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Biotopes / Ecosystems Nomenclature

Habitats of South America

 Report                                                    May 1996

INSTITUTE OF TERRESTRIAL ECOLOGY

INSTITUT ROYAL DES SCIENCES NATURELLES DE BELGIQUE



Habitats of South America

9

This classification is part of a report to the Commission of the European Communities, Directorate-



General for Environnement Nuclear Safety and Civil Protection, prepared by the Institut Royal des

Sciences Naturelles and the Institute of Terrestrial Ecology with the collaboration of Peter Kunkel,

Charles Vander Linden, Roseline C. Beudels, Marie-des-Neiges van der Elst, René-Marie Lafontaine,

Susan Iremonger, Marie-Odile Beudels, Yves Laurent, Chris Kerwyn, Dorian Moss, Wendi Craig and

Anne Devillers. 

Front cover. Upper Paranean riverine forest (unit 4A.3 1). Waterfalls (unit 24.141). Iguazu, Argentina, August, 1995. Photo, Anne S. Devillers.



Biotopes/Ecosystems/Nomenclature

10

INTRODUCTION



Scope and geographical coverage 

This extract of the PHYSIS database of the Institut Royal des Sciences Naturelles de Belgique offers a framework

to organize in an ordered sequence, according to the same methodology as that used for the CORINE Biotopes

Habitats of the European Cominunity typology (Devillers et al., 1991), the habitats of the South American realm.

The geographical area covered encompasses South America and its nearshore islands, Trinidad and Tobago,

Margarita, Aruba, Curaçao, Bonaire, Los Roques, Blanquilla and associated islands, the Galapagos Islands,

Malpelo Island, the Juan Fernandes Islands, the San Felix archipelago, Fernando de Noronha, South Trinidad and

the Martin Vas islands, St. Peter and St. Paul rocks, Easter Island and Sala y Gomez. 

Definitions and structure 

1. Habitat.

In the most common usage, a habitat is "the natural home of an animal or plant" (Collins English Dictionary), "the

normal abode or locality of an animal or plant" (Chambers Twentieth Century Dictionary), or "the natural abode

of a plant or animal" (Webster's New Collegiate Dictionary). Integrating these definitions over all species, for each

of which the habitat is the sum of the abiotic enviromnent and of all other species present, a habitat can be defined

as "a topographical expanse homogeneous in its physical and biotic components at the scale of the phenomenon

studied" (Blondel, 1979, 1995). This is the defînition that has been adhered to in the CORINE Habitat typology

and the Physis data-base. Thus, a habitat is a three-dimensional spatial entity that comprises at least one interface

between air, water and ground spaces, it includes both the physical environment and the communities of plants and

animals that occupy it, it is a fractal entity in that its définition depends on the scale at which it is considered. The

conditions encountered by a given species are similar on the entire surface of the habitat, but not all characteristics

of the habitat are ever assembled at any one point (Blondel, 1979, 1995). 

2. Scale. 

The définition of a habitat depends entirely on the scale at which it is considered (Blondel, 1979, 1995). Thus a

steppe can be defined, as in the Physis Database, as a "formation dominated by medimn or tall perennial tuft-

forming grasses or suffrutescents, with lacunar ground cover, together with its associated therophyte

communities". At a higher level of resolution it can be seen as a mosaic of habitats, bunches of tail perennial

grasses on the one hand, bare surfaces temporarily supporting communities of therophytes on the other hand. At a

lower level of resolution, the grass steppe and the steppe woods that dot it can be regarded as a single habitat, the

wooded steppe. The level of resolution that has been used in the CORINE typology is that of the ecological

requirements of small vertebrates, large invertebrates and vascular plants. A few units, clearly labelled, have been

introduced to permit rendition by the use of single codes or combined codes of the ecological requirements of

larger organisme or of the most mobile ones among the smaller ones. 

3. Habitat typology unit. 

A unit in the CORINE habitat typology and the Physis database is a habitat type, thus a characterization of a

collection of spatial entities that are considered equivalent as habitats, spacially separated but sufficiently alike in

abiotic conditions, physiognomy, composition of plant and animal communities. To play similar roles from the

point of view of nature conservation. 

4. Limits of units. 

There is an unavoidable degree of arbitrariness in deciding whether two communities are sufficiently

similar to be referred to the same unit, or conversely, sufficiently different to warrant distinctive

treatment. In that respect, habitat classification does not differ from any other taxonomic process. The


Habitats of South America

9

difficulty is only slightly increased by the fact that intrinsic criteria, such as reproductive isolation can



never be invoked to separate communities. The guideline followed in the CORINE typology was that

two habitats should be distinguished if the plant or animal communities they support were sufficiently

distinct to confer to them different significances in the préservation of sensitive species. For plant

communities, phytosociological criteria, recorded by the Braun-Blanquet approach, or any equivalent

one, were used, where usuary available, to assess degree of divergence and its relevance to sensitive

species. For animal communities, data are often lacking precisely for the groups most in need of habitat

conservation rather than species-specific programmes. Physiognomy, plant dominance, ecological

conditions and biogeographical parameters, including geographical séparation, have been used to

assess distinctness. 

5. Hierarchical arrangement. 

All habitat classifications use, alone or in combination, similarities in physiognomy, abiotic conditions,

plant community composition. plant dominance, plant community succession and, sometimes, animal

community composition to combine elementary units into collective entities of successively higher

rank (Dierschke, 1994). Contrary to classical species taxonomy, habitat classification cannot claim to

include, arnong others, a "natural" system of ordering based on the best available perception of

phylogenesis, thus on the history of the evolutionary process. Thus, the priority given to the various

criteria and the ensuing classifications are necessarily a rnatter of choice. The guideline of the

CORINE habitat classification bas been to use, 1, large- feature physiognomy, 2, plant-community

composition and 3, biogeographical or ecological factors underpinning animal-community

composition, in that order. This principle has, however, been occasionally departed front, when

necessary, to ensure maximum compatibility with existing local schemes. Sequence and level within a

division carry no intrinsic meaning as to conservation significance or affinities with other units of

equal rank. Only the series of nested ensembles of successively higher ranks is an indication of

cladistic relationship, the rank of individuel units derives from varying needs for finer subdivision, and

their sequence from the historical growth of the list. 

Rules of evolution 

Legitimate steady-state evolution comprises the amendments that will naturally occur in the course of

time as understanding of the habitats of the geographical area improves, as new needs of identification

arise, as an increasing portion of the total geographical area concerned is examined in detail. These

amendments should not, and do not, affect previous applications of the list. They are limited to two

opérations: subdivision of units, improvement of unit description. 

1.  Subdivision of units

It is the opération by which a fonnerly undivided unit is partitioned into a number of sub-units the sum of which

equals the original unit. It does not affect the content of the unit nor its geographical distribution although it is, of

course, possible that some of the sub-units will occur only in part of the area occupied by the undivided unit.

Exactly as in the case of the taxonomic splitting of a species, any legal status that was attached to the undivided

unit is automatically transferred to all its sub-units. 

2. Improvement of unit description 

It is the operation by which the description of a unit is completed or clarified, with, as appropriate, a more accurate

description of its geographical distribution, its ecology, a more complete list of characteristic species, a more

adequate title. It does not in any way affect the content of the unit. 



Global framework 

The South American habitat classification fits within a global system of habitat description which constitutes the

framework of the PHYSIS database (Devillers and Devillers-Terschuren, 1993). This system rests on the matrix-

use of two existing sets of upper category describers, the biotic realms of the I.U.C.N. bio-genetic reserve network

system (Udvardy, 1975), on the one hand (Table 1), and a list of upper units of habitats derived from the two-digit

CORINE categories (Wyatt et al., 1982; Devillers et al, 1991) on the other hand (Table 2). 



Biotopes/Ecosystems/Nomenclature

12

For realms, the geographical definitions of Udvardy have been adhered to as much as possible. Fer practical



reasons, however, they have been modified somewhat ta make limits of realms coincide either with marine gaps or

with political boundaries. Thus, in the arrangement proposed, no political boundaries have been crossed except

those that separate France, the United Kingdom, Australia and the United States of America from some of their

overseas territories. Precise definitions of the realms are explicited by Devillers and Devillers- Terschuren (1993:

13-14). 

The higher habitat units are directly derived from those proposed by Wyatt et al. (1982) as revised in the course of

the CORINE Biotope project (Devillers [et al.], 1992). Their extension to global applicability has largely drawn

upon the ecological analyses of Walter (1979) and Walter and Breckle (1986, 1991a, b, c), upon the

characterisation of major plant formations of the world by Rieley and Page (1990) and, for marine habitats, upon

the synthesis of Barnes and Hughes (1988). The definitions are summarized by Devillers and Devillers-

Terschuren (1993: 15-24), and can be found under the appropriate codes in the South American PHYSIS database. 

Upper units of habitat within any realm are designated by combination of a realm digit in the third rank to the left

of the decimal point with a biotope class number of two digits in the second and first ranks. Thus, Palaearctic

temperate conifer forests are designated by 042 (42), North American ones by 142, South American ones by 742.

Lower divisions, characterized by digits to the right of the decimal point, are specific to each realm and not

necessarily homologous between units formed with the same second and first rank digits, but different first rank

digits. Thus, Palaearctic juniper and cypress woods are coded as 42.A, with 42.Al, for instance, corresponding to

northern Hellenic Grecian juniper woods, while in an experimental draft prepared for North America, Juniperus



osteosperma-Juniperus monosperma-Juniperus deppeana-Cupressus arizonica woodland of central Arizona is

coded as B42.517, as part of piñon-juniper woodlands (142.51) in south-western summer drought forests (142).

There is, however, no reason not to preserve the lower unit hierarchy as far down as possible for types of habitat

with more evident cosmopolitan homologies. Thus, coral reef communities in the African realm can be expressed

as subdivisions of 211.256 as those of the Palaearctic realm are of 11.256. 

Table 1. Biotic realms. 

0.  Palaearctie

1.  Nearctic 

2.  Afrotropical 

3.  Indomalayan

4.  Oceanian 

5.  Austratian

6.  Antarctic 

7.  Neotropical 

(from Udvardy, 1975) 

Table 2. Biotope classes. 



1

Coastal and halophytic communities

11

Ocean and seas, marine communities



12

Sea inlets and coastal features

13

Estuaries and tidal rivers



14

Mud flats and sand flats 

15

Salt marshes, salt steppes, salt scrubs, salt forests



16

Coastal sand dunes and sand beaches

17

Shingle beaches 



18

Sea-cliffs and rocky shores 

19

Islets, rock stacks, reefs, banks, shoals



1A

Coastal agrosystems 



2

Non-marine waters

21

Coastal lagoons 



Habitats of South America

9

22



Standing fresh water 

23

Standing brackish and salt water



24

Running water 



3

Scrub and grassland 

31

Temperate heath and srub



32

Sclerophyllous scrub

33

Phrygana 



34

Steppes and dry calcareous grasslands

35

Dry siliceous grasslands 



36

Alpine and subalpine grasslands 

37

Humid grasslands and tall herb communities 



38

Mesophile grasslands

39

Tundra 


3A

Tropical grasslands

3B

Tropical shrublands 



3C

Tropical alpine communities 



4

Forests 

41

Temperate broad-leaved deciduous forests



42

Temperate coniferous forests

43

Temperate mixed forests 



44

Temperate riverine and swamp forests and brush

45

Temperate broad-leaved evergreen forests



46

Evergreen rain forests 

47

Semi-evergreen rain forests



48

Monsoon forests 

49

Tropical montane forests



4A

Tropical swamp forests

4B

Dry tropical woodland 



4C

Mangrove forests and scrabs 



5

Bogs and marshes

51

Raised bogs 



52

Blanket bogs 

53

Water-fringe vegetation 



54

Fens, transition mires and spnngs 



6

Inland rocks, screes and sands

61

Screes 



62

Inland cliffs and exposed rocks

63

Eternal snow and ice



64

Inland sand-dunes

65

Caves 


66

Volcanic features 



7

Deserts

71

Polar deserts 



72

Continental deserts and semi-deserts

73

Subtropical deserts and semi-deserts



74

Cool coastal deserts 



8

Agricultural land and artificial landscapes

81

Improved grasslands



82

Crops 


83

Orchards, groves and tree plantations

84

Tree lines, hedges, rural mosaics



85

Urban parks and large gardens



Biotopes/Ecosystems/Nomenclature

14

86



Towns, villages, industriel sites

87

Fallow land, waste places 



88

Mines and underground passages 

89

Industrial lagoons and reservoirs, canals 



9

Wooded grasslands and scrubs

91

Parklands



92

Bocages


93

Wooded steppe

94

Wooded tundra 



95

Treeline ecotones

96

Savannas


97

Wooded deserts and semi-deserts 



Acknowledgements 

We are extremely grateful to R. Bonnewitz, I. Benoit, J.H. Brun, M. Canevari, P. Canevari, J.C. Chebez, T. Frisk,

R- Gajardo, L.O. Giusti, M. Grassi, O. Huber, A. Krapovickas, V. Loewe, M. Lugo, P. Marconi, A.M. Molina, E.

Peña,V. Quintanilla, D. Rodriguez, M. Rumboll, K. Schiaffino, R.Taubenschlag, K.Thelen, R.Vanni who provided

us with much of the documentation on which the typology is based and contributed to its preparation by their

advice and encouragement. The project was sponsored by the Commission of the European Communities,

Directorate-Gencral for Environnement Nuclear Safety and Civil Protection. The sustained interest of J.

Vasconcelos, R. Roy and M. Cornaert in habitat classification and description as a tool for biodiversity

conservation has been central to the development of the CORINE methodology. 

References 

Barnes, R. S. K. and R. N. Hughes. 1988. An introduction to marine ecology. Second edition. Oxford, Blackwell

Scientific Publications. 

Blondel, J. 1979. Biogéographie et écologie. Paris, New York, Masson. 

Blondel, J. 1995. Biogéographie. Approche écologique et évolutive. Paris, New York, Masson. 

Devillers, P. and J. Devillers-Terschuren, 1993. A classifîcation of Palacarctic habitats. T-PVS (94) 1. Srasbourg,

Council of Europe. 

Devillers, P., J. Devillers-Terschuren and J. -P. Ledant. 1991. Habitats of the European Community in B. Wyatt et



al, eds. CORINE Biotopes Manual. Luxembourg, Commission of the European Communities. 

Dierschke. H. 1994. Pflanzensoziologie. Stuttgart, Verlag Eugen Ulmer. 

Udvardy, M. D. F. 1975. A classification of the biogeographical provinces of the world. IUCN Occasional Paper

18: 1-48. 

Walter, H. 1979. Vegetation of the earth and ecological system of the geo-biosphere. Second edition. New York,

Springer Verlag. 

Walter H. and S.- W. Breckle. 1986. Ökologie der Erde 3. Spezielle Ökologie der Gemässigten und Arktischen

Zonen Euro-Nordasiens. Stuttgart, Gustav Fischer. 

Walter H. and S.- W. Breckle. 1991a. Ökologie der Erde 1. Grundlagen. Second edition. Stuttgart, Gustav Fischer. 


Habitats of South America

9

Waller H. and S.- W. Breckle. 1991b. Ökologie der Erde 2. Spezielle Ökologie der Tropen und Subtropen. Second



edition. Stuttgart, Gustav Fischer. 

Walter H. and S.- W. Breckle. 1991c. Ökologie der Erde 4. Geniässigte and Arktische Zonen ausserhalb Euro-

Nordasiens. Stuttgart, Gustav Fischer. 

Wyatt, B. K., D. Cabot, S. ten Houte de Lange, H. Koeppel, U. Pinborg, F. de Beaufort, J. P. Ribaut. 1982.

Biotopes of significance for nature conservation. XI/94183. Brussels, Commission of the European Communities. 


Habitats of South America

9

1 .       COASTAL AND HALOPHYTIC COMMUNITIES



Oceanic, inshore and offshore waters and their open-water and bottom

communities; marine communities of the littoral zone and of coastal

lagoons, bays, inlets, estuaries and tidal rivers; coastal and azonal interior

halophyte or gypsophyte communities of saltmarshes, salt steppes, salt

scrubs, and gypsum scrubs; coastal sand dunes, sand beaches, shingle

beaches, sea cliffs, rocky shores, coastal islets, rock stacks, reefs, banks

and shoals; characteristically coastal agrosystems incorporating

seminatural elements. 



Habitats of South America

11

11.       OCEAN AND SEAS, MARINE COMMUNITIES



Oceanic and continental shelf waters of the world ocean and its connected seas, their associated open-

water and bottom communities, and marine vascular vegetation beds; marine communities of the

littoral zone and of coastal lagoons. Included within the South American realm are waters of the

Pacific east of the East Pacific Rise and, in the south, of the Pacific-Antarctic Ridge, of the Atlantic

west of the Mid-Atlantic Ridge, of the Caribbean Sea between the continent and the mid-line with

islands included in the North American realm; in the south, the demarcation arbitrarily follows the

mid-line between lands included in the realm and into the Antarctic realm. 

(Murphy, 1936; Cabrera and Willink, 1980: 106-107; Barnes and Hughes, 1988; Tharp, 1992: 42-43).



11.1      Open marine waters  

Pelagic biocoenoses of the world ocean, its connected seas and coastal lagoons.  They can be

characterized by their planktonic communities and by the composition of their nektonic or surface-

feeding faunas of cephalopods, fish, sea mammals and seabirds. 

(Murphy, 1936; Cabrera and Willink, 1980: 106-107; Nelson, 1980; Löfgren, 1984; Lowe-McConnell,

1987: 226-232; Barnes and Hughes, 1988; Stanley, 1995: 13-14; Diegues et al., 1995: 71-72, 74-75;

Hurtado, 1995: 131-132, 135). 

11.11     Oceanic waters  

Waters beyond the continental shelf. 

(Murphy, 1936; Watson, 1966: vii-xvi; Lowe-McConnell, 1987: 229-232; Lüning and Asmus, 1991:

10-11; Santelices, 1991). 

11.111      Blue ocean waters  

Oceanic waters of the tropical and subtropical zones, north of the subtropical convergence, situated, on

an annual average, around 40°S latitude except in the Humboldt Current zone of the east Pacific where

it swings northwards to about 25°S; they are generally of low productivity and high species diversity. 

(Murphy, 1936; Koepcke, 1961: 106; Watson, 1966: vii-xvi; Nelson, 1980; Löfgren, 1984: 134-135;

Barnes and Hughes, 1988: 37-42; Lüning and Asmus, 1991: 10-11; Santelices, 1991; Stanley, 1995:

13-14; Diegues et al., 1995: 71-72, 74-75; Hurtado, 1995: 131-132, 135; Bleakley, 1995: 14-15). 

11.112      Cool ocean waters  

Oceanic waters of the subantarctic zone, south of the subtropical convergence, at around 40°S latitude,

30°S in the Humboldt Current zone of the east Pacific, and north of the Antarctic convergence. 

(Murphy, 1936; Watson, 1966: vii-xvi; Nelson, 1980; Löfgren, 1984: 134-135; Barnes and Hughes,

1988: 37-42; Lüning and Asmus, 1991: 10-11; Santelices, 1991; Diegues et al., 1995: 71-72, 74-75;

Hurtado, 1995: 131-132, 135). 

11.113      Ocean upwellings  

Oceanic areas over which the warmer surface water is displaced, allowing cooler water rich in nutrients

to rise to the surface. This occurs when water is driven away from a coastline by wind action, when

two contiguous water masses are moved apart, or when deep currents impinge on an obstacle such as a

mid-ocean ridge. Zones of upwellings are characterized by higher biological productivity and lower

diversity. Major upwelling zones, involving, at least in part, oceanic waters, occur along the west coast

of South America from about 44° S to the equator, under the influence of the Humboldt Current, along

the east coast of South America, under the influence of the Falkland Current between approximately

44° S and 34° S, and in the vicinity of Saint Paul's Rocks. 

(Murphy, 1936; Watson, 1966: xiv; Nelson, 1980; Löfgren, 1984: 134-135; Barnes and Hughes, 1988:

11-12, 38-39; Hurtado, 1995: 135; Diegues et al., 1995: 75). 


12

Biotopes/Ecosystems/Nomenclature

11.12     Shelf and slope waters  

Waters of the continental shelf, underwater plateau extending from the coast to a depth of about 100

fathoms, beyond which the continental slope falls steeply toward the ocean bottom. 

(Murphy, 1936; Watson, 1966: vii-xvi; Cabrera and Willink, 1980: 106-107; Nelson, 1980; Löfgren,

1984; Barnes and Hughes, 1988; Santelices, 1991;  Stanley, 1995: 13-14; Diegues et al., 1995: 71-72,

74-75; Hurtado, 1995: 131-132, 135). 

11.121      Inshore waters  

Waters within the strong influence of land masses, both in terms of physical parameters and of fauna,

often arbitrarily defined as waters less than 5 km from low water mark, waters between the continent

and islands or islets well in sight of shore, and any seas of depth less than 6 metres. They include the

inshore waters of the South American continent and its nearshore islands, of Trinidad and Tobago,

Margarita, Aruba, Curaçao, Bonaire, Los Roques, Blanquilla and associated islands, of the Galapagos

Islands, Malpelo Island, the Juan Fernandez Islands, the San Felix archipelago, Fernando de Noronha,

South Trinidad and the Martin Vaz islands, of Saint Paul's Rocks (Penedos de Sao Pedro e Sao Paulo),

of Easter Island and Sala y Gomez. 

(Murphy, 1936; Koepcke, 1961: 106-107; Watson, 1966: vii-xvi; Cabrera and Willink, 1980: 106-107;

Nelson, 1980; Löfgren, 1984; Barnes and Hughes, 1988; Lüning and Asmus, 1991; Santelices, 1991;

Tharp, 1992: 42-43; Stanley, 1995: 13-14; Diegues et al., 1995: 71-72, 74-75; Hurtado, 1995: 131-132,

134-135). 

11.122      Offshore waters  

The zone extending from the limits of inshore waters to the continental slope. 

(Murphy, 1936; Koepcke, 1961: 106-107; Watson, 1966: vii-xvi; Cabrera and Willink, 1980: 106-107;

Nelson, 1980; Löfgren, 1984; Barnes and Hughes, 1988; Lüning and Asmus, 1991; Santelices, 1991;

Stanley, 1995: 13-14; Diegues et al., 1995: 71-72, 74-75; Hurtado, 1995: 131-132, 135). 

11.1221       Subantarctic offshore waters  

Waters covering the continental shelf of South America in the subantarctic zone, extending northward,

in the Atlantic, to Golfo San Matias and, in the Pacific, to Chiloe Island. 

(Murphy, 1936; Cabrera and Willink, 1980: 106-107; Löfgren, 1984; Lüning and Asmus, 1991;

Santelices, 1991; Diegues et al., 1995: 71-72). 

11.1222       South subtropical offshore waters  

Waters covering the continental shelf of South America in the subtropical maritime zone, extending, in

the Atlantic, from Golfo San Matias north to about 25°S and, in the Pacific, from Chiloe Island north to

northern Peru. 

(Murphy, 1936; Löfgren, 1984; Lüning and Asmus, 1991; Santelices, 1991; Diegues et al., 1995: 71-

72; Hurtado, 1995: 131-132). 

11.12221        Subtropical South Atlantic offshore waters  

Offshore waters of the subtropical zone of the Atlantic. 

(Murphy, 1936; Cabrera and Willink, 1980: 106-107; Lüning and Asmus, 1991; Diegues et al., 1995:

71-72). 

11.12222        Subtropical South Pacific offshore waters  

Offshore waters of the subtropical zone of the Pacific. 

(Murphy, 1936; Cabrera and Willink, 1980: 107; Lüning and Asmus, 1991; Santelices, 1991; Hurtado,

1995: 131-132). 

11.1223       Tropical offshore waters  

Waters covering the continental shelf of South America in the tropical zone, extending southward, in

the Atlantic, to about 25°S and, in the Pacific, to Punta Parinas in northern Peru. 

(Murphy, 1936; Cabrera and Willink, 1980: 106; Löfgren, 1984; Lüning and Asmus, 1991; Santelices,

1991; Stanley, 1995: 13-14; Diegues et al., 1995: 71-72; Hurtado, 1995: 131-132). 

11.12231        Tropical Atlantic offshore waters  

Offshore waters of the tropical zone of the Atlantic and of the Caribbean Sea. 

(Murphy, 1936; Lüning and Asmus, 1991; Stanley, 1995: 13-14; Diegues et al., 1995: 71-72). 

11.12232        Tropical eastern Pacific offshore waters  



Habitats of South America

13

Offshore waters of the tropical zone of the Pacific. 



(Murphy, 1936; Lüning and Asmus, 1991; Santelices, 1991; Hurtado, 1995: 131-132). 

11.123      Continental slope  

Waters situated over the continental slope, the steep descent from the continental shelf to the ocean

bottom, an area where upwellings, water mixing or shearing and other anomalies often develop. 

11.124      Offshore and coastal upwellings  

Zones of the continental shelf where the warmer surface water is displaced, allowing cooler water rich

in nutrients to rise to the surface, often generating much increased biological productivity. 

(Murphy, 1936; Watson, 1966: xi; Lowe-McConnell, 1987: 226-229; Santelices, 1991; Diegues et al.,

1995: 75; Hurtado, 1995: 135). 

11.1241       Major offshore and coastal upwellings  

Coastal and offshore waters of the major upwelling areas of the world ocean. 

(Murphy, 1936; Santelices, 1991; Diegues et al., 1995: 75; Hurtado, 1995: 135). 

11.12411        Humboldt upwelling zone  

Waters of the continental shelf off the mediterranean, desert and tropical coasts of Pacific South

America subjected to intense upwellings under the influence of the Humboldt Current. A large number

of organisms are entirely or largely dependent on this high-nutrient zone, in particular, the fishes



Engraulis ringens,  Sarda chilensis,  Trachurus murphyi,  Ethmidium chilcae,  Austromenidia regia,

Neptomenus crassus and the birds Spheniscus humboldti,  Oceanodroma markhami,  Oceanodroma

tethys kelsalli,  Oceanodroma hornbyi,  Oceanites gracilis,  Pelecanoides garnotii,  Phalacrocorax

bougainvilliiPhalacrocorax gaimardiSula variegataPelecanus thagusLarosterna inca

(Murphy, 1936; Koepcke, 1961: 107-108; Löfgren, 1984: 134; Lowe-McConnell, 1987: 226-229;

Santelices, 1991; Hurtado, 1995: 135). 

11.12412        Patagonian upwelling zone  

Waters of the edge of the continental shelf off the Patagonian desert and Pampa coasts of Atlantic

South America, subjected to upwelling under the influence of the Falkland Current. Important

populations of marine birds and mammals, including outposts of Humboldian ones, feed in these

waters and breed on the adjacent coasts. Characteristic species include Phalacrocorax atriceps,



Phalacrocorax bougainvilliiPhalacrocorax gaimardi

(Murphy, 1936; Watson, 1966: xi; Löfgren, 1984: 134; Diegues et al., 1995: 75). 

11.1242       Minor offshore and coastal upwellings  

Waters of the continental shelf in which local upwelling conditions are generated by features of the sea

floor or the circulation of water masses. They are recorded, in particular, from the north coast of Isla

Margarita, in the Venezuelan Caribbean, from French Guiana and from the Cabo Frio region off the

coast of Brazil. 

(Watson, 1966: xi; Lowe-McConnell, 1987: 229; Dawes et al., 1991: 215; Tostain et al., 1992: 12;

Diegues et al., 1995: 75). 

11.125      Shoals  

Shallow waters over permanently submerged elevated features of the continental shelf sea-floor,

whether shoals, banks or reefs. 


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