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Pacific Coast: CPD Site SA43

LOMAS FORMATIONS
OF THE ATACAMA DESERT

Northern Chile

Location:  Series of almost 50 isolated localities in Atacama Desert along coastal Chile, between latitudes 18°24'-29°54'S.
Area: 
Less than 5000 km² of vegetation formations dispersed in western fringe of 291,000 km² of arid lands.
Altitude: 
.
0-1100 m.
Vegetation: 
Community compositions are highly variable: mixtures of annual, short-lived perennial and woody scrub vegetation.
Flora: 
c. 550 spp. of ferns, gymnosperms and flowering plants; very high endemism (over 60%); many threatened species.
Useful plants: 
Unusual ecophysiological characteristics make the plants valuable as potential genetic resources.
Other values: 
Communities highly endemic; support for mammal, bird and insect communities; tourism.
Threats: 
Urbanization, mining, pollution, road construction, livestock grazing - especially numerous goats, fuelwood gathering, commercial plant collecting, erosion.
Conservation: 
La Chimba National Reserve (c. 30 km²); Pan de Azúcar National Park (c. 438 km²).

Map 77: CPD Site SA42
References

Geography

The Atacama Desert forms a continuous strip for nearly 1600 km along the narrow coast of the northern third of Chile in its Regions I-IV: Tarapacá, Antofagasta, Atacama, northern Coquimbo, from near Arica (18°24'S) southward to near La Serena (29°55'S). Adjacent to the Pacific are uplifted marine terraces or the coastal range arising directly from the ocean; farther eastward is a large alluvial plain, which is bordered by the Andes. Although arid conditions extend in the Andes up to c. 3500-4000 m, the Atacama Desert is usually deemed to end at their base or near 1500 m on the drier slopes (Börgel 1973). The faulted coastal mountains (mostly 500-1000 m high) are composed of Cretaceous sediments (limestone and sandstone) over more ancient masses of crystalline rocks. Whereas desert conditions are continuous along the coast of Peru and Chile, the topography, climate and vegetation of each desert are distinct (Harrington 1961; Rundel et al. 1991; Duncan and Dillon 1991; Rauh 1985). For some generalities and comparison with the coastal Peruvian Desert, see the Data Sheet on Lomas Formations - Peru, CPD Site SA42.

The conditions for development of hyper-arid habitats in this region are similar to those in coastal Peru, including the influence of cool sea-surface temperatures associated with the northward-flowing oceanic Humboldt Current, and strong atmospheric subsidence associated with a positionally stable, subtropical anticyclone. The result is a mild, uniform coastal climate with regular thick stratus cloud banks well below 1000 m during the winter months. The northern seaport Iquique has average monthly temperatures between 14.5°C (September) and 21°C (March); the recorded absolute extremes have been 8°C and 31.3°C. The Tropic of Capricorn crosses the region slightly north of the city of Antofagasta.

Where the coastal topography is low and flat, the winter stratus layer dissipates inward over broad areas with little biological effect, but where isolated mountains or steep coastal slopes intercept the clouds, a fog zone develops with a stratus layer concentrated against the hillsides. These fogs ("camanchacas") are the key to the extent and diversity of vegetation throughout the desert. The moisture allows the development of fog-zone plant communities termed "lomas" (small hills) between sea-level and 1100 m. These plant formations also have been called the fertile belt, fog oases or meadows on the desert. In Chile almost 50 discrete localities support such vegetation (see combined Map 77). The Chilean area actually covered by this vegetation probably is less than 5000 km², even during periods of optimal weather and maximum development.

Much of western South America is influenced by brief periods of heavy rainfall and relatively high temperature associated with rare but recurrent El Niño events (Dillon and Rundel 1989; Quinn, Neal and Antúnez de Mayolo 1987; McGlone, Kershaw and Markgraf 1992). However, heavy storms are virtually unknown along the north coast of Chile, whereas northern Peruvian coastal cities occasionally receive torrential rains. The maximum precipitations for a 24-hour period recorded in Chile at Arica, Iquique and Antofagasta are 10, 13 and 28 mm respectively (Miller 1976). Johnston (1929) reported 17 mm at Antofagasta during a single day in 1925, a strong El Niño year. However, El Niño years can fail to produce increased precipitation in northern Chile. The El Niño event of 1983 caused heavy rains from northern to southern Peru, and a relatively high 7.3 mm in Iquique, but no notable rains farther south (Romero and Garrido 1985; Rutllant 1985). Nonetheless, that year's fogs appear to have been unusually dense, resulting in excellent flowering in the Chilean lomas formations (Prenafeta 1984).

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Vegetation

Lomas formations occur as discrete communities or islands of vegetation separated by hyper-arid territory devoid of plant life. Since growth is dependent upon available moisture, the climatic patterns determine plant distributions. Topography and substrate combine to influence the patterns of moisture availability and areas of suitable habitat. The vegetation is largely restricted to the coastal escarpment and lower portions of numerous gorges ("quebradas") that traverse it. Ecological requirements and tolerances of individual species ultimately determine community composition; species endemism exceeds 60% (Rundel et al. 1991). Broad overviews of the vegetation have been provided by Schmithüsen (1956) and Rauh (1985). The following account briefly describes the larger lomas formations, including the data available on species diversity.

Northern Atacama Desert

The coastal zone from just north of Arica (near Peru) to the seaport Antofagasta is essentially barren, having extremely low species diversity and density. The values of mean annual precipitation reported for northern Chile are the lowest of any long-term records in the world, with Arica and Iquique averaging 0.6 mm and 2.1 mm respectively; in several consecutive years there may be no rainfall. Along the escarpment above 500 m grow scattered individuals of cacti - the columnar Eulychnia iquiquensis, and Copiapoa species. Near Iquique the vegetation is only weakly developed; some 20 species have been recorded. An exception 10 km east of Iquique at 990-1100 m is a large community of Tillandsia landbeckii on pure sand. The communities near Tocopilla with over 60 species recorded and Cobija with 15 species have only a few endemics, including the rare Malesherbia tocopillana, Mathewsia collina and Nolana tocopillensis (Jaffuel 1936).

The slopes and coastal plain around Antofagasta are virtually free of any vegetation except scattered individuals of Eulychnia iquiquensis and Copiapoa species. North of the city within Quebrada La Chimba vegetation tends to occur along the margins of the dry river canyon where runoff concentrates. Johnston (1929) collected around the city in the spring of 1925 following heavy rains and found 34 species; herbarium records and recent fieldwork indicate that nearly 60 species occur (Dillon, unpublished).

Cerro Moreno, a prominent rocky headland north-west of Antofagasta, has a small fog-dependent plant community on its south-western slope at a relatively high elevation (Follmann 1967). The vascular plant species recorded number just 28, less than half as many as at nearby La Chimba. The lower margin of the fog zone on Cerro Moreno is near 600 m and marked by scattered populations of Copiapoa. Although Eulychnia iquiquensis is dominant, its vigour is poor. The shrub stratum is most notably absent within the zone - only a few shrubby species occur, such as Heliotropium pycnophyllum, Ephedra breana and Lycium deserti. Annual and perennial herbs include Cynanchum viride, Viola polypoda and Argythamnia canescens. Two essentially non-desert species disjunct from central Chile have small populations at the site: Acaena trifida and Colliguaja odorifera. Vegetation along the coast below this fog zone is virtually absent, with only a few scattered individuals of Nolana peruviana and Tetragonia angustifolia.

Southern Atacama Desert (pictures)

The richest development of the fog-zone vegetation in northern Chile is 50 km north of Taltal near the mining village Paposo (25°03'S), with no fewer than 230 associated species of vascular plants. The central portion of this fog zone is dominated by Euphorbia lactiflua and Eulychnia iquiquensis (Rundel and Mahu 1976). Other important shrubby species are Echinopsis coquimbana (Trichocereus coquimbanus), Oxalis gigantea, Lycium stenophyllum, Proustia cuneifolia, Croton chilensis, Balbisia peduncularis and Tillandsia geissei. Annuals include Viola litoralis, V. polypoda, Cruckshanksia pumila, Alstroemeria graminea, Malesherbia humilis and Chaetanthera glabrata. Above and below the central fog zone the coverage of species drops sharply and growth forms of the dominants change. Whereas drought-deciduous tall shrubs dominate within the zone, the importance of cacti and low semi-woody subshrubs increases away from the centre.

Dense mounds of the bromeliads Deuterocohnia chrysantha and Puya boliviensis are locally common along the coastal flats and mark the lower margin of the fog zone at c. 300 m. The coastal plain below the fog supports broad stands of Copiapoa cinerea var. haseltoniana and scattered shrubs, in communities similar to those at Pan de Azúcar to the south. Above the uppermost limits of regular fog at c. 800 m, Copiapoa stands again become dominant along with Polyachyrus cinereus, Oxalis caesia, Nolana stenophylla, N. villosa, N. sedifolia and N. peruviana. Some shrub species such as Gypothamnium pinifolium and Chuquiraga ulicina occur below and above this fog zone (Rundel and Mahu 1976; Johnston 1929). Large areas of coastal terraces from north of Paposo to Pan de Azúcar are covered by annuals, predominantly Nolana aplocaryoides.

Pan de Azúcar National Park (c. 26°04'-26°09'S, 29 km north of Chañaral) has been described in some detail; the mean annual temperature is 16.4°C. Two types of plant communities are present (Rundel et al. 1980). One is composed of low succulents, with virtually mono-specific stands of Copiapoa cinerea var. cinerea (picture) (form "columna-alba") or C. cinerea var. haseltoniana (Mooney, Weisser and Gulmon 1977; Gulmon et al. 1979; Ehleringer et al. 1980; Anderson et al. 1990). Small mostly subterranean cacti such as Neoporteria sp. (= Neochilenia malleolata, Thelocephala krausii) are also present (Weisser 1967a, 1967b). A second type of community consists of open stands of less succulent shrubs on "bajadas" and in washes. The dominants are Nolana mollis, Heliotropium linearifolium, Gypothamnium pinifolium, Oxyphyllum ulicinium, Euphorbia lactiflua, Tetragonia maritima and Eremocharis fruticosa.

The inland region from Chañaral south to the Copiapó River marks the northernmost portion of El Norte Chico (Muñoz-Schick 1985). This region has open communities of scattered low shrubs of Skytanthus acutus, Encelia canescens, Frankenia chilensis and Nolana (Alona) rostrata. Annuals and short-lived perennials in favourable habitats include Perityle emoryi, Oenothera coquimbensis, Adesmia latistipula, Astragalus coquimbensis, Cruckshanksia verticillata, Fagonia chilensis and Tetragonia angustifolia. Numerous geophytes (with bulbs or corms) are also conspicuous in the region when sufficient moisture becomes available.

Immediately to the south of Chañaral several species have their southernmost distribution, including Gypothamnium pinifolium, Dinemandra ericoides, Nolana aplocaryoides and Tiquilia litoralis. The coastal strand and dune formations of the littoral belt from Chañaral southward to Caldera and the Copiapó River have many of the same species that occur in the interior, and additional characteristic species such as Nolana divaricata, Heliotropium stenophyllum, H. linearifolium, H. pycnophyllum, Oxalis gigantea, Ophryosporus triangularis, Ephedra andina, Euphorbia lactiflua, Bahia ambrosioides, Senna cumingii var. coquimbensis, Tetragonia maritima, Echinopsis coquimbana and Eulychnia acida (Kohler 1970; Reiche 1911; Opazo and Reiche 1909).

Quebrada El León (26°59'S) in the coastal area 15 km north of Caldera is a canyon with extraordinary rocky formations, running water and pools. The rich and diverse vegetation includes Euphorbia lactiflua, Opuntia sphaerica (O. berteri auct.), Eulychnia breviflora, Copiapoa spp. and the perhaps endemic Neoporteria sp. (= Neochilenia transitensis) (Anderson et al. 1990).

The broad lower valley of the Copiapó River funnels sufficient moist maritime air masses nearly 50 km inland to support scattered populations of coastal lichen species on the rocky hills around the city. However between Copiapó and Vallenar, vegetation is virtually or totally absent over extensive areas. Thin veneers of sand cover rocky gentle slopes over much of this region. The climate as well as the effects of human activities have been important in producing this arid landscape. The only significant species along many km of the Pan American highway are perennials, such as Argylia radiata (picture), Bulnesia chilensis, Encelia canescens, Frankenia chilensis, Caesalpinia angulata, Polyachyrus fuscus, P. poeppigii, Nolana rostrata and N. pterocarpa (Kohler 1970).

Quebrada Carrizal Bajo (28°08'S) is in the coastal area between Totoral and Huasco. The coastal formations in this region have vegetation corresponding to the Desierto Costero Huasco (Gajardo 1987). Due to occasional precipitation, this vegetation has more permanence and continuity than the vegetation occurring farther north. The endemics include Copiapoa echinoides, C. malletiana, Neoporteria carrizalensis and N. villosa var. laniceps (Anderson et al. 1990). At times this region is dominated by geophytes, which can cover expanses of the desert spectacularly. Near Huasco, dune formations support a well-developed cover of halophytic species, notably Sarcocornia fruticosa, Carpobrotus chilensis and Distichlis spicata (Kohler 1970).

The geographical southern limit of the Atacama Desert is subject to a variety of views, with most opinions favouring either just north of La Serena (29°54'S) or 280 km farther north at the Copiapó River (27°20'S). La Serena receives erratic precipitation averaging less than 130 mm per year, compared with only 29 mm at Copiapó and almost 0 mm through much of northern Chile. On a floristic basis, La Serena is clearly the most appropriate periphery. Near La Serena north of the Elqui River Valley, semi-arid coastal scrub vegetation is replaced by desert succulent communities with floristic affinities to the regions northward (Rundel et al. 1991).

Desert vegetation of interior valleys between Copiapó and La Serena is sparse but regularly present, compared to the more arid regions farther north. Beginning c. 40 km south of Vallenar, the vegetation coverage increases markedly and continues to north of La Serena. Succulent communities with mound-forming species of Eulychnia and Opuntia (Tephrocactus) are dominant, and the shrubby Heliotropium stenophyllum, Balsamocarpon brevifolium and Bulnesia chilensis are common. In wash habitats important shrubs include Cordia decandra, Adesmia argentea, A. microphylla and Ephedra.

North of La Serena along the coast, precipitation is sufficient to support relatively high vegetation coverage in a transitional community with representatives from the coastal matorral of central Chile (Rundel 1981). Frequently encountered species include Echinopsis coquimbana, Oxalis gigantea, Lobelia polyphylla, Myrcianthes coquimbensis, Puya chilensis, Lithrea caustica, Heliotropium stenophyllum, Nolana coelestis, N. crassulifolia and N. sedifolia. Coastal dunes are dominated by Nolana divaricata and Tetragonia maritima (Kohler 1970), and associated species include Nolana (Alona) carnosa, Ephedra breana and Skytanthus acutus. Numerous ephemeral geophytes are manifest when rare rains fall, including Alstroemeria spp., Tecophilaea spp., Leucocoryne spp., Rhodophiala spp. and Leontochir ovallei (Muñoz-Schick 1985).

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Flora

Recent catalogues on the vascular flora of Chile have been compiled (Arroyo, Marticorena and Muñoz 1990; Marticorena and Quezada 1985; Gajardo 1987), and a floristic inventory of the c. 7000 km² of lomas formations in Chile and Peru has been completed (pictures) (Dillon, in preparation for publication). The northern Atacama (El Norte Grande) is a centre of pronounced endemism - a large number of species range only slightly to its north or south. The flora of this region was the focus of R.A. Philippi's first collecting efforts in Chile in 1853-1854, culminating in the Florula Atacamensis (1860). This is the type locality for nearly 200 of his newly described species. Reiche (1907, 1911, 1934-1937) contributed a series of papers with descriptions and notes on the flora of northern Chile. Johnston (1929) and Werdermann (1931) described the flora of the northern area from field studies in 1925 (a strong El Niño year). A photographically illustrated Flora includes about the southern quarter of the Atacama, in El Norte Chico (south of Chañaral to north of Petorca-La Ligua) (Muñoz-Schick 1985).

The estimated total numbers of taxa of vascular plants represented within the Atacama Desert's lomas formations are 80 families, 225 genera, 550 species (Dillon, in prep.). The families with the most species diversity are Asteraceae (33 genera, 65 spp.); Nolanaceae (picture) (2 genera, 37 spp.); Leguminosae (13 genera, 35 spp.); Cactaceae (7 genera, c. 30-40 spp. - cf. Hoffmann-J. 1989, Anderson et al. 1990); Boraginaceae (5 genera, 26 spp.); Solanaceae (9 genera, 19 spp.); and Apiaceae (8 genera, 17 spp.). Only 68 of the species (c. 7%) reach the lomas in Peru north of 18°S latitude.

The lomas formations from Arica to Antofagasta have little floristic affinity to the Peruvian formations immediately to the north. Of the 117 species of vascular plants recorded from this region by Johnston (1929), only five species (4%) were reaching their southern limit. In contrast, 89 species (76%) represented northern extensions of the richer flora south of Antofagasta. The remaining 23 species (20%) were endemic to the region, most notably species of Nolana.

The area around Arica itself clearly is related floristically to the Peruvian lomas, but many genera and species drop out at or immediately to the south of Arica. For example, the four most important groups of coastal Peruvian cacti either do not cross into Chile (the Loxanthocereus group of Cleistocactus, Neoraimondia) or do not occur south of Arica (the Islaya group of Neoporteria, Haageocereus - except H. fascicularis occurs well above the lomas inland in Tarapacá); and similarly three of the most important cactus groups of northern Chile either do not cross into Peru (Copiapoa) or hardly do so (Eulychnia, Neoporteria sensu stricto) (Rauh 1958; Hoffmann-J. 1989). Terrestrial Tillandsia species are a dominant aspect of the coastal vegetation of southern Peru, but of very limited distribution in the Atacama Desert. The southernmost limit of these bromeliads is between Iquique and Tocopilla, where pure stands of T. landbeckii occur.

Important shrubby species primarily confined to the Atacama Desert include Berberis litoralis, Anisomeria littoralis, Atriplex taltalensis, Adesmia viscidissima, Croton chilensis, Balbisia peduncularis, Nicotiana solanifolia, Teucrium nudicaule, Monttea chilensis, Stevia hyssopifolia, Senecio almeidae, Gutierrezia taltalensis and Haplopappus deserticola. The region is a centre of diversity for the Nolanaceae (Johnston 1936), with several species typically occurring sympatrically. Frequently encountered nolanas include Nolana mollis, N. salsoloides, N. sedifolia, N. peruviana, N. leptophylla, N. villosa and N. aplocaryoides.

A number of genera with coastal perennials cross the floristic barrier south of Arica, including Alternanthera, Ophryosporus, Tetragonia, Oxalis, Calandrinia, Senna, Palaua, Tiquilia, Heliotropium, Caesalpinia, Opuntia (Tephrocactus sensu lato) and Urocarpidium. Nolana is distributed from northern Peru (as well as the Galápagos Islands) to central Chile, with peaks of species diversity near Mollendo in Peru and Paposo in Chile - only N. lycioides is distributed in both countries.

Despite the barrier, a number of species also are successfully established over the entire length of the Peruvian and Atacama deserts, such as Apium (Ciclospermum) laciniatum, Encelia canescens, Chenopodium petiolare, Mirabilis prostrata and Loasa urens. Alstroemeria paupercula (picture) occurs from Chala in south-central Peru to Caldera in Chile. Pasithea coerulea and Fortunatia biflora occur from southern Peru to central Chile. Although Solbrig (1976) drew a strong significance to the presence of floristic elements of the Chaco region of Argentina in the Atacama Desert, few of the species mentioned actually enter the coastal desert region - most of them are found in central Chile or on the higher slopes of the Andes.

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Useful plants

This highly endemic flora is of inestimable importance. There are some traditional uses of species by the local inhabitants (Aronson 1990; Bittmann 1988), e.g. food from Oxalis spp., medicinals from Salvia tubiflora and Ephedra spp. Some plants of the Atacama Desert have developed unique ecophysiological traits through their evolutionary histories that will be of interest particularly in the genetic engineering of crops for agriculture and horticulture.

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Social and environmental values

Global warming is difficult to document, yet important. Floristic distributional data combined with long-term precipitation records suggest that there has been a steady loss of biodiversity in northern Chile over the last 100 years (Rundel and Dillon, unpublished). While the trends will be difficult to reverse, these communities provide unusual opportunities to study the ongoing process of global climate change.

The intrinsic value of the Atacama Desert's plant and animal communities lies in the unique nature of their composition, the high levels of endemism and some species' remarkable adaptations for survival in some of the planet's most demanding conditions. Even rolling lichens are found (e.g. Rocella cervicornis) (Follmann 1966). These islands of habitat are essential for very many of Chile's threatened species (e.g. Benoit-C. 1989), and the lomas species have often been little studied or are even undiscovered. Recent scientific interest has led to more detailed research on the dynamics of this flora.

Some areas nearby have archaeological importance. The beauty and rarity of the lomas formations provide opportunities for tourism combined with scientific studies. If the impact on the delicate communities is controlled through supervision, lomas formations can be enjoyed by the public and preserved for the coming generations. Environmental education, on the importance, the rarity and the unusual characteristics of these natural resources, is desperately needed.

For example, Quebrada El León needs some recuperation from overuse and could become a lasting and informative oasis as a nature reserve for residents of Caldera and Copiapó. Punta de Teatinos (29°48'S), just north of La Serena, has interesting flora (e.g. Neoporteria jussieui var. dimorpha) and fauna - an estuary attracts many freshwater and marine birds. Both the land and water ecosystems require adequate protection, and this area could be a focus of nature recreation for residents of La Serena and Coquimbo.

Economic assessment

The primary utility of the Atacama Desert has been in mineral extraction and harvesting marine resources. The lomas formations are not generally considered of great economic potential due to the harsh, highly variable and unpredictable environmental conditions in the coastal desert.

Investigations of the potential water from wind-blown fog have shown that considerable moisture is available (Muñoz 1967). The generation of water by condensation from the available fog sources recently has stimulated local agriculture, which supplements costly produce brought in from several thousand km to the south.

Nonetheless, the poor soils and scarce water reserves in the northern Atacama Desert will not allow large-scale agricultural production immediately along the coast. Persistent agriculture or silviculture in lomas formations would be extremely damaging.

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Threats

Since many sites have become accessible by road only recently (e.g. within the past c. 12 years), the Atacama's specialized ecosystems remained well preserved until recent times. Road construction in association with mining operations is increasing human occupation in the region. With the rise in copper prices during the 1980s, reactivation of mining activities utilizing large quantities of sulphuric acid has had an essentially undocumented impact on terrestrial and marine life (e.g. Anderson et al. 1990).

The mining, overgrazing by goats, extraction of fuelwood and commercial gathering of plants and animals have all combined to impact upon most regions in northern Chile. Efforts are needed to curtail hunting of rare species, such as some cacti and bulbs (Benoit-C. 1989; Anderson et al. 1990), guanacos, chinchillas and various birds including Andean condors, and in the regulation of the exploitation of marine species, e.g. shellfish, fish and algae.

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Conservation

La Chimba National Reserve (c. 30 km²) was recently established; it is c. 15 km north of Antofagasta and regularly visited. Creation of a National Park in the exceptional area around Paposo (2000 km², with c. 345 species) would be a major step for protecting a substantial portion of the region's diversity and endemics (Reyes and Zizka 1989; Hoffmann-J. 1991; Anderson et al. 1990). One large nationally protected area exists within the extreme desert region - Pan de Azúcar National Park (picture) (established in 1986), which covers c. 438 km². It has been recommended (Anderson et al. 1990) that this park be expanded northward to include Quebrada Esmeralda (25°50'S) and Quebrada de Las Lozas (25°41'S), which would protect areas very rich in cacti, and 15 km southward.

The recent discovery of rare and highly endemic species (e.g. Copiapoa spp., Griselinia carlomunozii and Tillandsia tragophoba - Anderson et al. 1990; Dillon and Muñoz-Schick 1993; Dillon 1991) shows the need for additional conservation measures. Since Atacama plants have narrowly confined geographic distributions, there needs to be a series of National Reserves or parks to help ensure the future of these unique communities (Benoit-C. 1989; Anderson et al. 1990). Many of the cacti are propagated in nurseries at least abroad, which diminishes pressures to collect them from the wild, and export of cacti is regulated under CITES.

The estimated 550 species of vascular plants, most of which are largely endemic to the Atacama Desert, represent over 10% of the estimated 5100 continental species in Chile (Marticorena 1990). Conserving the flora of the Atacama Desert should be an immediate priority for local and national authorities and is internationally worthwhile.

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Map 77. Lomas Formations, Peru (CPD Site SA42) and Lomas Formations of the Atacama Desert, Northern Chile (CPD Site SA43)

References

Anderson, E.F., Bonilla-F., M., Hoffmann-J., A.E. and Taylor, N.P. (1990). Succulent plant conservation studies and training in Chile. World Wildlife Fund-U.S., Washington, D.C. 136 pp.

Aronson, J. (1990). Desert plants of use and charm from northern Chile. Desert Plants 10(2): 6574, 79-86.

Arroyo, M.T. Kalin, Marticorena, C. and Muñoz, M. (1990). A checklist of the native annual flora of continental Chile. Gayana, Bot. 47: 119-135.

Benoit-C., I.L. (ed.) (1989). Red Book on Chilean terrestrial flora (Part One). Corporación Nacional Forestal (CONAF), Santiago. 151 pp.

Bittmann, B. (1988). Recursos y supervivencia en el Desierto de Atacama. In Masuda, S. (ed.), Recursos naturales Andinos. University of Tokyo, Tokyo. Pp. 153-208.

Börgel, R. (1973). The coastal desert of Chile. In Amiran, D.H.K. and Wilson, A.W. (eds), Coastal deserts: their natural and human environments. University of Arizona Press, Tucson. Pp. 111-114.

Dillon, M.O. (1991). A new species of Tillandsia (Bromeliaceae) from the Atacama Desert of northern Chile. Brittonia 43: 11-16.

Dillon, M.O. (in prep.). Flora of the lomas formations of Chile and Peru. Fieldiana: Botany.

Dillon, M.O. and Muñoz-Schick, M. (1993). A revision of the dioecious genus Griselinia (Griseliniaceae), including a new species from the coastal Atacama Desert of northern Chile. Brittonia 45: 261-274.

Dillon, M.O. and Rundel, P.W. (1989). The botanical response of the Atacama and Peruvian desert floras to the 1982-83 El Niño event. In Glynn, P.W. (ed.), Global ecological consequences of the 1982-83 El Niño-Southern Oscillation. Elsevier Oceanography Series, Amsterdam. Pp. 487-504.

Duncan, T. and Dillon, M.O. (1991). Numerical analysis of the floristic relationships of the lomas of Peru and Chile. Abstr. Amer. J. Bot. 78: 183.

Ehleringer, J.R., Mooney, H.A., Gulmon, S.L. and Rundel, P.W. (1980). Orientation and its consequences for Copiapoa (Cactaceae) in the Atacama Desert. Oecologia 46: 63-67.

Follmann, G. (1966). Chilenishe Wanderflechten. Ber. Deutsch. Bot. Ges. 79: 452-462.

Follmann, G. (1967). Die Flectenflora der nordchilenischen Nebeloasen Cerro Moreno. Nova Hedwigia 14: 215-281.

Gajardo, R. (1987). La vegetation naturelle du Chile: proposition d'un systeme de classification et representation de la distribution géographique. Ph.D. thesis. Université de Droit, d'Economie et des Sciences d'Aix-Marseille. 282 pp.

Gulmon, S.L., Rundel, P.W., Ehleringer, J.R. and Mooney, H.A. (1979). Spatial relations and competition in a Chilean desert cactus. Oecologia 44: 40-43.

Harrington, H.J. (1961). Geology of parts of Antofagasta and Atacama provinces, northern Chile. Bull. Amer. Assoc. Petroleum Geol. 45: 168-197.

Hoffmann-J., A.E. (1989). Cactáceas en la flora silvestre de Chile. Ediciones Fundación Claudio Gay, Santiago. 272 pp.

Hoffmann-J., A.E. (1991). Paposo: aguas cristalinas. Eco-Ambiente, El Mercurio (Santiago) 31/3/91.

Jaffuel, P.F. (1936). Excursiones botánicas a los alrededores de Tocopilla. Rev. Chilena Hist. Nat. 40: 265-274.

Johnston, I.M. (1929). Papers on the flora of northern Chile. Contr. Gray Herb. 85: 1-171.

Johnston, I.M. (1936). A study of the Nolanaceae. Contr. Gray Herb. 112: 1-83.

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Authors

This Data Sheet was written by Dr Michael O. Dillon (Field Museum of Natural History, Center for Evolutionary and Environmental Biology, Department of Botany, Chicago, IL 60605- 2496, U.S.A.) and Adriana E. Hoffmann-J. (Fundación Claudio Gay, Alvaro Casanova 613 - Peñalolén,

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