Pérez et al. 2003). Moreover, these forests have a high amount of carbon stored in long-lived biomass, soils, and slowly decaying coarse woody debris (Battles et al. 2002). Distribution: Conifer-dominated forest ecosystems occur between central Chile (Austrocedrus chilensis) and Cape Horn (Pilgerodendron uviferum). They cover an area of 3,424 km2, which is equal to 0.5% of the country’s mainland.

13. DROUGHT-DECIDUOUS DESERT WOODLAND ECOSYSTEM: These ecosystems cover a limited area dominated by deciduous trees that loose their foliage in the dry season. They represent a seasonally dry subtropical vegetation type, which is largely absent from Chile because of the presence at subtropical latitudes of the dry Atacama Desert. Biota: Common drought deciduous trees are Geoffroea decorticans, Lycium chanar, Prosopis chilensis, Prosopis strombulifera, Prosopis tamarugo. The fruits of both Prosopis and Geoffroea have high nutrient contents and are consumed by humans and wild and domestic animals (Becker 1983; Escobar et al. 2009; Carevic et al. 2012).These forests provide valuable ecosystem services (forage and food) in areas where productivity is limited by extremely dry conditions. Abiotic parameters: The presence of a shallow water table that trees can access via tap root system is a condition for the development of these narrowly distributed desert forests. Functional ecosystem model: Tree species in this ecosystem are considered specialized stress tolerators (sensu Grime 1979), living under strong thermic stress, prolonged water deficit, high salinity, and heavy impact of herbivores (Arce et al. 1990; Medina and Cardemil 1993; Cazebonne et al. 1999). Current threats: The limited area and marginal condition of these forests makes them highly susceptible to processes of increasing aridity and water



extraction for irrigation of agricultural land in arid regions. In areas where timber is rare, these trees have been exploited for various uses, such as making fence posts, firewood and charcoal (García and Ormazabal 2008). In the heart of the Atacama Desert (Pampa del Tamarugal), large-scale plantations of Prosopis tamarugo have been developed over several decades to provide forage to livestock (Carevic et al. 2012). Distribution: Patches of drought-deciduous forests are found scattered in the regions of Tarapacá and Copiapó. They cover an area of 1,180 km2, which is equal to 0.16% of the Chile´s continental area.

14. WINTER-DECIDUOUS, WARM-TEMPERATE FOREST: This ecosystem is dominated by 25-30 m tall trees that drop their leaves during the cold season, with an understory that often includes evergreen trees and shrubs. Numerous vines and epiphytes cover the canopy branches and trunks, including the endemic Chilean national flower Lapageria rosea (Philesiaceae) and its pollinators (Valdivia et al. 2006). Biota: Canopy dominant trees include Nothofagus oblique (roble), N. glauca (hualo), N. alessandrii (ruil), and N. alpina (rauli), mixed with evergreen trees such as Lomatia hirsuta and Azara petiolata, among others. This forest ecosystem is also known as Maulino forest because it occupies the Maule River region, a transition zone between sclerophyllous and deciduous forests (Bustamante et al. 2005). It also includes a mixture of sclerophyllous forest species such as Cryptocarya alba or Aextoxicon punctatum. The Maulino forest is characterized by the presence of fragmented populations of narrow endemic tree species, including Gomortega keule, Pitavia punctata and N. alessandrii (San Martín and Donoso 1996). The avifauna of the Maulino forest ecosystem, as affected by forest fragmentation and habitat degradation has been described by Vergara and Simonetti (2004). Abiotic parameters: Canopy dominance by deciduous species indicates that this environment is subjected to strong seasonality, under the influence of the dry summers of the Mediterranean climate and cold, rainy winters (San Martín and Ramírez 1987). On the other hand, in this region, we find some of the richest soils in the country with respect to nutrient content, and trees in this zone are therefore among the fastest growing, native temperate trees. After massive logging of the Maulino forests in the 1700 and 1800s, soils were cultivated for more than century loosing much of their fertility. Functional ecosystem model: Frequent anthropogenic fires have affected this ecosystem for more than a century, greatly altering nutrient cycling, and reducing organic matter contents and water storage capacity (Litton and Santelices 2003). Exchangeable phosphorous, soil pH and inorganic nitrogen content are incremented. Today, this forest ecosystem has lost its biodiversity and potential ecosystem services due to its extreme fragmentation cover loss (Bustamante et al. 2006). Current threats: Between 1975 and 2000, the Maulino forest cover declined ostensibly, as 67% of the land originally forested was planted with commercial stands of Pinus radiata (Echeverría et al. 2006). Several species of endemic trees are now threatened or endangered (Villa-Suazo and Benoit-



Contesse 2005). This ecosystem has high national priority for conservation measures because of the high threat level, high endemism, and species richness (Simonetti 1999). Part of the problem is due to the near absence of public protected areas and their small surface (Armesto et al. 1992). Forestry plantations, which now exceed by much the land covered by remnants of native forest and are adjacent to narrow protected areas, can have effects on the recycling of nutrients (evergreen vs. deciduous foliage), the diversity of understory plants and animals, the expansion of exotic species, and the incidence of fire (Staelens et al. 2011). Distribution: Restricted primarily to two small regions of the country, O’Higgins and Maule (Fig. 1), this warm-temperate deciduous ecosystem extends southward along the central valley to the latitude of Valdivia. It occupies a land surface of 17,871 km2, which is equivalent to 2.48% of the country’s mainland.

15. WINTER-DECIDUOUS COLD-TEMPERATE FOREST: In this ecosystem, winter-deciduous tree species are almost exclusive dominants, with few or no evergreen tree species present. The canopy can be 20-24 m tall, declining in height at higher elevations and locations where strong westerly winds blow all-year round. Biota: The two most common tree species in the canopy, which rarely occur together, are Nothofagus pumilio and N. antarctica. Occasional understory trees are Drimys winteri and N. betuloides, and the understory often has a rich herbaceous cover that includes colorful orchids and perennial herbs. Abiotic parameters: This ecosystem is restricted to the coldest environments, forming the treeline on the Andean mountains from 35o S southwards, and occupying the most “continental” climates in Patagonia, and on the island of Tierra del Fuego (Arroyo et al. 1993), characterized by snowpack formation during winter. Functional ecosystem model: These forests provide multiple ecosystem services to a vast region of Patagonia, including stream flow regulation, carbon and water storage, and soil protection from erosion at high elevations. They are also found bordering wetlands and ice fields, hence dominating the terrestrial-freshwater transition. Therefore, they have a significant role in the control of the regional effects of climate change (Arroyo 1998). According to Caldentey et al. (2001), timber harvest in deciduous forests alters the annual and seasonal flux of organic residues into streams, as well as litter deposition patterns. Ibarra et al. (2011) showed that silvicultural intervention of Nothofagus pumilio forest changes the internal microclimate due to the loss of canopy cover, in addition to the loss of long-lived carbon stored in soils and biomass (Klein et al. 2008). Despite large-scale disturbance through logging and fire in the 19th and 20th centuries, deciduous sub-Antarctic forests still include some of the largest areas of old-growth and pristine ecosystems in mainland Chile, with a great intrinsic value as biogeochemical baselines from pre-industrial conditions (Hedin et al. 1995). Current threats: In the late 1800s and in the 20th century, large areas of winter-deciduous cold-temperate forests in the two southernmost regions of Aysen and Magallanes were devastated by



settlers, who used fire to open land for gazing livestock. Many deforested areas were later abandoned and have remained without tree regeneration for several decades, causing severe contraction of this ecosystem in southern South America. Lack of regrowth is primarily due to the null sprouting and colonizing ability of the dominant Nothofagus pumilio after fire. The boundary between the sub-Antarctic deciduous forest and the Patagonian steppe has changed over the last century, because of burning and logging practices, leading to contraction of deciduous forests and expansion of steppe (Veblen et al. 2011). More recently, in the mid 20th century, the introduction of beaver (Castor canadensis) and its subsequent population explosion have greatly disrupted hydrologic cycles and destroyed thousands of hectares of riparian vegetation in Tierra del Fuego and adjacent islands, with limited capacity of Nothofagus pumilio or N. betuloides forests to recover from this novel disturbance (Anderson et al. 2006). Distribution: Sub-Antarctic forest ecosystems occupy a broad latitudinal range between the tree line zone of south-central Chile to the lowlands of Tierra del Fuego, for a total range of 32,994 km2 representing 4.6% of the area of continental Chile.

16. SALT-FLAT ECOSYSTEMS: Salt deposits occupy significant areas of high altitude desert in northern Chile. These ecosystems represent remnants of the gradual desiccation of ancient Andean lakes, found in extensive basins without drainage, which accumulated water during pluvial cycles of the Late Quaternary, from 15 to 10 thousand years ago (Gayó et al. 2012). Biota: These areas have sparse plant cover, characterized by species that tolerate high concentrations of salt in the soils, including Puccinellia frigida, Scirpus atacamensis y Sarcocornia pulvinata (Teillier 1998, 2000; Teillier and Becerra 2003; Faúndez-Yancas and Escobar-Vera 2005). Among the aquatic species adapted to this high stress environment are freshwater crustaceans of the genus Artemia (Zuñiga et al. 1999), along with diatoms and extremophile cyanobacteria (Demergasso et al. 2003, 2004, 2007). Abiotic parameters: These dry lakes occur on flat basins, containing shallow ponds of salt water, which make soil conditions strongly hyaline, where few organisms can live (Tellier and Becerra 2003). Functional Model: Lack of knowledge about ecosystem functions in these environments limits the possibility of developing a functional model. Productivity seems highly dependent on seasonal rains, which are normally restricted to the austral summer period. Because of extreme aridity and low rainfall, vegetative activity is strongly limited by low P and N availability (Daniel Alfaro, personal communication). Current threats: Primarily due to mining activities, which include extraction of water, boron, and other types of minerals and salts (Acosta and Custodio 2008). Distribution: These ecosystems are limited to the high altitude desert of northern Chile, from 18°S to 27°S, occupying a surface of 8,360 km2, equivalent to 1.16% of the country’s mainland.



17. SAND DUNE ECOSYSTEMS: This ecosystem is represented by the extensive littoral dune fields, which are identified in the CONAF et al. (1999) survey. Some of these dune fields extend inland several kilometers from the coast (Ramirez et al. 1992). Biota: Vegetation cover contributes to the stabilization of the coastal dune systems, particularly low shrubs (e.g., Ambrosia chamissonis, Carpobrotus chilensis), grasses (Ammophila spp.), and geophytes. Some of these species have wide global distributions on coastal sand dune environments. Abiotic parameters: Dune systems are highly variable in vegetation cover depending on successional stage, which causes high variability in microclimate, soil development, and nutrient contents. Because of the low water retention of the substrate and the marine salt spray, many plant species in this ecosystem can be characterized as drought or salt resistant or halophytes (San Martin et al. 1992). Functional model: Because of their extensive range along the coastline, these ecosystems offer important ecosystem services (Barbier et al. 2010; Everard et al. 2010), protecting land from wind erosion, as they become vegetated, in addition to creating natural barriers against storms and tsunamis. Current threats: These important ecosystem services, and the biodiversity of dune systems, are becoming impaired by the construction of large house developments on sand dunes especially in central Chile, a process that needs further assessment. Distribution: They extend discontinuously along the coastline of mainland Chile, all the way to Chiloé Island (42o S), for about 2 thousand kilometers. Their surface is estimated in 1,910 km2, equivalent to 0.27% of the country’s land.

18. ECOSYSTEMS ON LAVA FLOWS AND VOLCANIC ROCKS AND ASH: These ecosystems occur on scoria from lava flows or volcanic ash substrates, with sparse of no plant cover depending on the age of the substrate. Vegetation development can take a long time because the substrates lack some essential elements for plant growth such as nitrogen (Gallardo et al. 2012). Biota: Pioneer states of ecological succession on volcanic substrates in Congullio volcano (Gallardo et al. 2012) are characterized by a sparse cover of shrubs (Gaultheria, Ericaceae) and extensive areas covered by the moss Racomitrium sp., and the lichen Stereocaulon sp. Functional model: These moss and lichen species have associations with nitrogen-fixing bacteria that enrich the soils and facilitate the establishment of vascular plants on these volcanic substrates (Pérez et al. 2014). The development of vegetation has been described by Gallardo et al. (2012). Distribution: All along the country (from the northern highlands to Patagonia), associated with volcanic cones, from which many are active today. They occupy an area of 1,474 km2 (0.20% of the country), which is variable depending of the extent of volcanic activity and the rate of vegetation succession.

H.4. (19.) LOW ELEVATION DESERT ECOSYSTEM. This arid ecosystem is found at the southern margin of the Atacama Desert, where large numbers of



ephemeral plants (annuals and geophytes) flower in response to irregular pulses of rainfall, which are variable among years, accompanied by the presence of pollen feeders and decomposers. Rain pulses occur in winter and are strongly linked to the positive phase of ENSO (Armesto et al. 1993; Cereceda et al. 2000). During dry years between rain events, there is a complete lack of biological activity. Biota: Annual plant species constitute rich plant communities (Armesto et al. 1993; Vidiella et al. 1989), with a sparse cover of drought deciduous shrubs. Among the annuals, diverse genera are Calandrinia and Cistanthe, Phycela among the geophytes, and Skytanthus acutus among the prostrate shrubs. Seed banks are persistent and shrubs remain dormant during dry years (Gutiérrez 2008). Along with the cover of ephemeral plants, a rich diversity of detritivores, pollen and nectar feeders, and herbivores become active, including coleopterans, snails, and moths. Functional model: Productivity at the southern margin of the Atacama is highly constrained to the short periods of moisture availability, during which a complex trophic chain develops for a limited time. Current threats: The development of an ephemeral plant cover is also associated with transhumance, especially goat grazing, which has negative impacts on the potential response of plants to future rain events. These ecosystems, because of their species richness and fragility, are considered a conservation priority (Squeo et al. 2008). Recent models of regional climate change suggest that these areas will suffer severe declines in rainfall (CONAMA 2006); further work is needed to assess the differential susceptibility of desert flora and fauna to these changes. Distribution: These ecosystems occupy an extent of 29,025 km2 (4.0% of Chile’s mainland), approximately between 24 and 30 S.

20. ANDEAN DESERT ECOSYSTEMS ABOVE TREELINE: This ecosystem is equivalent to the Subnival altitudinal belt as described by other authors (Castro et al. 1982; Villagrán et al. 1982; Squeo et al. 1994; Villagrán et al. 2003), located above the altitudinal limit for woody growth forms in the Andes, with sporadic presence of herbaceous cover. Soils are rocky, nutrient-poor, and often disturbed by runoff and landslides. Abiotic parameters: Its climatic conditions are those of cold desert. Plant establishment is difficult because of periglacial effects due to frost formation in the substrate. It is located above 4250 m at 19 S, above 3600 m at 33 S, and above 1500 m at 40 S. Biota: Few plant species reach the upper limit for vegetation, with sporadic individuals. The most common growth forms are low (<10 cm tall) perennial herbs, with a rosette shape and deep root systems, or cushion plants stuck to the substrate. Some species frequently found at the upper margin of vegetation in the Andes are Oxalis exigua, Pycnophyllum molle, and Viola spp. Distribution: Because of the latitudinal extent of the Andean chain, this ecosystem is present all along the mainland from north to south, below the permanent snowline, occupying an area of 37,116 km2, equivalent to 5,16 % of the country’s land.



21. ABSOLUTE DESERT: This ecosystem corresponds to the lowlands of the Atacama Desert (below 2000 m), characterized by the complete absence of vascular plant cover and the nearly total lack of significant rainfall. The absolute desert extends from southern Peru until the latitude 24o S (Arroyo et al. 1988; Marquet et al. 1998) and the lack of rain is a product of the rain shadow effect of the Andes on the moist easterly winds, the presence of the cold Humboldt current, and the latitudinal position of the eastern Pacific anticyclone. Distribution: In Chile, it extends between 19o (Arica) and 24o S (Antofagasta), covering 100,708 km2, which is 14% of Chile’s mainland.

22. GLACIER ECOSYSTEMS: These are ecosystems without vascular plant cover, where substrates are permanently buried under a thick layer of ice and snow. Biota: However, a significant presence of microorganisms has been detected on the ice surface, buried in the ice, or on emerging rocks. Castello and Rogers (2005) suggest that between 103 and 107 living microorganisms can be found per liter of ice during defrosting. Takeuchi and Koshima (2004) have described seven micro-algal species, including Chlorophyta and cyanobacteria in the southern ice fields of Patagonia, southern Chile. Functional Model: Even though it may seem obvious, it seems necessary to emphasize the capacity of water storage in glaciers and large ice field. Millennial glaciers in the Andes of northern and central Chile hold water reserves that are critical for maintaining lowland ecosystems including people (Hodson et al. 2008). It must be noted that glacier ecosystems are presently being affected by climate warming and human activities (e.g., mining), causing significant reductions in their mass and water holding capacity, especially in the drier zones of northern and central Chile (Marden 1997; Winchester and Harrison 2000; Herrera-Ossandón 2005; Mardones et al. 2011). Massive ice melting in the case of the Patagonian ice fields and Antarctica can also be a factor leading to global sea level rise (Aniya1999). Distribution: Mainland Chile holds large masses of ice and snow in the high Andean peaks and in the extensive ice sheets present in the regions of Aysén and Magallanes. Their surface amounts to 26,889 km2, or 3.4% of the country.

A.2. FRESWATER BODIES AND WETLANDS.

We only distinguished two types of freshwater ecosystems, considering mainly the level of water saturation of soils and the provision of ecosystem services. This classification will be revised in the near future by the Chilean Ministry of the Environment, using a higher resolution.

23. TELMATIC ECOSYSTEMS. This ecosystem-type corresponds to wetlands or seasonally flooded areas where the water table often reaches the surface (Ramirez et al. 2002), but without a permanent waterbody. There is heterogeneity of ecosystems within this category, which includes inundated woody vegetation, temperate and sub-Antarctic moorlands, and high Andean



peatlands. Biota: The rich biodiversity of invertebrates and cryptogams associated with these ecosystems, especially high-Andean bogs and extensive sub-Antarctic moorland, remains poorly documented. Vertebrates and vascular plants are better known, including a rich migratory avifauna associated with wetlands. A complete list of works reporting on this biodiversity can be found in CONAMA (2008). Abiotic parameters: Sediments accumulated in bogs and moorlands are rich deposits of organic carbon, maintained by low rates of organic matter decomposition. In Magellanic moorlands, carbon has accumulated for thousands of years, providing a unique record of past climate changes and their effects on local vegetation (e.g., Hauser 1996). Strong environmental differences depend on the location of the wetlands, with marked differences between high Andean bogs from north-central Chile, which are subjected to contrasting temperature differences under less than 400 mm of rainfall, and austral moorlands, concentrated at latitudes higher than 40 S, under moderate wet oceanic climate, receiving from 2 to 4 meters of annual precipitation. Functional Ecosystem Model: Because of the great water storage capacity of the dense moss carpets, wetlands are key ecosystems for the regulation of hydrologic cycles at the landscape level. In addition, wetlands provide specialized habitats for the flora and fauna and are valuable sources of water for the human societies living in their vicinity (Boavida 1999; Zedler and Kercher 2005; Kandus et al. 2010; RAMSAR 2013). In areas that do not have ice capped mountains, such as Chiloé Island, water discharge from moorlands originate streams and rivers and refill the water table, supplying water to rural populations (Díaz et al. 2008). Current threats: In the Lake District of southern Chile, ecosystems are seriously threatened by the rapid expansion of Eucalyptus plantations, established over drained wetland soils. Once desiccated, habitats can be invaded by exotic Ulex europaeus. Massive harvest of Sphagnum moss biomass for commercial purposes is altering the hydrologic cycle over large regions (Díaz et al. 2008). In the high Andes of northern Chile, the use of groundwater reserves by mining complexes has depressed the water table causing the rapid desiccation of wetlands (Squeo at al. 2006). Distribution: They are found throughout the country, but more abundantly from the Lake District (40 S) to the south, with the most extensive areas of Sphagnum moorlands occurring in Tierra del Fuego (Arroyo et al. 2005), occupying a surface of 45,146 km2, equivalent to 6,3% of the area of the country. More patchily distributed are high-Andean bog ecosystems, with their greatest extension in the Andean highlands (18-22o S).

24. AQUATIC ECOSYSTEMS. Freshwater ecosystems include lakes, ponds, reservoirs, and streams, integrating the neighboring riparian environment. Ponds, lakes and reservoirs are considered in the land cover survey of CONAF et al. (1999), but riparian environments, representing the land-water transition were only mapped in areas with cover by low herbaceous vegetation. Therefore, riparian ecosystems are clearly underrepresented in the present



cartography and their extent and distribution need to be re-assessed in the near future. Biota: Freshwater bodies are characterized by a rich diversity of organisms, many with restricted distributions. Among the endangered taxa in these environments, we can cite 29 species of endemic amphibians (Lobos et al. 2013), the freshwater otter (Lutra provocax), and the majority of freshwater fishes (Habit and Victoriano 2005). Main factors driving the loss of these species are the alteration of aquatic and riparian habitats, damming of rivers, and the massive introduction of predatory fish (salmon and trout). Except in the most inaccessible areas and in the sub-Antarctic region, most aquatic ecosystems in Chile are greatly transformed by human impact and can be considered “novel ecosystems” (sensu Hobbs et al.). Associated riparian environments are also important habitats for the reproduction of numerous species of migrant and resident birds. Functional ecosystem model: Aquatic ecosystems are critical components of the Earth’s hydrologic cycle. They harbor complex food webs, and are sites of energy and matter transfer with the neighboring terrestrial systems that can be critical for sustaining biodiversity (Bailey et al. 2004; Postel and Carpenter 1997). Relevant services of aquatic ecosystems to society include the provision of clean water for human use and agriculture. Riparian zones contribute to lowering the risk of erosion, thermic regulation of water in streams and lakes, filtering sediments, and storage of carbon and nutrients (Lyon and Gross 2005). A riparian ecosystem consisting of a 16-m wide band of vegetation can retain 50% of the nitrogen and 95% of the phosphorus transported downstream. Riparian habitats also provide substrates (e.g., submerged wood) and organic material (e.g., litter) to aquatic bodies. Current threats: Human habitation of riparian environments poses threats to both the integrity of these ecosystems and human life due to the increased risk of floods and mudflows associated with the removal of natural vegetation. Because of recent damming of rivers, changes in the management of reservoirs and lakes, and loss of riparian habitat it is critical that the conservation status of these ecosystems could be monitored in the future, including physical and chemical parameters (Sweeney and Czapka 2004). Distribution: Freshwater bodies are present throughout the country, but they are far more numerous in the temperate region, particularly in the Rivers and Lakes Districts (38-43o S), and extend though the Channel Islands all the way to Cape Horn. Their areal extent is 9,043 km2 (1.26% of Chile’s continental area).

25. ECOSYSTEMS DOMINATED BY INVASIVE SPECIES: These are ecosystems where invasive exotic species have come to dominate energy and matter exchanges without deliberate human intervention. In the satellite-based land cover survey of Chile (CONAF et al. 1999), these sites are covered almost exclusively by spontaneously established exotic plants. However, detailed field data may be necessary to assess additional areas where exotics are now expanding, which are not accounted for by the current survey. Ecosystems dominated by exotics are one of the most dynamic components of global



change. Functional Ecosystem Model: Alteration of nutrient cycles (e.g., by nitrogen-fixing invasive species) and habitat changes associated with these “novel ecosystems” (Hobbs et al. 2009) represent a major driver of biodiversity loss, leading to species extinction worldwide (Mack et al. 2000). Examples of invasive species causing major changes in Chilean ecosystems, which are detectable using satellite images, are the following: (1) Expanding exotic pines (Pinus radiata) invaders from nearby forestry plantations, or deliberately introduced, especially in deforested or burned areas occupy nearly 3 million ha in the Chilean mainland. This process has also been reproduced in other regions where pine species has been deliberately introduced (Gómez et al. 2011). The real magnitude of pine invasion in south-central Chile is still a subject of debate (Bustamante and Simonetti 2005). (2) Widespread presence of Teline monspessulana over large areas of the Mediterranean-climate region of Chile, particularly occupying the understory of exotic tree plantations, native forests with a deciduous canopy, and road and trail networks (Pauchard et al. 2008). Anthropogenic fire has a stimulating effect on the spread of this novel ecosystem. Especially because its N-fixing capacity, its rapid spread may cause major alteration of soil nutrients (García et al., 2007). (3) Dense stands (up to 100% cover) of the shrub Ulex europaeus are distributed over large open, disturbed areas, preventing the establishment of native species. Its recent expansion is associated with its re-sprouting ability after frequent fires used to eradicate it and lack of biological control (Norambuena et al. 2007). Soils are modified permanently because of its capacity to fix nitrogen. (4) The impact of North American beavers (Castor canadensis) has caused the death of many thousands of trees in riparian and aquatic ecosystems of Magallanes, Tierra del Fuego and surrounding archipelagoes, due to the effects of flooding, altered forest cover and soils, and induced changes in floristic composition of wetlands (Anderson et al. 2006; Wallem et al. 2010). Distribution: Novel of ecosystem types are widespread all over the country, from Mediterranean to temperate latitudes (0.15% of the mainland area), but are especially concentrated in south-central Chile (35-45 S). Lack of field data suggests that the real extent of these ecosystems may be underestimated.

26. URBAN ECOSYSTEMS: Cities are defined as population and industrial concentration sites (CONAF et al. 1999, code 1100), where energy and water fluxes, resource demands, and waste products are dominated by human-related activities (Marten 2001). Urban dynamics is dominated by socio-economic drivers and extreme biophysical conditions (Alberti and Marzluff 2004). Research on urban ecosystems is urgently needed to improve understanding of the drivers of their accelerated expansion and population growth (Pickett et al. 1997), both factors having negative impacts on regional biodiversity and human wellbeing. Knowledge of urban ecosystem dynamics in Chile has started to accumulate only recently (e.g., de la Maza et al. 2002; Pauchard et al. 2006; Correa-Araneda et al. 2010). Studies need to focus particularly on unknown



ecosystem parameters, such as energy and matter exchanges with the atmosphere and surrounding land and water, and urban feedbacks on regional climate (Georgescu et al. 2014). Distribution: Urban population in Chile has grown up to nearly 90% in the last decades. Large urban centers (>50,000 people) are distributed in all regions of the country, with >75% of the population and resources concentrated in central Chile (Metropolitan Region, and Valparaíso, 33-34o S), where more then 7 million people live in an area of 2,036 km2 (0.28% of the country). In contrast, the two southernmost regions in Chile have been identified as some of the least populated in the world (less than 5 people per km2), thus holding some of the largest remaining wilderness in temperate regions worldwide (Mittermeier et al. 2003).

27. INDUSTRIAL-MINING COMPLEXES: These anthropogenic ecosystems, marked by massive interventions on land and waters, were mapped based on Chile’s land use and vegetation survey (CONAF et al. 1999). Because of the amount and type of waste products, these ecosystems can have serious impacts on their immediate environment. They impose high demands of energy, water and human population, presumably beyond those of some large cities. Water consumption is particularly important, as these mining centers are concentrated in the arid regions of the country, where much water is derived from underground fossil sources. Recent studies have documented their ecological impacts on terrestrial environments (Lagos 1997; Castro and Sánchez 2003), atmosphere and water pollution (Guevara et al. 2006), soils (Ortiz-Calderón et al. 2008; Ávila et al. 2009), and contamination of coastal areas (Castilla 1983; Ramirez et al. 2005). Distribution. Mining complexes are concentrated mainly in north-central Chile, with direct impacts on 163 km2, which is equivalent to 0.02% of the mainland, but with an extensive and still unaccounted area of indirect impacts.

28. INTENSIVE-USE, AGRO-PASTORAL ECOSYSTEMS: At the time of the survey (CONAF et al.1999), these ecosystems were dedicated to intensive agriculture or pastures for livestock grazing. The main crops in the country are cereals, horticulture, and fruit trees. Livestock include mostly cattle, goats and sheep. In the case of goats, they graze freely in semiarid rangelands partly or fully covered by native vegetation. Functional ecosystem model: These areas dedicated mainly to the production of food for human consumption (including local and international trade). The main ecosystem service associated with these managed ecosystems is provisioning (MEA 2005). However, the quantification of negative externalities of ecosystem management in these lands requires additional studies, including fertilizer use efficiency (Eastman and de Fuentes 2010), and greenhouse gas emissions associated with cattle raising and burning of residues (Li 2000; PNUMA 2007). Oyarzún and Huber (2003) have documented high fluxes of nitrogen (NO3-N) from grazing pastures into streams of the temperate region. Distribution: These intensely managed ecosystems are concentrated in south-central Chile, occupying 7.35% of the



country’s land, precisely in the region where endemic biodiversity is concentrated (Armesto et al. 1998).

29. CONIFER PLANTATIONS: Extensive areas planted with fast growing pines from North America. The regeneration dynamics (rotation period), nutrient fluxes and carbon storage in these ecosystems are different from areas with native forest cover or eucalyptus plantations. Biota: The main tree species planted (>95% of the area) are Pinus radiata, but smaller areas have been plated with P. ponderosa and P. contorta. Some of these plantations can be considered “novel ecosystems” (sensu Hobbs et al. 2009), which are used by a small number of native species, depending on management practices, and are sites where invasive species are expanding (Estades and Escobar 2005). Functional ecosystem model: Field studies indicate that plantation ecosystems have hydrologic cycles that differ markedly from native forests, due to greater rain intercepting surfaces, increased evapotranspiration, and reduced water percolation (Huber et al. 2008, 2010). Regarding nutrient cycling, pine-dominated ecosystems have much slower litter decomposition rates than litter of deciduous Nothofagus that dominate native forest the same regions (Lusk et al. 2001). However, litter mass loss under pines is accelerated with respect to the decay rates of litter from sclerophyllous species in neighboring native woodlands (Lusk et al. 2001). These results suggest that large-scale substitution of native ecosystems by conifer plantations can significantly alter the rates of nutrient return to soils, with respect to forests dominated by deciduous or sclerophyllous tree species. Although plantation ecosystems, receiving government subsidies, are linked to national efforts to mitigate climate change through carbon capture (Gilabert et al. 2007), recent studies suggest that carbon gains from fast-growing plantations can be offset by substantial carbon losses due to litter and woody debris decomposition and soil erosion (Houghton 2005). Current threats: Large-scale forestry based on exotic species can threaten native ecosystems due to invasion of neighboring native communities, increasing the risk of forest fires and externalities related to the use of pesticides and fertilizers (Gomez et al. 2011; Rejmánek 1996; Rejmanek and Richardson 1996). Distribution: Plantations of Pinus radiata presently occupy 21,041 km2, equivalent to nearly 3% of the country.

30. BROAD-LEAVED PLANTATIONS: Forestry plantations dominated by evergreen, broad-leaved eucalyptus trees originated from Australia are massive monocultures in south-central Chile. They are used primarily as resources for the fiber and pulp industry. One species, Eucalyptus globulus comprises more than 60% of forestry plantations, with other less extensive areas dominated by monocultures of Eucalyptus nitens, Eucalyptus camaldulensis, and Acacia melanoxylon. Functional Ecosystem Model: Litterfall is the main process returning nutrients to the soil in both native forests and plantations. Schlatter et al. (2006) reported accumulations of nitrogen, phosphorous, potassium, and calcium in litter under E. nitens of 90-100, <15, 20-40, and 140-230 kg ha-1,



respectively. Some of these nutrients may be present in mobile forms that can be subjected to hydrologic loss. Gilabert et al. (2007) suggest that eucalyptus plantations may be able to fix more carbon per hectare than other exotic tree plantations. An analysis of rotation times is necessary to assess the storage capacity of soils and biomass because these plantations are often proposed by private industry as part of mitigation plans. Because of the high rates of evapotranspiration, plantations of eucalyptus over bog and moorland sites can increment hydrologic losses (Huber et al. 2010) and enhance soil desiccation. Distribution: Eucalyptus monocultures are found mainly in the regions of Bio Bio, Araucanía, and Maule, but are recently expanding southwards reaching the Island of Chiloe (42 S), covering an area of 4.2 km2, about 0.6% of the country.



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