Mountain Recrch and Dadopmant, Vol. 1, No.2, 1981, pp. 113-184 GEOECOLOGY OF THE ANDES THE NATURAL SCIENCE BASIS FOR RESEARCH PLANNING Bouaxco Gostex Mou! ao Apniene V. Larsts? INTRODUCTION “The term "mountain geoecology” wae fist introduced by Carl Troll, 4 geographer who devoted trl of his ie to the study of the mountainous areas of the world and thote of Latin America im particular. Tt war while stad ing air photography and phocograrometry in 1938 that he developed the idea of *Landschaltshologie” (landacape cology), by which he meant a comprehensive physica geography. He later coined the word “gececolagy” ax a Syoonym ‘er "Lancschaftsokologe,” and defined i to in ‘ude the various branches of geology, climatology, ped ag, psa orzapy, oy and bly (Tel, 1968) ‘Why not simply re the term ecology” which bse mach the sane meaning? Ives (1979) suggests that ecology is ‘often regarded as province ofthe biologists and, there fore, tends toward a neglect of the abjotie components of ccosystems, The abiotic components are an integral pai fof any environment, as are humman populations. Ives calls for'a more balanoed approach with a renewed emphasis jon the geosciences of geography and geokury and a greater concern by ccologits Jor the study of human populations, ‘THE ANDEAN REGION Overview ‘The Andes Mountsine exiend through 70 degrees of lasinude along the wesern margin of South Ameria, They compnse a section of the 15,000 kilometre New ‘World Cordillera, are 7.250 kin in length, and cover ‘continsous area of more than to milion kon". ‘The ‘Andean Cordillera extends from the coast of the Carib> bean Sea in Veneauela and Colombia ot about lastade TI°North to Tierra del Fiego at abot 59°Seuth. Hon. ‘ever, for the purposes of MAB Project 6, the south ‘eramost border ofthe Andean region as been defined by fn imaginary Line extending from Valdivia, Chile, t2 Noaquen, Argentina. This excludes the Andean areas ‘covered by forests of the Andean-Patagonic type, but in cludes the subtropical and dry terrain in Cie and Arges ‘una north of Latitude 40°South (Figue 1) “Atough the core ofthis region isthe high Cordileca de los Andes, # complete understanding af its cology ‘would be impossible without also considering certain con- tiguous lowland areas, These areas include the coastal plains of the Carihbean Sea and the Pacifie Ocean, large "Gente de Fenigia, Univerad Navional de Cordoba, Cordoba, Argentina valleys and basins connected with these plains, and those 15 of the Orinoco, Amazon, and Parang-La Plata vers which originate inthe foothill of the Andes. The downslope position makes them the recipients of number of natural processes such ay hydrological action, colluvial nd alluvial depestion, cold air eurzenis, and.ex: treme geopbysical events such as landhlides. The more Dronouneed the orogeaphic relief of an upland area, the ‘eater wil be the potential effect a neighbouring low Tacs. MAB Projec 6 also inches some South American smountzine not strictly inthe Andee. These ate ranges in Chile anc Argentina north of latace 40° and parallel 19 the Andean chain, such as the Sierras de Cordoba ‘Cereain characteristics of the Andes mountains make them unique among high-alitade arese of the Works ‘These include (1) Abrupt changes i altitude resulting in vertical succession of ecozones ranging Irom ran forest and ese at the lowest levels #0 mourn tandea and ultimately permanent stow and ice at she highest. hinghamto, NY 15909, USA,116 / Mouvearn Resrazca aso Devetornit (2) Am enormous range of lartude, making for consid- rable variations ia climate, sol, vegetation, and land ute through the region (8) The great longitudinal breadih of the Andes, par~ tically in Colombia and inthe central subregion, “Tae maximum vith (more than 700 km) occurs between latitudes 15 and 20°South atthe Andean once” in Bolivia, (This area i also one of the bighescin cerme of mean stad.) (1) Te above three factors combine to give a picture cof great massivenes. ‘The otal bull of the Andean hain is about Five million cubic Klometzes. (8) Such ruggedness sugges very Youthful moun tains. Tn consequence, soils are immature and ger cally undiffertotiated and biogeographical de Selopment is relatively recent © Finally. the noclrsouth bartier position of the ‘Andee places it perpendicalar tothe great atmos Pheric ‘and oceanie circulations, This creaes ‘considerable aacro- and micro-climatic differences between the eastern and western slopes and the adjacent lola AI those factors interact to preduce a very complex set of climates and, therefore, a complex disrbution of eco: zones, Most important are the effect of latitude, altitude, ‘and topography. ‘Before presenting more deailed information about our state of knowledge of the cology of the Ande, the three ‘nin subregions arecefine! xo that they may serve a8 re erence point for later discussion (Figure ). Noxritest Apes The predominant characteristics of that part of the Andes situated between coastal Venerucla ard northern Peru (Cajamarca are ity great diversity and its moeaic of ‘ecosystems The Northern Andes subregion is dis finguinhed from the est ofthe eegion by the higher rela tive burmidity and by the greater limatic symmetry be tween the esstem and western flanks of the range."The large high plateaus found in the Central Andes do not ‘This ecological diversity is dependent on the flloning variables: altitude, steepness of slope, precipitation and. zelative humidity, and synoptic climatology. The comb tation of these variables produces the broad range of diverse ecosystems in the Northean Andes which conver: ‘Scnally are grouped inthe fllowing ecazones aramo or Tierea helada (land of ‘ox 4,800 to 3,200 ‘Tierra fia (cool land) 31200 ro 2,200 m ‘Tierra templada ((emperate land) 2.200 to 1,000:m ‘Tuerracaliente (hot or warm 1,000t0 0m land) Purther distinctions are usually made within these main fecazones according to tilerencet in altitude, haridiy, and steepness of slope. Tn terms of human use of these fren, chee differences ate offen a important as the di ‘inetions between the main ecozores, ‘The oorthern Andean region is dissected by deep valleys running parallel to the main crest of the range and, therefore, the vertically artanged ecozonal belts ‘reupy areas of comparable size. Accordingly, human Secupttion snot concentrated within any one belt ait ‘onthe high plateaus above 3,800 min the Centzal Andes, ‘and has to adopt to several aliucinal bets At the upper levels 9 combination of high humidity and low tempera tore becomes the main limiting factor for human ceeu pation of the Northern Ander Consequently, the upper Timi of cultivation is abot 3,200 rm, which is 1,000 m lower than in the Central Andes. Below ths upper limit, agriculture And estle-raiting have beceme important in all coomines. Nevertheless, there are two ecosystem: Sich are more suitable for agriculture than the others lind which at present have the highest population density. (Onc of them compris the les mountainous parts of the “hieras frat” where potato and crop cultivation, dairy farming, and increasing’, vegetable farming are foond ‘The second is represented by those parts of the “tierras, templadae” which have moderaie humidity and excellent conditions for the ealtivation of cofle, size, and rut Cermnat Anos “The Central Andes, extending from Cajamarca, Peru, 0 Antofagasta in Chile and Catamarca (north of Belen} in Argentina, are characterized by a preponderance of Tange high plateaus above 3,500 tm variously called “pana” cr “atiplano "The puna vegetation of these high pletcaus ie a response to a short rainy season and low tempera tures, particaarly at night [is further influenced by the specific effets of solar radiation, sharp variations in tem: perature, low humidity, and low exygen pressure. The Plateaus ae surrounded and separated by higher moun fain ranges and volcanoes, ‘The weetor flake of there mountains and the coastal lowlarvs are cry, oshereas the humid eastern Hank is distinguished by a succession of vertical ecozones from the Zona Nevada rising up 10 5,800 mn to the topical forests in the Amazon andl Paranae {Us Plata basins. While the heights ofthese eeozones vary with latnde and slope aspect, the esstecn escarprocnt generally shows the following Zeladonehip between alt fde and vegetational bet “Zon Neva 6.8004, 800 m Puna 4800:3,500 m Goja de la montana (cloud forest) 3,300-2 000m ‘Yangae (humid mountain forests) 2/000-,000-m Selva (topical forests) #,000-0 ‘The high plateaus above $,900 m in Pera and Boivia tare the most densely populated areas ofthe Andes. Four million people (33 percent of the population) live in the Peruvian pimat snd 3.5 million (88 percent ofthe pope lation) in’ the Bolivian aliplanos, Some yalleys in the central and eastern anges of the Cental Andes aswell as fn the eastern topes are also densely populated. Conte- ‘quently, 50 percent of the population of Pert and 50 Percent of the population of Bolivia live in rmountain reat. Because of icity the western slopes have lowJFeuns 1. Map of South Ameccs with nethorn, coal and southern regions enti Sed ates ofthe Andes it 3,00 m or sbove118 / Moowrais Reseanen ano DrvRiopwnir levels of amin occupation, The coastal iver valleys were densely populated even in pre-Colombian times apd, be- cans of the recent rapid growth of sowe urban eenires, specially Lima, the population has increased consid ferably Less than 10 percent of the papuition of both ‘countries lives in the astern Towlands. ‘The Central Andes were the “heartlana” of the preColombiae indig- ‘enous eropires and even roday they represent the only rea in the whole Andean region where the indigenous population exceeds 80 percent ofthe toal High-pateau residents grow potatoes, chenopodia, Daeg, and other froscresistant"ezops. ‘Their sheep, llamas, and alpacas graze perennially even neat the scow line. The extreme upper limit of potato cukivation is 4.200 m, In pre-Colembian times, mest high plateau ‘communities posressed land fights and cultivated Iapd at lower elevations inthe areas with teraperate and tropical climates on the exstera slopes of the Andes. In more recent times, escarpment communities often utilized mul tiple-altitude bets, but this system has been largely re- placed by exchange between groups residing at different alstudes and by markets, The eastern border areas of the [Andes in Per and Bolivia day mere than ever. provide Uwopieal, subtropical, and temperate zone products for ‘markets in mountain areas. Sourtans Aves Although the Andean mouptains extend to the south- cernmost ip of South America (Tierra del Fuego}, forthe purposes af this volume the suthern border ofthe sub= Fegion is taken t9 be somewhat farther north. Te South- frm Andes begin ar the edge ofthe alplang in Anvof fasta and Caamarca and extend © another imaginary line drawn through Valdivia in Chile and: Neuquen in Argentina, The altitude of the mountain summits de ‘reaees from north to south, The extent of cistection by valleys is very high and there are no high plateaus. The area is characterized by a dry or seasonaly-dry subtrop- ‘al limace and also by an increased daily and seasonal temperature range, The later becomes more pronounced with increasing larituce. ‘The vegetation corresponds to the climate and the different alitainal levels. Relatively humid forest ecosystems occur only in che extra-Andean ‘coastal Cordillera it Ohile and the Argentinian Sierras de ‘Cordoba, ‘The xerapbilie forest covers those parts of the caster cxarpments which receive at Teast 400 mm of finfall, In the inner areat in the rubiropical climatic one, vegetation becomes exuemely scare, The limiting factor ig arcty in the lower parts and temperature in the bigher parts tie not surprising thet the inner areas of the Andean range lacking exploitable vegetation are almost devoid of hhuman occipation, ‘The only exceptions are some isolated mining districts which are nevertheless of great im- portance. They are totally dependent on an external food ‘upp, and water supply can avo be a great proslem, However, important human seulerpents ia this region are concentrated inthe lower sections of major valleys and fon the piedmont of both Manks ofthe Andes a8 well 38 in the lower parte of mountain chaine parallel € the main range. These setlements are located in irrigated areat where maize and fodder crops are produced and where Vegetable and fruit farming is important, Becrute of the avallbiliy of fodder crops and the exploitation of forests ‘nd grasslands as pastures, te rising of exile, mle, and theep is widespread. Important tourist. settlements and Infrastructures can alto he found inthe region, inching the extra: Aridean mountains GEOECOLOGY OF THE REGION Geoecology ie the study of interrelaionshipe bur, in ‘order te understand the basic components gcology, li ratology, biology and to organize the information and false ic use or eoference, the material inthis sation wil be categories! and discussed under separate head Inge; some overlapping i unaverdable. Grovocteat Hisrony ov nis ANDES “The Andean Cordillera and the Perw-Chile Trench are par of the “ring of fire," 2 great band of veleanic and ake activity which circles the Pacific Ocean. The Peru-Chile Trench is a deep trough in the ocean floor Which rune parallel to the mountain chain Team lainide {North 10 40°Sounth, This trench isthe ste of tectonie~ plate activity which provides nev evidence for how the ‘Andes mountains were formed, Orogeny’, the process of ‘mountain-huilding, is now known 10 be closely linked wrth the volcanic and seismic activity which secomparin tectonic-plate colisions ‘The lithosphere, the outer roc layer of the earth, i thought to consist of large tectonie plates which ean be set In motion by the upwelling of wokeanie magma along their ‘margins from deep within the earth. This spreading ‘motion moves the plates in one direction, causing the ‘kading edge to be: consymed (subducted) by bending down atthe point of contact with another plate, thereby forming a tench. Inthe Ander, the wo pltes invalved are the Nave Plate (an oceanic plate) and the South ‘American Plate (¢ continental plat). These two plates ‘come together just beyond the present coatline to forma the PeruChile Trench, ‘The subcuction of the Nazca plate is responsible forthe gcological history and present- fay orogeny ofthe Andes (James, 1973). "Phe eldest rocks found im the Andes today are sei= mentary rocks laid dewn ina grea? geosyncline (off what was then the coastline of South America) during the Palacozoic Era nbout 250-850 million years ago. Texts, these sedimentary beds may be found far inland in the ‘eastern ranges of the Andean Cordillera, At the end of the Palacoeoic Era and the beginning of the Mesozoic, the supercontinent Pangsea began to shor signs of breaking ‘up. This period of daquiet tet the plate-tectonic eyele inmotion. The edge of the South American continent ‘became highly anstable, causing the geonynctinal layers (0 bend and buckle upward, forming the eastern ranges. At the same time, magma invaded these beds, creating Dbatholiths anc vleanocs ‘Daring the Jurasie period of the Mesozoic (short 190 million years ayo), th lithosphere began to «zack along the continental edge and subduction of the Nazca Plate Doegan, forming the Peru-Cohile tench, Magma walling 0 the autface eae of the trench formed line of atahore vel- ‘canoes. By the beginning of the Cretaceous period. Arica fand South America had become separate continents and nother volcanic are formed east ofthe frst one. By 50 or 50 million years ago, voleanie activity reached a peak ond fencrmavs quantives of magmaa invaded the erst, form ing the great Aadean batholith and pushing eastward t9 further fold and lift the extern ranges The carly part of the Cenozoe Bra was a time of considerable erosion of both exstern snd western ranges, Tn the Central Andes, substantial quantities of sedi mentary debe (up t0 15 ken in thickness) filled the area ‘beeween the ranges to form the aliplano. During the Pix eae and Plstocene epoche of the Cenozoic 9 period of Uuplieoccarved which formed the mountains which exist today. The surface, which had heen eroded (9 near 9a level, was that up above 4,800 ma with moreinocks up ‘6.550 m. Fekding and thrustfuuting created large bane A vast ake (known at Lake Ballivin) was created during the Loner Pleistocene and then drained during Final upift of the Andes leaving Lake ‘Triicac sr ste presenteday remnant (James, 1973; Winterbalder and ‘Thomas, 1978), Gopmonpntotoey axo Toocearay “The reat diversity of reel in the Andean Region torlay in result of other processes in alition o tectonic plate motion and its revultant orogente proceses. Two re lated geological forees waich conteibuce to the landscape are volcanic activity and earthquakes, The igneous activ ‘ny whichis amociated with mountain building continues ‘uninterrupted today, with volcanoes, bot active and ex tunet, tucéing the mountain chain from Tolima in the nnosth (Colombia) w Velcan Corcovado in the south (Chile), Some of the highes peaks inthe Andes are va: Canoes, such as Cocopani (5.696 m) in Ecuador, Tecora (6,050 ma) in Peru, and Sajama (6,520 mm) in Bolivia. South America boasts the worlds highest volcanoes in three categories: possibly the highest sxifw: volcace ‘Aconcagua, about 7,000 m, in Argentina, 2ls che high ‘it peak inthe Western Henaphere), the highest dormant Nolzano (Liullailaco, 6,750 m, on the Chile- Argentina border], and the highest acter volcano in the work! (Gsually considered ro be Antoalla, 6,440 m, in Argen tina, but more accurately Guayatin, 6,060 m, in Chile, ‘hich erupted in 1950) TThe Andean region, cated near a tectonieplare edge, isthus subject ra vulesnism and earthquakes. In addltion. cearthquakes are eequeatly the cause of other distuptive imatural ccents such ae landslides, avalanches, and Needs, although these phenomena may’ also have other causes. E.G. Mouna ano A.V, Lirtte / 119 [An interesting comparizon may be made between 10 ‘earth tremors whic stuck the Annes recent peat in May 1960 an earthquake registering 8.3 on the Richter ‘Scale hit southern Chile, causing abot 3,000 deaths and ‘considerable destruction in Valdivia and Puerco Monte, ten yesre later in northern Pera, an earthquake of lescer smagnitude (77 Richter) caused moce than 66,000 dearhs land destroyed the tonne of Yungay anc Raneahires and the city of Huaraz. The reason for this greater loss of life land property ws tha the 1970 tremor touched of a im tense avalanche which inundated everything in is path (Rebinson, 1967; Romero, 1072; Delury, 1977; We fl, 1972), Topography: Gens ‘An overall piture of the variations in relict in dhe ‘Andean chain e shown in Figure 2. The Andes are nar rowest and lonest at the southern end and widest in the ‘central ection (Bolivia). A second broad area is at the northern end (Collmbia and Venezucla) where the Andes fan out into four distinct ranges. Generally, the trend of the cordlleras is north-south, but_at abou: labeude TPSouth the Andes torn westwaed o form anes sem circular curve paral to the west coast. Tn southern Colombia the system divides into three distinct ranzes, the easternmost of which bifurrates, one branch trending north along the Colombia-Veneaucla border, the other fortheas toward the Caribbean Sea, The Pacific coastal plain extends along the entre seaboard and in Ecuador the western range rive ikea great wall abruptly from the broadened plain Detsiled examination of the Andean chai show that is made up of many interwoven ranges which enclowe high intermontane plateaus and valleys The conjunction of "seo oF more ranges i sae or knot: a mire of these Contribute & the constricted appearance Df certain pats ofthe Andes, especially in the Narthern subeegion opogaphy: Nene Andes Tn Veneeuca, the most northern and eastern por the Anis, the motntains rise gradually from the Dean Sea and extend on either side of Lake Maraceibo toward the Colombia border. These ranges are, in fact 4 structural continuation of the Ceribbean Ilancs, Two 9 Vernesels's mountain chaine join to form Colombia's «astern range (Cordillera Ocienial) while two other ranges "ise parallel to it. These three distinc chains, separated by the reat meridional valleys of Mogdalena and aia ‘Cauca, meet at the Nudo de Patt, relorming a8 to ranges Here, in Eevadr, the Andean vault ist ts na rower, about 150 to 200 km, and is dissected by several perpersdicular valleys. Iris mace tsp of two corileras and an inter Andean valley beeween, where the city of ‘Qh ie located. From the Rado de Paso, Ecuador's ridges, cronned with yoleanoes, branch southward to en close the Fenasorean Plateab, an area sted with act ‘wine basin (Marrero, 1964; Robineon, 1967), From a structural viewpoint, the Northern Andes show a high surfacetowolume ratio at a consequence of ex120 / Mountarn Reseanc ap DeveLopMent aia aT Mg a aie aa cure 2. Crsrsections through the Andes at several ltt. Lewivea, 1856: Marrero, 19:48; rol 1968) tensive sloping surfaces. A second! important feature is the carsity of level surfaces (ltiplanos) above 3,000 ma, Topography. Cetra Andes Ecuador's ranges converge at the Nudo de Loja, and continue inte Pere where Une ranges are Formed which ‘ome together ar the Nuc de Paseo. Here tee moun- tain chain again arise which focus atthe Nudo de Vilea ‘ota, Further south at about latnude 14°South the Andes diverge into two major ranges, enclosing the altplano of Pers, Bolivia, and northern Chile and Argentina. The altiplano is the vast treeless and undiseected tblsland ‘tbove 3,600 m which surrounds Lake Titcace and Lake Pope and which ip characterized by interior drainage, Wwinkswept Nas, salt marshes and saline lakes (Ogilvie, 1929; Sauer, 1950). The atiplano is extensively underlain by lava ows and Quaternary sedimenss. The Central ‘The Contour line isa 1,000 ra. (data form Bowman, 192448 Andes have a loner surface-e-volume ratio than the [Northern Andes due to the grest massiveness and high plateaus. In addition, there are a numberof glaciated cor- Silera and veleanoes exceeding 6,000 m, Tepegroply: The Ect Flank TTae eastern slopes of both the Northern and Central ‘Andes show a great natural unity which i largely una feted by subregional nuances. From the Venezuelan Anvdes 10 Bolivia, the eastern flank exhibite grest geo rorphologial and scologieal homogeneity. The immenee fox, consisting of schists intersected with bathotids and dlsected by deep valleys with humic-forest-covered ‘Slopes, exende aver 5.500 mo, Some ecosystems, sich a the soe nublads, or dvd forest, are almost continuous from Veneauela to Bolivia,Topography. Souter Andes “The southernmost portion ofthe altiplano extends into Chile and Argentina as the Puna Ge Atacama, a cold dry plateau dotted by salt Hats which are fed by small sear, Tin much of Chile and the bordering areas of Argentina the Andes form single, narrow cominant range. How fever, in the northwestern provinces of Argentina the Puna de Atacama is bordered by aeeriee of ranges which are functionally but not geclogieally related to the Andes. ‘The orogenic activity which prockcee! the main Andean, chain buckled these outlying ranges into & series of long parallel north-south blocks enclosing high narrow busine Regional seismic activity i intense inthis area Th Chile and along the western boundary of Argentina ‘single chein, deeply ditected by streams and lacing the level surfaces found in dhe General Andes includes the highen summit in the continent, Aconcagua, 7,000 rm ‘This dominant chain is separated from 2 loner coastal ange by the Valle Longitatinal, a long depression about 20 km wide which becomes « string of small basins be foweer Santings and Cogaimbo, Rewer latitudes 28° and 19°South no permarent stream reaches the ocean This lifelee coaeeal region isthe Aracama Desert. Herein the northern coastal area are also the grea nitrate basins (of Chile (Sauer, 1950; Robinson, 1967; Miler «ef 1977) Sonus Mowniins a Sot Gers “The process of sol formstion is contralled by: (1) char acter ofthe bedrock, (2) climate, (3) ising organisms, fie or degre of lope, and (5) the time or duration that sikforming processes have been preceeding. without ‘major duturbance (Retaer, 1974). Parent material in high raountain arcasis often high in voleanic, igneous, and metamorphic material. Climatic characteristics at high altitudes inclade low lemperatutes find high wind! speed which increase mechanical weather ing processes, Biotic communities within rvountaia fysteme are highly variable with “much local differ: Catiation: at very high elevations plant communities are limited by permanent frat and snow and ie cover. Relief brings into play the action of gravity and consequent ownward movement of eo, detritus, snd water Ane, since prolonged time is needed for the development of ‘mature soils, continual downslope movement and rapid soil depletion dhrough cresion prevent this maturation, ‘isthe variation in elevation and slope that principally Influence sol genesis over the meiniain landscape (Buck Hal, 1078-116). Variations in elevation relat to climate ‘ane variations in Slope relace to gravity; both have mani fol ett ‘The inverse relationship between temperature and ali tude produces active. weathering anc. 2oil-sorting. by Ieeze-than cycles atthe higher elevations (Corte, 1953 Fahey, 1973 and 1974; Mellor, 1979). Daiy feezing and thawing, at clevations where nighttime frovs occur ‘hroughout the year, produce “drnal slifhiction” lead E.G. Mousa ANDA.V. Livsit / 121 ing to the denudation of plant cover on sopes (Troll, 1968), Again, however, local landscape pattern influence conditions of removal and accumulation of deposits by wind aetion, drainage and erosion, ard vegetation action ‘on soil genesis (Buol taf, 1973117), Soll temperatures, ‘which relate to biological acivity and vegetation growth, are higher on north-facing slopes in the Southern Andes (Gouthern hemnsphere) and on south-facing slopes in the Rockies (northern hemisphere) (Buol «al, 1973:118) Pronounced! rie and steep slopes are characterized by intensive erosion, ereep of the regolith, and more dra rmatie processes sich af free Fall of terials, modsides, ‘and avalanches, Such events are localized and may alo be triggered) by seismic or volearic activity (Beek and Bramao, 1968). At Buol and his collexgucs {1973:119) rote, the age ofthe soil is decermine! largely by the sta bility of the surlace. Since marked relie! procuces un stable surfaces, mountain soils are usually young and shallow Ip gcocral, soils show distinct aldrudinal zonation from sexlevel to the level of permanent snow and ice (Bridges, 1978:90), Most, but pot all, mountain soils ae of low fer tlty, shallow "Lithosole, sremgiy acid, very stony, ‘medium to coare textured, and well-drained (Buringh, 1970:97; Reteee, 1974), Andeen Sole Soll description and classification in the Andes is par ticularly dificult because of extreme slitainal nd kat tudinal diversity. Hence, only scl systems in the broadest sense willbe deal with in this section, ‘Norihra And: ‘The humid climate of the Northern ‘Anges produces plan cover that ie limited not by water, but pncipally by low temperature conditions which are 9 function of elevation. Thee are, however, marty inet Andean. valley systems with sub-humid 1 semi-arid ‘microclimates (Beek and Bramao, 1968). The semi-arid valleys are characterized by Sodie, or salt containing, Solonet2 suis, while the sub-huonid valley systems ten toward the more mature Lzohumnic sls, with more rapid forganie master turnover, such as Braniaens abd Chest- rut sols. Stony Lithorols characterize the steep dopes hile Acid Brown sels are more represenative of yeatler Slopes. Atthe highest and teslss regions of the Nether ‘Annies, sile ate clasifed a0 Paramo humic or Andean hhumse sods (Cuatrecseat, 1968). These soil which sre highly fertit, are acidic, dark in colour, very deep and compact, and saturated with water, Pararyo sole are rich in organic matter and predominanily Andosls, devived from voleanie material high in vitreous elements, and Alpine Rankers, derived from crystalline rock (Dachau four, 1978-28). Cental Arcs: The western ranges of the Central Andes are relatively arid with deer like sole. Book and Bramao (1968) idemify darninant soil types as Regoans (imma ture and himestone-derived), Lichosos, Alluvial sil, and the calcimagnesian Sodic soils krown as Solonchak. ‘The ‘easter Central Andean ranges are relatively more huraid than the western ranges ane have mixed Brow Forest fils, Lithosole, ac heavy-textured Alluvial sol2 / Mourrain Reseanctt ano Devevorstent ‘The aniplano can be divided int the seasonally dry sorthern plateau surrounding laker Titicaca and Poops toa the semiarid southern plateau of Bolivia, Chi, and ‘Argentina, Much ofthe aca ofthe northern aliplaa t covered with the sediments of shrunken or exis, tnchiding the gret rch soln or alt fate (Bese and Bramao, 1968) Around Lake Tacaca Humic Gey sols are common; thve are hydromerpie in nature and are Characterized by temporary on permanent saturation tall pores (Duchauious, 1918108), which lead ttle ‘composition of erganic master de tox cil onygen de tit Socal such ss Solnet and Solonchak ae present in depressed areae. By far the mom prevalent sls are Chestnut soil. Rendeines (alpine meadow sol), and Hume and Stappe Chemozems (Prine tll), Winter fuer and Thome (1578.2) sent the mineral erlity Of coe of hove groupe ae good, since ite teaching has taken place, Tn the semiarid soulern antiplaro, si re devel from volcan ath with cart sandy topo Dunes, salt Bate, and Umestonedeived Rego pe dominate further oath "tm nds Beck ae Bramao (1968 10) point cout, “Much ofthe northern sector i thickly manted Br fayers of lean ash originating from along chain ot intermitendy ace vleanoes" Woleanic aah depos in this eeio ae qute deep even on relay sep does In the nomterm ane central areas of the Cental Range, Lihovls and frown Forest soe mixed with a voleanic jah rine are common, Further auth in he Gen Range, volanicderived Andosasof low fry, Litho sels alpine Ranken and Padls are the mgjr tall The western Andean foothill of Chile are charoter: ‘niealy madteranenn nelimate and sal ypet. Hyorotooy ive Drainage Sens The Andes serve to divide the South American cont: rent imo Atlantic and Pacific drainage systems. With a Few exceptions, rivers draining the eastern rangea and slopes have petennial lows and are long: those draining, the western ranges tthe Pacific ate shore and many are seasonal (Robinson, 1967). Most of the eatward rain- age of the Northern and Central Andes is via the three great river systems: Orinoco, Amazon, apd Parana-La Plata. Rastward drainage of the Southern Andes involves several river systems, westward drainage of the Andes, on the other hand, includes several hundred independent ‘Nester Ander: The Venenatlan Merida Cortilers hat three principal drainage systems: (1) the Tocayo River ‘lowing north to the Caribbean Seaj (2) the Chama and Mouatan Rivers draining into Lake Maracaibo; end (3) rnamersus rivers, inclcing the Uribante and the Becomo, which are a part of the Orinoce watershed. Eastward ‘drainage of the Colombian Andes ie divided between the Orinoce and Amazon watersheds. The Central Cordillera feeds the Cauca and Magdalena Rivers, whieh flow northwards to the Caribbean, whereas the Western Cor Ailsa drains into several rivers including the Sina, the ‘Atrato, and the Patia, Ecuador's Cordillera Real has east- ‘ward flow to the Amazon and westward flow toa number of Pacific civers including the Esmeralda in he north and the Guayas in the south (Cental Andes: Nortcen and central Peru show a typical ‘Amazon-Pacifc drainage pattern, There ate more than forty perennial or seasonal rivers that spill trom the ‘western flarks of the Cordilera Occidental: These rivers are used to irrigate the arid Pacific coast, producing fertile valleys where all major and minor population cent are located. ‘The central and western ranges are dieected By rivers such as the Maranon and Huallaga in the north, ‘and the Apurimae, Urubambs, and Mantaze inthe south wich Now northward into the Amazon. Further south, fon the Peruvian/Bolivian entiplano, all drainage i in ternal and feeds large highland lakes such a8 Titicaca and Poopo and saline marches auch athe Salar de Uyuni in Bolivia, The Bolivian slopes of the ansiplano are drained by the Pleomayo River, the Parana system, and tae Beni land Grardle Rivers ofthe Amazon system, Soutien Ander: Because the continent is aarcow in the south, the rivers flowing to the eait are considerably shortened. Rivers in northern Argentina lead ito th Parana, while southern Argentine Andean drainage oc- ‘curs through aseries of southwestowing streams sich as the Salaco, the Colorado, the Negro, and the Chubut rivers, Numerous rivers drain the western sloper of the Chilean Andes. In northern Chile, where the coastal anges reach the sea, many of the rivers are seasonal in low. In central and southern Chile rivers are perc and numerous. Stnding and Stornd Wate High-akitude lakes, snowfelds, and glaciers are ew porcan: as sources of tzeams and storage depots of fresh ‘water (Lafler, 1968; Osrem, 1974), Tee and snow can nod the landscape, transform sols influence the li ‘mate, and have a profound effect on living forms, particu Jatly humans Slaymaker, 1974; Mellor, 1979) Glacial lakes in the Andes often supply water for urban centers in nearby lealvies (Ogilvie, 1922:88). [mn the Northern and Central Andes, the lower limit of ‘permanent snow ranges from 4,600 m st the equator to ‘more than 5,000 m in the latitude of La Pax, Bolivia (Gchvwercceger, 1976). Very high lakes in the Andes ate found at elevations beeween 4,00) and 5,000 mi and many ‘of these were formed during the last retreat of glaciers ‘about 12,000 years age, and, as Laffer (1968) notes, ete sill indirect contact with the ie. The damming of mout= tain lakes by glacial ice or glacial moraines proces inste~ bility with often tragic results, Kinal (1968) cites eases of sudden discharges of glacial lakes leading to the lose of thousands of lives on numerous oetaionsin Per ‘With the exception of the very arid northern Chilean ‘oest end the southern altiplano, the water supply in the Andes is sufficient for present needs, However, popu lation pressure on existing water resources is now being {itn Bogota, which has the largest urban popillation in the Andes (Guhl, 1968). As urban cents, stich at Bogowe, Quito, Cuzeo, and La Paz, continue to grow,slacal lakes and melt-wate will become of iereasingim- portance. Certainly, the expanding need kr hyelroelet rie power will lead to more engineering works to exaloit ‘Andes sraterway resources CCuineare ane Weare (Climate variables, sich as temperature, precipitation, air pressure, solar radiation, and wind speed, are the “Geiving variable” thas structure and influence the eta lishment of ecorystorm. At a more fundamental level, ‘there are several global determinants of climate that mast be considered in any discussion of the climate and weather of a major geegraphie region. Thete are (1) fat ‘ue, (2) magnitude oF mast of lan area and proximity 0 water bodies, (3) atmospheric patterns, and (#) relief of the land macs (Rebineon, 1967). The Andes are charae= terized os (1) primarily tropical ond subtropical by la tule; (2) in lose proximity tthe Paciie Osean and the svestern margin of a large continent; (3) subject to the [Xmoepherie conditions of the trade winds and equatorial Interropical convergeace zone (ITCZ); ond (4) having fe of the highest and most massive mountain systems in the world (et, 1968), Matin Chats Theres in alte ae ascompanie ty the loverng of ir termpertar and huty sd incre nnd Specs rbd and lr raaton Thee ae ‘ated cara teas, lagater with vanatons i plat ier anil topoprapy, produce compl cal ane onal weather oni ‘Aldagh sree an inverse elatoneip bene em rate and skinid, there no enatant vue fer te {Emperor deine per anit eatin (mpc te) he stepet apc rats ocr daring he werent mone, Por ‘ape tne Alps ach 1000 tol elvan produce a 40°C drop in tempera dng January wheres in July a 1000 cae ge lpr of BPC (Geiger, To¢s-i), Win the eqonerial woe onthe Pera iglina, empeatregradints are versteep, a 10.9°C per 1,000"m dirng te cooler scaon and 25°C per T0'm ring te ware ene Lie an Hanoy 27 Senna cmperaute varton ks athe ce atone than tse eve, his tre for ped igh Iande and even more tin temperate selon ‘There ‘Rees nen om conte mat a level mative ype cana high astude, Seasonal ‘satin at high estos are mined a al ‘ations are aioe, Marked aaa verano in deserts, results rn high fv of ongwave ation that ocums ander doudfte Gaytine ses andthe rapid cacape of is radiation under dear nighctime ses {Eater dowdy ain, the una! trpertar tage wl decrease markedly. = = Tec variations in topography influence tempersture and hy, partial with veapect ure anges. Daytime temfernare valley ope pen gatens oe tree wih nereatingeevation.Atsig, weve thee iss tempernureinverton, where bh valley speak temperatures are low aad slope temperatre are fl E.G, Moumma ano AN Livre. / 123 tively high. The same general pattern forthe theoe relief features holds foe humidity, except tha hrmiity i maxi- smal at night when tempzratures are low. Therefore, valley floors, despite their relatively low elevation, tend to display the greatest diurnal temperature and humidity fuctuations Solar radiation, particulary short-wave radiation, ine crcaacs with altitude Becaune ofthe reduction in atmos phere oxygen, nitrogen, and ozone molecules hich absorb apd seatter radiation (Prohaske, 1970). Tem: perate highlands show most pronounced seasonal vars on in solar radiation whereas in tropical highlands sca sonal varction sles pronounced and more influenced by Goad caver, haze, and dust attocietod with crynese Ulteavicet radiation at elevations greater than 3,000 min the tropics i more than double that reccrded at aoa evel (Buctener, 1963), Andean Clinaes “The atmospheric circulation, which translers energy through the movernent of ai masees, contro the weather pattern ofthe Andes region. Pigueeilstrates air masa fonitiens ever South Ameriea during December and January (upper figure (a)-southern hemisphere samimce Solstice) and June and aly (lower figure (b)-eouthera hemisphere winter sobtice). The cominant atrspherie featurer that influence South American climate ore sev- eral high and low pressure “cells the intertopical con- vergence zone (ITCZ), and the trade winds which com Verge toward the TTCZ. The ITCZ. i 4 low-pressure trough which exchanges energy. through a vertcal coor ection cycle (Hadley cell, with the trade wands. The ‘equatorial postion of the TTOZ, contributes directly to the perennial hursdity of southern Colombia and northern Ecuador (Sick, 1969). ‘The seasonal «hit of the ITCZ northward i June and July and southward in December and January dominates othe seasonal rainfall patetns in the Central Andes Between July and January, the southern Pacific and southern Allantic high-pressure air masses shift south. ward leading to a corresponding southward movement of the molsture-den low pressure DCZ (Figure 3). This shift produces the Central Andean wee season i the high- Jands curing the menths of November to March when the ITCZ is ints southern position. The aridity of the Cen taal Andean costal one ie te result of mowncain rai shadow effect, the cold Pacific Humboldt current an the fauthern Pacific high-pressure cell. The. rain-shaow cic is produced by maature-besring ar from the sth erm Allantic high-pressure cll releasing it mitre a8 ascend) the eastern slopes of the Andes. As the already moisture-depleted air descends the western slopes, warn Ing ard the rie in pressure increate the moistire hiding. Capacity of the ai and precipitation is prevented A rain: thadow effect alone would not produce the extreme central coastal aricityrather, the position ofthe Him: Doldt current adds a further dessicating feature to the coast. AS the cok) Humboldt current fows northieard along the western coast, Uae cool-dey Pacific high-pressure aris cooled further at it erases the Humbolde current124 1 Mouweats Reseancit avn DeveLowent ‘icuRe 3. Atmospheric condition in Seuth America: (2) ding {he wuthern hroviphereeammer teiie (Oecersbe!Jaruary) (©) daring he southerm hemisphere winter solstice June!Ju), TTCZ isthe iteropical convergence ee: atroms ignily the direction ofr mate exerts heeled aren i he lowe figure i the aid sane ofthe coat, stiplana, and Patagonia, (Ghodiled om FAO/UNESCOIOMM, 1575; Gabel. 1947) and a very stable temperature inversion occurs in which Tain is almost eliminated (Eide, 1968). The £! Nino up- ‘weling current of warm water displaces the Humbold. ‘current every few years, and gives ee to heavy raine and flooding along the otherwise arid coast. A shift in the FECZ further south than normal has been saggested at ‘onc caus ofthe appcarance of BI Nite (Johnson, 1976). ‘Along the Chilean coat, south of the Atacama desert, the dimate is influenced by polar maritime circulation patterns, In thi area of central and southern Chile, polar fir mamees cross the coast and ascend the Andes with heavy precipitation resulting Prasrintion: Rainfall patterns ate iustrated for the Andes and adjacent coast in Figure +. Inthe north, pre “ipitation is wniformly high throughout the year both in the highlands and on the coast. Southwards trom central euador, at sbout the latitude of Guayaquil, ccaetl arid: ity develops and increases, culminating in the coaly-dry [tacara desert of northern Chile. Central ighland pre- ‘ipitaton at parallel latitudes sifs into a seasonal pattern of rainfall where the anna totale are about half those in the Northern Andes. The arid puna of Bolivia Chile, and ‘Argentina occurs where the arid 2one erostes the Andes (see also Figure X0))- The west-central coast of South [America ie an Andean rain-ehacow at precipitation fale fom the eastern slopes; further south in Patagonia, the ce verte oncone and the rainshadow Hes onthe eastern slope \Vaktvia, Chile, at about latiuale 40°South, is repre sentative of tie very wet region of the couthern Andes Coas,Partielany vieent thunderstorm ate common in ‘he highlands, especially the central highlands, ‘The snowing is given by latitude and altiude for the Andesin Figure §, Deviations of the height of permanent snour-felis from lines of freeving temperatures between Tatiudes 15° and 30°South are caused largely by ow pre tation. Precipitation i the form of snow or hail de- pends upon clevation, latitude, apd time of day. How: ver, below the permanent snovline in the Northern snd Cental Andes, snow cover sekdom persists into the late fernoon “because” daytime temperacures, even a 4,500 mi, are well above fcceing Tnprotire Latitude, elevation, and the Humboldt ceargent are imaportant determinants of Andcan climate Homever, latitude is only 4 minor factor in the Northern and Southern Andes, Temperature lapse rates seem 10 Tange from 1°C 10 6.5°C, but with much local variation (Johnson, 1976). ‘The Huroldr current exerts a marked Cooling ellect on the Central Andean coast and western ‘Andean slopes. Ta fae, temperatures may be warmer A 1,509 m on the western slopes than at sea level. This illus ‘ater 2 revertal of the normal late rate pattern Daily temperature means and ranges at four locations are listed in Table 1. When ald Se held ronghly con: ant latitude effects on mean annual temperature are ‘minimal’ Bogota, Quito, Cajamarca, and Arequipa, with ‘latitudinal spread of more than 20 degrees show a mean nnual temperature range of less than I°C. What do become apparent are the southward «ends of slight tere perature seasonality and increased diurnal variation (Table),‘i iGHLANOS see Wild _ “ie ws Uarirupe Frama Anal rainfall are) along lng gradient {rom TOMA to 40°S ter the Pale Cont ad the Anda High lind, Rain is expremed on log vale. Locations are. ‘Merida, Venezuela b. Medlin, Colombia; "Boga, Gaio> bia 4 Quito, Hrundor|« Cajarare, Ben f Caren, Peng [ee Peay Bolivia, h. Onur, Bava Cxint Redenor, Arsen tina: Salta Argentina: hos Mall, Argentina: Barras ‘quill, Colombia: m. Ciudad Muti, Colombia, a. Boens eonura, Colombia! 0. ‘Tumaco, Celombia: . Exmeralde. evador)q. Manta, Eeosdon; ©! Labite, Peru: + Chiclayo, Perot Limo, Peru. Arica, Chiles. Fgque, Chew. Anto- fags, Chile; . Valpmaieo, Chie; y. Coneepeon, Chie; 2 Valdivia, Che (data from Jonson, 1975; A Miler, 1976; Prakash, 1976 Stow, 1976) Frovne §, Summer and winter lit of feesing temperatures (OPC ines) and altos o permanent snawline (open Hgures). {(nodited tom Sebwerdeger 1976) E.G, Moura ano A.V. Liem. 125 Annual temperature values, however, can be mis leading, since there is considerable year-to-year variation Figure 6 ilutratce a {0-year monthly temperature record at site an the Peruvian aliplane. Seasonal varie tions this ste are principally the result of heavy cloud ‘over during the wet seaoon and clear skies during the dry featon (May-Angust). Ann variations in temperature ange ffor a bigh of 6.0°C in 1998 to a bor of 3.6 in 1951, Thie remarkable ameunt of temperature variation is plowed in Fogure 7. The years from 1950 to 1953 rub have been dificult ones for native farmers ‘Barooric and scygm pressor: No discussion on Andean climate would be compte witht comment on al tudinal effects on ar pressure. At altitude increases, baro- metric presire decreases at a constant, but not linear, fare, Ongen pressure, which is a constant Fraction of barometric presaue, decreases atthe same rate. Accord ingly, oxygen at 2.000 m elevation shows a pressure redvction of 20 percent below sea-level values. At about 5,400 m elevation, there isa 50 percent reduction in the svallable oxygen. The effects of reduced oxygen pressure Inthe Andes ie dixcused hy Lite, pages 152 160 Veoersrion “The disitatian of climate zones combined with al ‘de is the most important factor in determining the dis tribution of vegetation types Similar plant communities ‘nay be found in widely separated parts of South America ‘which have simular weather patterns, There are, however Other facore which influence vegetation which ae not dec related to ciate, notably sel type, slope, drain tage, and human interention (Robinson, 1967), Pant fifeforms show remarkable adaptations 0 the severe crvironments found in the Andean region, Tit Dnly the movt extreme conditions that tually provent plane growth such as the almost total dryness ofthe Ata tama Desert and the permanent cover of snow and gla ‘ers at higher alieades. ‘A map indicating the major vegetation types of the ‘Andes is shown in Figure 8. The term given in the cop tion ate largely those of Tro (958) and are intended ae descriptive labels. The vegetation bets 10 be described below are algo showa in Figure 9, which is schematic longitudinal section through the exstem and western cor illras “Te small inset map of South Ameria indicates @ line on either side of which the two sections are oeated. ‘This figure alio inckudes many smaller areas wo 1 merous to deseribe here, many of which raul from shel tered conditions or special local ciate factors, Figure is particularly useful for compating the symmetry or anymmetry of vegetation types to changes in moisture on citherside ofthe Andes Northern dnd "Mort of the Northern subregion can support a lush growth of vegetation on account of the high hurnidity ‘Tropical rain forests and other types of evergreen and decituous forests dominate this subregion with consid- table symmetey of vegetation types on the eastern and tweeten flanks. On the lowest slopes are the tropical rain126 / Moras Restanct aso DeveLorwent Tame 1 Temperate cate (°C) a four sites of rh uae aitad but dif latiude nthe Andes Bogota Quite Cajamaren Aceguipa Laude exon oss res ‘Alea, meter 2.888 2.318 aon _ Daily Day Daly, Daly, Dally, Daly, Dally, Daly Mert Mesn Range Mean Range _—Mean—_—Range Mea Range Jamary peo 0 ie 0 ir Be “ Febeary be 16 8 mo uw 19 a March Bro 28 5 Wo 4 as 4 pe ir 3 iso 2 #0 i ie 3 June Be 3 150 B ns 19 132 a July Re 8 ne 6 m3 9 1a ia ‘Aust Re ° i 1% 50 3 Ba 0 September Be oo 132 6 BS "7 Wa a Oeteher De 3 ns ir es 15 136 a November i > re rr Ko 18 139 December 13 19 rT 5 0 16 14a 16 Annual 13 6 Fiore 6, Forsy-year record of ‘mean monthly "temperatures (1931 16°7970) at Chaquibarn Tsar Tae teas Toes 1968 I9ae leq 988 TON) 1900 ll, Peru (3910 mm. 1443S luz) 'Valuce af the top of cach |2-month period are mean an- nual temperature, (rdied {rom Litle ard Hanna, 1978) Tear test toss tesa Tastee Test Taste toed is6)” 1962 Toes Ioea ives vee leer 1008 1969 TOMEAN ANNUAL ‘TEMPERATURE. 3019401980 YEAR 19801970 route 7. Fory-year trend in mean annul trperatres (1931 1970) at Grugutbarnbila, Pena Gnodied foe Late and Hann, 1978) forests which support agriculture year-round, producing bananas, mnie, eaeao, and other ere ‘Above the rain forest isthe cooler cloud forex, calle! variously the tera fia, sta rublaia, Neelwald, moss forest, égde montan; this (net erm sued primarily in eastern Peru and Bolivia, This belt consbes of dense ands of ferns, epiphytes, and bamboo, Beyond the tree= Tine isthe paramo bel: consisting of graalanl and herba- ceous plants (Sick, [959; Weber, 1969}, and above this is the swine, Cental Arce Here inthe mid-vection of the Andean Cordillera arig- ity begins o be a significant factor in vegetation pater ‘Along the coastal plain, dryness inereater southward 0 colina inthe Atacama Desert, one ofthe dries places fon earth. Except for riverine oates (where agricultare and. Targe population centres are now found), desert ‘yper of vegetation predominate. ‘Theve inelase thorn-bush and succulent desert, and loma vegetation, which is 2up porte! by the gare, 2 heavy mist which moves north long che coast of Pera (Tra, 958; Sick, 1950), The wast high plateas, or akiplano, exhibits what is called “pana” vegettion, but changes in humidity from north to South create three different puna types. Nowernmos!, but ting across the Central Anes from norihwes: to south= ‘est is the butnid or moise puna characterized by an n- incerrupted carpet of grassy plants. Next isthe dry puna swith the bunch grastes called wu (Ratacs app. and many ‘thers) and resis Tola shrubs dominant. Desert par, found in the Puna de Atacama of Chile and Argentina, has typical xerophytie planes such as cactus and thorny snubs (Troll, 1968). The puna ares Gemonetrstes most dramatically the “arid diagonal’ of yegcttion nese zones ‘extend inland rather than run pall tothe coast cue © ‘en inercase in dryness (rom both north to south and east “The eastern slopes of the Andes have a type of veRe- aiem simile t0 that of the cle forest of tra fo Ae Ficeas f Map af vegetation inthe Ant fom 4°N 0 57*S as, Vegetation bet and other symbol ar! rp ical aia fore (including montane forens); 2. opie ‘nbuopial semideciuaus fore (Tucumane-Baivian ‘cergrcen hgh mountain raland (Paral). 4 peal mountain meadows; 3 moss or homies pa dy puna bell, 7 thar and desert pana elt na de Atoms) 8 ceaal deer; 9. open sarin ‘within the ran forents: 10.alerphyllo woodland and Sheubled of central Chil, 1 thorn savanoe (rest fd gresdnd) inthe Fevadorian and Peron ees Tegions; 12. mist savannas (dcedvout fret and grat Ian) along the coat cegon of Ecuador 15 sah panne sole 14 atid areas, 1 camera mergin of tke Anders 16, westein boundary of Iygrophytie vegetation areas, (afer ral, 1968)128 J Mouwzats Reseancit xp Devevorwent seribed above. In the Central Andes this region i called ‘gi del moniaa eyebrow of the mountain, Southern Andes "This aubregion is largely in the temperote zone and temperatures decrease southward. More important for ‘vegetation ie a change in the st ie circulation which reverses the arity pater of the Gentral Andes; that is, incrensing dryness on the eastern eloper and in: creasing wetness on the coast “Along the coast, south of the Atacama Desert, is an aca of thorn serub, Atabout latitude 0°South, westerly ‘winds lead to an increase in moisture, In this area of central Chile there is cyclonic winter rain are sarmer Gry- ‘nes (mediterranean-type climate), which produces 4 sa tropical hard-leaf forest and yerubland. Extensive agricul ture in the Longitudinal Valley depends on irrigation (Sick, 1969) From noch fo south at higher ahitudes, ie the Puna de Atacama, detcribed above ax having desert puna vege tation, siceeded by an ares of mountain grateande snd mined forests. On the eastern slopes, dryness gradually inereases. In the northern part of the area, circulation patterns produce rain during the southern summer and, therefore, gratands and some mixrd forests. Southard, the vegetation gradually becomes scrubbier and more xerophytic Awineat Lire "The native fauna ofthe Andes is rich and diverse with innumerable vertebrate and invercebrate species adapted. to mountain ccorystems, Fittan (1969) identifies South ‘America at having the richest acimmal population of all continents. Certainly, the vas tropical forest contsibutes ‘uch to this faunal diversity; However, the importance of ‘Anan forms should not be underplayed (Mana, 1968). “The zoogeograptic divisione ofthe Andes are known 28 (1) the Patgents Chiles subregion, from northem Peru to ‘Tierra del Fuego (Ceotral and Southern Andes), and (2) the Gsiane-Braciie sabregion which includes highland ‘Venenuela, Colombi, and Ecvador (Northern Andes). Giltmore (1950) ist some ofthe characteristic fauna ofthe Ptagonia-Cilea subregion at: oposauins; some distinct armadillos; a number of kystcomorgh rodents (chin~ ‘marae, chinchlline rate, octodon rats, tuco-tuees); wild” and domestic cavies (guinea pig), hhuemal deer; will and domestic canmelide (veuna, guar reco, alpaca, llama); theas; ducks, geese, conders, and ‘many other species of birds. Facelent reviews can be Found! of Andean anhropods (Beach; Bicherl, Tes, Kuschel, Noodt; Schuster); of fish (Gery), of birds (Clrog) ard of mammal (Simpson) n the second velumne (Of Biogeography and Beslgy in South America (Pitas ea, 1966), SUBREGIONAL ECOLOGY Of al terrestrial compesite ecooyntems, the Andes are probably the most complex, since each altitudinal belt has ts own structure, both vertically and horizontally, anc plant and animal lie is incredibly diverse. [n general, the lack of inregestion among different environmental studies has resulted in partial and oversimplified approaches t0 Andean ecosystems. Such fragmented and oversimplified ‘ers have often led environmental mismanagement Multiple and intensive use of Andean environments is ievitable inthe future, Hence, the more thorough and comprehensive the knowledge of Andean environments is, the better the prospect for wise exploitation and long: term comservation of i natucal resources, nthe section that fllons, 2 combined subregional (Northern, Central land Southern Andes) and altudinal approach will be taken in order (0 deal with the onanifeld kinds of eco= “ystems found. Whereas the previous section dealt with environmental parasceters in a very general way, an al- tempt will be made here to. integrate environmental parameters and to provide more resolution and deal ‘The Noaruew Axons In this section, three raain ecozones axe discussed: (1) the cold pararao belt (nia blade above 3,C00 rm; (2) the Intermediate, temperate belt (ora fra) between 2,000 ma and 3.200 m: and (3) the low elevation, marta bels (ea Unnplada and tora cli) below about 7,200 rm. Each of these northern Andean ccoeones is characterized by high precipitation and humidity, minimal seasonality i tem perature and rainfall, and’ high primary plant produc tivity and species diversity. Cold Bzsyions: Parcs ‘Paramos may be divided into “High Paramo” (from 8,600 m te well above 4,000 m) and “Low Paramno" (oe- ‘ween 3,000 m and 4,000 m) ecazones (Figure 10}. ‘High Poramos- This econyetem is restricted to afew hu due squate kilometres that are scattered on the upper: most levels of the Costa Rican mountains and the Calor, bian, Venezuelan, and Ecuadorian Andes. Some [Parsimostike environments, however, are alto found in the ‘extrome northeastern part ofthe Peruvian Andes (Ease nuco, approximately laticude 8°South): this environ: ‘mental area is called “Jalea.” The high paramos are env ronments not gready anodified by human activities and ‘may be considered able” under present conditions. ‘Mean annual temperature ranges benween 1°C. and 17°C with almost no seasonal variation. Conversely, daily variation, although largely tempered by dhe high hur rmidity levels, may show a range a large at 25°C. (form -38C to 20°C) with a mean annual maximum of about 42°C, an absolute maxienom of 20°C, and an absolute ‘minimum of -8°C. Night and morning frosts occur on ‘more than 150 days a year. Precipitation is fairly well i tributed throughout the year, interspersed with two short, less humid periods: mean annual amounts range from