Palaeogeography, Palaeoclimatology, Palaeoecology, 80 ( 1990): 107-127

Elsevier Science Publishers B.V., Amsterdam

107

Ice age vegetation and climate of subtropical Chile

C. J. H e u s s e r

Department o]" Biology, New York University, 1009 Main Building, New York, N Y 10003 (U.S.A.)
(Received October 3, 1989; revised and accepted April 17, 1990)

ABSTRACT Heusser, C. J.. 1990. Ice age vegetation and climate of subtropical Chile. Palaeogeogr., Palaeoclimatol., Palaeoecol., 80: 107- 127. Vegetation and climate of subtropical Chile over the past > 50,000 yr are reconstructed from pollen, spores, and other microfossils contained in a 10.7-m section of lacustrine deposits of Laguna de Tagua Tagua (34~30'S, 71 10'W). Controlled by 14 radiocarbon age determinations, five pollen assemblages covering stratigraphic zones in the section represent plants on the upland and species inhabiting the lake basin. Surface pollen spectra from 58 stations (approximately 30-40S) form an adjunct for interpreting past vegetation and climate from the fossil record. Temperate, semi-humid woodland of southern beech (Nothofi~gus dombeyi type) and podocarp (Prumnopitys andina) was apparently established about Laguna de Tagua Tagua during the Pleistocene. Contrasting existing semi-arid, broad sclerophyllous vegetation, the woodland was extensive about the lake at the time of the last glacial maximum (25,000-14,000 yr B.P.) and earlier (> 43,000-34,000 yr B.P.). Pollen from the families Chenopodiaceae and Amaranthaceae, which characterize intervals of the Pleistocene (centered around 33,000 and >43,000 yr B.P.) and the Holocene, suggests intervening episodes of relative aridity. Lake level fluctuations appear to follow the late Quaternary climatic pattern implied by pollen assemblage data. Water in the lake was relatively high during development of southern beech-podocarp woodland, whereas during intervals of chenopod-amaranth dominance, lake levels were comparatively low. Excess precipitation over evaporation, which during the Pleistocene favored development of woodland of beech and podocarp, is attributed to greater storm frequency. Conditions were apparently governed by strengthening of atmospheric circulation in the belt of southern westerly winds. In the late-glacial, the temperate and more humid, ice age climate with limited seasonality underwent transition to a subtropical, semi-arid, Mediterranean type that identifies the postglacial. Fluctuations in intensity of the southern westerlies, marked by latitudinal movement of the polar front, can account for past vegetation changes at Laguna de Tagua Tagua.

Introduction
G l a c i a l d e p o s i t s in the A n d e s M o u n t a i n s , eraplaced d u r i n g the Pleistocene by w i d e s p r e a d a d v a n c e s o f glaciers ( C l a p p e r t o n , 1983), attest to ice-age climate in c o n t r a s t with the present. C l i m a t i c c o n d i t i o n s were a p p a r e n t l y c o l d e r a n d m u c h snowier t h a n they are t o d a y . A t 33S in the s u b t r o p i c a l A n d e s o f C h i l e - A r g e n t i n a , glaciers at present at altitudes o f > 4500 m h a d d e s c e n d e d to a r o u n d 2800 m d u r i n g the last g l a c i a t i o n (Caviedes a n d Paskoff, 1975; C o r t e a n d Espizt~a, 1981). P r e s e n t - d a y climate in the high c o r d i l l e r a at this l a t i t u d e (Cristo R e d e n t o r , 3250'S, 3829 m), by c o m p a r i s o n , is strikingly d r y with m e a n a n n u a l 0031-0182/90/$03.50 ;~:~,1990

p r e c i p i t a t i o n m e a s u r i n g only a b o u t 350 m m (Miller, 1976). F e w p a l a e o e c o l o g i c a l d a t a b e a r i n g on the clim a t e a n d v e g e t a t i o n o f the last glaciation are r e c o r d e d from s o u t h e r n S o u t h A m e r i c a (Heusser, 1987). A t higher latitudes, where glaciation c r e a t e d d e p o s i t i o n a l basins, m o s t records are f r a g m e n t a r y o r begin after late-glacial w a s t a g e o f the ice, as glaciers p u l l e d b a c k into the A n d e s M o u n t a i n s . O n l y rarely in t e m p e r a t e s o u t h e r n Chile, as on Isla G r a n d e de Chilo6 (42S), does a r e c o r d (Villagr~m, 1988) e m b r a c e the millennia o f m a x i m u m glaciation. C o m p a r a b l e sites for establishing the env i r o n m e n t a l setting at lower s u b t r o p i c a l latitudes d u r i n g the ice age are equally rare.

Elsevier Science Publishers B.V.

108 Laguna de Tagua Tagua, in the interior central valley of subtropical Chile, is the only regional source of continuous vegetational and climatic data from before the last glacial maximum. Its lacustrine deposits, with their macrofossil and microfossil content (Casamiquela et al., 1967; Varela, 1976; Heusser, 1983), provide a unique interpretive base for assessing the vegetation and ice age conditions not only at low altitude but also in the high cordillera. Its late-glacial paleo-Indian record (Montan6, 1968, 1969) is one of the earliest in southern South America. The purpose of this paper is to describe and interpret the fossil record of Laguna de Tagua Tagua, emphasizing its pollen and spore stratigraphy in the context of the past environmental setting.

c.J. HEUSSER evidence of drift-like deposits formed by mudflows, is now discounted (Segerstrom et al., 1964; MacPhail, 1973; Varela, 1976). During the latest of three glaciations, an advance in the neighborhood of Aconcagua (Fig.l) that reached as low as 2800 m during the last glacial maximum (Caviedes and Paskoff, 1975) is correlated with an advance of the same glacier system in Argentina, outwash from which is dated at 10,000 yr B.P. (Bengochea et al., 1987; Espizfla, 1989).

Climate
Climate of subtropical Chile is hot and dry in summer, when the region comes under the dominating control of the Pacific anticyclone. Influenced by frontal systems originating in the southern westerlies, it is cool and relatively wet in winter. Climatic data for stations at low altitudes (Table 1, Fig.l) indicate mean temperatures of approximately 18-21C in summer and around 8-12C in winter; total annual precipitation is between 100-800 mm with 84-90% falling during autumn and winter. Cool air, moving inland from the Pacific Ocean, orographically loses its moisture on crossing the Coastal Mountains, leaving the interior valleys relatively dry (Miller, 1976). Precipitation amounting to about 800 mm in the vicinity of Laguna de Tagua Tagua contrasts 1200 mm to the west of the Coastal Mountains and 2500 mm at this latitude on the Pacific slope of the Andes (Almeyda and S~tez, 1958). Fog and low stratus clouds are a common feature of the coastal sector in the northern part of the region. A southward trend of increasing summer precipitation is apparent in the data (Table 1), shown at the latitude of Concepci6n and at stations farther south (Fig. 1), where summers are increasingly wet. This condition is brought on by greater control of the polar maritime cyclone, resulting in an increase of storm frequency and Cloud cover throughout the year. At latitude 40S, annual precipitation increases to 1300-2500 mm at low altitudes (40005000 mm in the mountains) and mean temperatures fall to 15-18C in summer and 8-9C in winter. Temperatures between the latitudes of Santiago and Puerto Montt usually vary by only 1-2C

Subtropical Chile Physiography and glaciation

Subtropical Chile (Fig.l) lies roughly north of the latitude of Concepci6n (3650'S) and south of La Serena (2955'S). Topographic features inland from the Pacific Ocean are the Coastal Mountains, Central Valley, and the Andes Mountains. Consisting to a great extent of low hilly terrain of Mesozoic granitic and older metamorphic rocks (Brfiggen, 1950; Zeil, 1964), the Coastal Mountains rise northward in altitude from a few hundred to > 2000 m. Dissecting this upland and crossing the Central Valley, an intercordilleran depression underlain by Quaternary sediments, is a succession of drainage systems originating in the Andes. The Andes Mountains to the east, formed by Mesozoic and Cenozoic sediments and volcanics, are at altitudes > 2 0 0 0 m with summits between 32-34S rising to > 6000 m. These summits, where the firnline is at about 4500 m, are occupied by a number of small valley glaciers, which have been receding overall during recent centuries (Lliboutry, 1956; Corte and Espizfla, 1981; Su~trez, 1983; Cobos and Boninsegna, 1983; Espizfla, 1986). Glaciation of the subtropical Chilean Andes during the Pleistocene reached altitudes as low as 1200m (Santana-Aguilar, 1973); glaciation to lower altitudes in the Central Valley, based on the

ICE AGE VEGETATION AND CLIMATE OF SUBTROPICALCHILE

109

711 - 30 La 7~2

710 i--!

Seren~
:."

..j"

?
Pacific Ocean

- c , . . / C e r r o Aconcagua

-~--'~'. 7021
Santiago
r'

z~ --'i/i-' "/: '~' :/i--i

./:--:

Laguna de Tagua ~rnando

35
High andean beech f o r e s t ( > 3 7 ) & tundra ( u n d i f f e r e n t i a t e d )

-Conce

Subtropical thorn-shrub - succulent vegetation

I Subtropical broad sclerophylloua

__ woodland (matorral)

Val 40 ~]~

Deciduous beech f o r e s t

Evergreen rain f o r e s t

100

200 km

74

73

72

71

Fig.1. Map of subtropical (30 37S) and adjacent temperate Chile showing locations of Laguna de Tagua Tagua, physical features, plant formations (Schmithfisen, 1956), and places where meteorological data are available (Table 1).

during the rainy winters; in summer, differences amount to 3-5C (Table 1).

Vegetation
Regional natural vegetation of the interior valleys and low hills of subtropical Chile (Fig.l) is characterized by broad sclerophyllous woodland

or matorral (Schmithfisen, 1954, 1956, 1960: Oberdorfer, 1960; Mooney, 1977; Thrower and Bradbury, 1977; Armesto and Martinez, 1978). Supplanted by extensive farming in the valley bottoms, which were largely settled beginning in the 19th century, the woodland covers mountain slopes to a maximum altitude of 1600m. Its latitudinal and altitudinal relationships are with

110 TABLE I Climatic data for selected stations in subtropical and temperate Chile" Station Av. temperature

C. J. HEUSSER

(of)

Av. precipitation (mm) Seasonal distribution (%) Annual Spring Summer Autumn Winter

Summer

Winter

Subtropical
La Serena (2955'S) Santiago (3327'S) San Fernando (3435'S) 18.3 20.6 20.0 17.8 17.1 17.6 15.3 11.7 8.0 7.5 9.1 7.8 8.3 7.6 110 360 780 1338 2510 1330 1960 8 15 14 16 18 18 21 2 3 2 5 9 12 16 22 24 25 28 29 29 28 68 58 59 51 44 41 35

Temperate
Concepci6n (3650'S) Valdivia (3948'S) Osorno (4033'S) Pto. Montt (4128'S) "Almeyda and S~ez (1958).

semi-arid thorn shrub-succulent vegetation, established to the north, and deciduous beech forest, lying mostly to the south (Fig.2). Owing to precipitation and temperature gradients and a human disturbance factor, broad sclerophyllous woodland differentiates over the extent of the region. Communities typified by Cryptocarya alba (Lauraceae) contain, in addition, the arboreal species Schinus latifolius (Anacardiaceae) and Peumus boldus (Monimiaceae) and in cloud-covered parts of the coast, Beilschmiedia

miersii (Lauraceae). Communities in relatively dry locations are distinguished by Lithraea caustica (Anacardiaceae) and include, in addition to Cryptoearya and Peumus, the overstory species Quillaja saponaria and Kageneckia oblonga (Rosaceae). The tree Maytenus boaria (Celastraceae) is generally of minor importance but wide distribution in the Lithraea-dominated communities. A large sector of the dry interior between the Coastal Mountains and the Andes at the border of broad sclerophyllous woodland is occupied by

High Andean Beech Forest

Evergreen Rain Forest

Elevation rn. _ ~ 3000Subtropical High Andean

Vegetation

["~Nothofagus pumilio ~]

N.

dombeyi

N. dombeyi-N, alpina Saxegothaea conspicua-N, Eucryphia cordifolia-N,

dombeyi

dombeyi

2000-

Andean Tundra

1000- _ _

--Thorn S h r u b ~ ~ ~ " \. \\\\\\. Succulent f ~ N . ~.~ Vege!ation ~ ~ / / ' o ~ . ~ . / V .

~\ .~ - ~ ~/~. ~

~ ~

. \ ~ \ ~ I

m.

r 1-10oo

+':i2+;ii,-ii:ii+++ :i'~5i5575i:; :2+2~ ~'' '~ii~ ~+

320 3'30 3'40 i I 3'5 3'6 3'7 3'B 3'9 4'0 4'1
Broad Sclerophyllous Woodland (matorral) Laguna de Tagua Tagua Deciduous Beech Forest

Fig.2. Plant formations on the west slope of the Andes Mountains shown in relation to Laguna de Tagua Tagua. From Schmithfisen (1960) with some modification.

ICE AGE V E G E T A T I O N A N D C L I M A T E OF S U B T R O P I C A L C H I L E

111

steppe-scrub or espinal. Featured are the shrubs Trevoa trinervis (Rhamnaceae), Colliguaya odorifera (Euphorbiaceae), and Cestrum parqui (Solanaceae), and grasses (Nassella chilensis) and composites (Gutierrezia paniculata). Containing scattered flat-topped trees of Acacia caven (Mimosaceae), this vegetation has been much disturbed by man and grazing animals and is considered to be part of a successional series leading to woodland (Oberdorfer, 1960). Colonies of Nothofagus (Fagaceae), outliers of southern beech forest to the south, reach northern limits in the mountains (Fig.3). A transect made between the ocean and the Andes at 3450'S, close to Laguna de Tagua Tagua, shows the location of deciduous Nothofagus at altitudes above the limit of woodland: stands of N. glacua on summits of the Coastal Mountains at about 600 m and of
m.

N. obliqua between 800-1650 m on the west slope

of the Andes (Donoso, 1975). According to Rodriguez et al. (1983), the northern limit of N. obliqua is at about 33S on the upper slopes of Cerro La Campana and Cerro E1 Roble at altitudes between 1500 2200 m; N. glauca extends to about 34S (500m) and N. alpina to 35S (1000 m); and other species of deciduous beech, N. antarctica and N. pumilio, belonging to the high andean beech forest, range to 3515 ' 3530'S (1600-1800m). Evergreen beech (N. dombeyi) in the Andes reaches the northern end of its range close to 3430'S (600-1000 m). Three arboreal gymnosperms, also ranging equatorward from a more southerly distribution locus, occur at scattered stations along the subtropical andean front. Growing as far north as 35 45'S are two podocarps Prumnopitys andina

0 -

~ 0

~ ~ ~ , '~, -.

,, N o t h o f a g u s pumilio - N. a n t a r c t i c a N. d o m b e y i

20001000O-,

~.,,

e -1000

:,:::,::ii!::,::~,::',::::!::i%
33 ~ ~ o ~ ~ ~ 34 I 35
i

---'-'%, ,~40

n-

32
m.

36

37

38

39

41S. Lat.

L. de T a g u a T a g u a ~ o Nothofagus obliqua N. g l a u c a D N. alpina r ~ m.

30002000-

00

.....

~o

32

33

34

I
i

35

36

37

38

39

40

4 1 S . Lat.

L. de T a g u a T a g u a
m.

* P r u m n o p i t y s andina < >P o d c a r p u s satigna

, o o o -

1000

32

33

34 ]

35

36

37

38

39

40

41S. Lat.

L. de T a g u a T a g u a

Fig.3. Distribution of species of Nothofagus, Prumnopitys, and Podocarpuson the west slope of the Andes Mountains with reference to plant formations (see Fig.2) and the location of Laguna de Tagua Tagua. Data are from various sources including Reiche (1907), Donoso (1975, 1978), Rodriguez et al. (1983), and unpublished records.

112 (1000-1100 m) and Podocarpus saligna (1500m). The third gymnosperm Austocedrus chilensis (Cupressaceae) is established most equatorward near 3240'S (1700-2000 m).

C.J. HEUSSER volcanic and marine sedimentary rocks of Cretaceous age (Mapa Geol6gico de Chile, 1982). The shallow basin of the lake was formed during the late Tertiary-early Quaternary and extensively filled by laharic deposits from the Andes Mountains (Varela, 1976). Later aggradation by the Estero Zamorano, a tributary of the Rio Cachapoal, apparently contributed further toward impoundment of the lake. Laguna de Tagua Tagua before being drained in 1841 measured some 3 0 k m 2 in area with a maximum depth of 5 m (Varela, 1976). In the early 19th century, the lake was widely renowned for its unusual floating islands, consisting of interwoven rhizomes and stems of cattail and waterweeds (Reiche, 1907). As the wind blew, they carried cattle as passengers from shore to shore (Gay,

Laguna de Tagua Tagua

Location and physical setting
The site of Laguna de Tagua Tagua (3430'S, 7110'W) is a tectonic depression, lying just northwest of San Fernando and 120 km southwest of Santiago in the Province of O'Higgins (Figs. 1 and 4). It is situated at an altitude of 200 m in agricultural land at the eastern edge of the Coastal Mountains. Except for a narrow opening toward the east, the site lies within a circular ridge of

Fig.4. Setting of Laguna de Tagua Tagua. Based on Instituto Geogr~ificoMilitar quadrangles, San Vicentede Taguatagua (1984)and Chimbarongo (1984) drawn to a scale of 1:50,000, with altitudinal limits of the floor of the lake from the Laguna de Tagua Tagua (1930) quadrangle on a scale of 1:25,000.

ICE AGE VEGETATION AND CLIMATE OF SUBTROPICAL CHILE

1 13

1833; Darwin, 1958). Drainage was effected by construction of a canal between the north shore and Estero Zamorano (Fig.4). This was done in conjunction with digging a network of ditches across the lake floor, making the basin suitable for cultivation of crops.

Previous work: macrofossil remains and stratigraphy

Remains of extinct Pleistocene mammals, especially bones of mastodon, encountered during excavation work, pointed to the antiquity of the lake deposit (Wyman, 1855; Oliver, 1926). Most notable has been the recent discovery of mastodon bones in association with human artifacts and charcoal dated at 11,380 yr B.P. (Casamiquela et al., 1967; Montan6, 1968, 1969; Pino and Varela, 1977). Stratigraphic studies made of exposures along the drainage canal indicated a complex late Quaternary history of lacustrine deposition, interrupted at times by low lake levels and erosion, over an estimated time span of 57,000 years (Varela, 1976). The basal unit described by Varela (1976) is laharic, consisting of a matrix of volcanic ash containing fragments of andesitic rock and pumice stones, with a thickness of > 6 m. Sand and gravel (member 1, up to 2 m thick), overlie the deposit and underlie 10.58 m of lake sediments, beginning as greenish-gray clay, containing at levels molluscs (Tropicorbis) and other fossils (member 2, 10.58 7.25 m). Successively higher in the section are beds of yellowish clayey mud and some sand, intercalated by mud and by sand and gravel, with fossil deer (Antifer), each showing signs of erosion (member 3, 7.25-5.61 m; member 4, 5.61-4.49 m); greenish clay, also showing evidence of erosion, containing bones of birds, fish, and other vertebrates (member 5, 4.49-2.35m); dark clayey carbonaceous mud with paleo-Indian chipped tools, scrapers and flakers, among abundant remains of extinct mastodon (Mastodon), horse (Hippidion), deer (Hippocamelus), birds, frogs, fish, and rodents (member 6, 2.35-2.07 m), yellowish gray mud with diatoms, sponge spicules, ostracods, fish scales, and bones of frogs and birds, along with seeds and related plant material

(member 7, 2.07 1.04 m); and dark clayey, carbonrich paleosol charged with vertebrates and objects remaining from human occupation, projectile points, stone knives, and grinding stones (member 8, 1.04-0 m). Dates reported from levels in the upper beds (Montan& 1968, 1969) are 11,380+320 yr B.P. at 2.3m and 6130_+115 yr B.P. at 1.0m. Laguna de Tagua Tagua, according to the stratigraphic observations made by Varela (1976), was at low level during intervals at the beginning of the lacustrine record, during the deposition of member 4, following deposition of members 3, 4, and 5, and late in the record of member 7. Episodes of relatively high water are contained in members 2 and 5 and stands of moderately high water in member 7 and the upper part of member 8. Reference is the level of the lake prior to drainage.

Pollen and microfossil studies

Two sections of the lacustrine sediments of Laguna de Tagua Tagua for the study of fossil pollen and spores, one of 10.7 m and the other of 3.2 m, are from the central part of the lake (Fig.4). The location is in the northeastern sector of drained lake bed, where a farm road, beginning about 1200m distant on the upland, crosses a major drainage ditch. The 10.7-m section extends to the approximate time of drainage, which was the preserved bottom of the lake resting beneath sediments dug up during construction of the ditch. Sampling of the 3.2-m section was done later for the purpose of close-interval dating of upper levels. The location of the section is the margin of a cultivated field in proximity to the deeper section, as the original site of coring was no longer accessible. The upper 50cm of sediments, disturbed by cultivation, were discarded, Fourteen radiocarbon-dated levels show an average sedimentation rate of 51.4 yr cm 1. age-depth relations are indicated by the equation Y=0.18192+ 1.9550e-4x (Fig.5), giving an estimated basal age of the 10.7-m section of 53,800 yr B.P. Sedimentation over the length of record was not uniform, as may otherwise be implied by a straight line fit of the dated levels. Owing to past variations in lake volume and water depth, in particular, rates varied. To establish changes in rates of sedimentation, a larger data set is needed.

114
Depth
m, O-

C.J. HEUSSER

5-1

Av. sedimentation rate

5 1 . 4 yr cm - 1

m r- -

~"o .
\ %

10

10

2'0

30

40

5'0
14C y r B.P. X 10 3

5'0

Fig.5. Age-depth relations for the 10.7-msection from Lagunade Tagua Tagua. See Table 3 for data. Regional near-surface samples were collected for study in conjunction with the fossil pollen studies. They are broadly representative of plant communities existing over the latitudinal and altitudinal extent of subtropical Chile. Pollen in these samples is intended to portray vegetation prior to European settlement, when, for example, Pinus, Eucalyptus, Rumex, Plantago, and Hypochoeris were introduced. Many (but not all) of the sample stations are lakes or ponds, and their pollen spectra thus complement the fossil spectra in the lacustrine setting of Laguna de Tagua Tagua. Although these data are from an extensive region, wherein lakes are not consistently present, it is significant to note that lake and non-lake stations in proximity frequently record comparable proportions of leading pollen taxa. Section samples at 10-cm intervals and nearsurface samples were processed in the laboratory for their pollen and spore content (Heusser and Stock, 1984). Identifications are from modern reference collections and published accounts (Heusser, 1971; Markgraf and D'Antoni, 1978; Villagr~in, 1980; Wingenroth and Heusser, 1984). Among species of Nothofagus, N. dombeyi type (including N. dombeyi, N. antarctica, N. pumilio,

N. nitida) and N. obliqua type (including N. obliqua, N. alp&a, and N. glauca) are alone determinable. Frequencies (%) of tree, shrub, and herb taxa are from sums of at least 300 grains and of aquatics and vascular cryptogams from sums of >300 total pollen and spores. Plant nomenclature follows Marticorena and Quezada (1985), except where plant names are given in cited references.

Modern pollen spectra

Near-surface sampling stations 1-58 are located between approximately 30-40S (Fig.6, Table 2). Stations in the north (1-13), mostly in broad sclerophyllous woodland, are distinctive. They show high proportions of Gramineae (grasses), Chenopodiaceae-Amaranthaceae (chenopodsamaranths), and Tubulifiorae (composites) with variable amounts of woodland indicators, for example, Quillaja, Maytenus, Schinus, and Lithraea (Fig.7). Stations on Cerro E1 Roble (3, 4), the northern outlier of deciduous beech, clearly reveal this lacuna by a maximum of 36% of Nothofagus obliqua type; minor amounts of N. dombeyi type found there and elsewhere in the north are wind

ICE AGE V E G E T A T I O N A N D C L I M A T E O F S U B T R O P I C A L CHILE

[ 15

-30

I L a S e r e n a ~ o " - _ ---i : 'e fl 1 iJ /] [" I --/ i /

mlr,
,

-32

,,,s

[]

g
_ 3 4 o
"

2 '; ~ i

Subtropical thorn-shrub - succulent vegetation Subtropical broad sclerophyllous woodland (matorral) Deciduous beech forest Evergreen rain forest High andean beech forest ( =,37 ) & tundra (undifferentiated)

3,~4i
' i

[]

o Santiago

Laguna

de /

-'t'8~gi
u,,

e~5 o 7 ',, .;]

[]

Tagua Tegua[~] e 1 1 , ', j /

/ "4 ~

[]

i' ~ '~ "-~

_36 ," ,',',~ / C) .~)! ~"

Concepci6n : ~ []

-i
-3B

1 ~ / e2/14ct7 "t "~'s-,o

"

'~"

',....../ ,,I-~,
34-3g; ,2t-26 ~- ,,t. 4"+.:77-33
4 4 ;)e,s43 ,J Vsldlvla
.51-56 / ~ i

-40-

~ L
0 100 i

;7 "4'24f-47 ~s %49 ~
5 0 r)

45e"

~ / ~ 712':
200 i i 300km. p

7O B

Fig.6. Subtropicaland temperate,near-surfacesamplestationsin Chileshownby number(Table 2) with referenceto plant formations (Schmithfisen, 1956) and modern pollen spectra (Fig.7). transported from beyond source regions of high andean beech and evergreen rain forest. Low percentages of other woody taxa, for example, Drimys, Myrtaceae, and Aextoxicon, reflect minor entomophilous constituents of the vegetation. Stations in the south or at altitudes higher in the mountains (14 58) are for the most part in deciduous beech forest and evergreen rain forest. They portray large percentages of Nothofagus: N. obliqua type, as much as 60%, and N. dombeyi type, reaching 92%. Woody plants, exemplified by Laurelia, Lomatia, Myrtaceae, and Aextoxicon, are also indicative of these forests and are, at times, well represented. Stations for the gymnosperms Podocarpus saligna (14-17, 51 56) and Prumnopitys andina (14-20, 27-33) reach percentages, respectively, of 43% and 84%. Percentages of another gymnosperm, Araucaria araucana, which grows in communities allied with high andean beech forest (3730'-4023%), are, by comparison, comparatively low (maximum 33%). This is apparently the result of poor pollen dispersal, owing to the large-size pollen grains (77-96 I~m), which sink close to source trees. Irregular percentages shown by Gramineae appear to be associated with varying degrees of openness in the vegetation, often caused by natural disturbances (fire, crustal movement, lahars). At certain high altitude stations (for example, 21 24), Gramineae are indicative of high andean beech forest-tundra transition.

Fossil record of the 10.7-m section from Laguna de Tagua Tagua

Five pollen assemblage zones (TT-1 to TT-5), the upper two divided into subzones, are recog-

116 TABLE II Surface pollen stations in relation to vegetation, temperature, and precipitation" Station Location Lat. 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 La Serena Panquehue Co. El Roble Co. E1 Roble Lag. Lonquen Lag. Aculeo Angos. Paine Rancagua Rancagua Tagua Tagua Teno Lag. Avendafio Lag. Mellizas Termas Chillfin Termas Chillfin Termas Chillfin Termas Chillfin Cen. E1 Toro Cen. E1 Toro Cen. E1 Toro V. Lonquimay V. Lonquimay V. Lonquimay V. Lonquimay V. Lonquimay V. Lonquimay Conguillio Conguillio Conguillio Conguillio Conguillio Conguillio Conguillio V. Llaima V. Llaima V. Llaima V. Llaima V. Llaima V. Llaima E1 Reloj El Reloj Huilipilun Villarrica Villarrica Rucafiancu Termas Palguin Termas Palguin Panguipulli Pto. Nuevo Lag. Ranco San Martin San Martin San Martin San Martin San Martin 2955 3248 3258 3258 3343 3350 3355 3415 3415 3430 3452 3645 3719 3648 3648 3648 3648 3717 3717 3717 3825 3825 3825 3825 3825 3825 3840 3840 3840 3840 3840 3848 3848 3845 3845 3845 3845 3845 3845 3905 3905 3910 3910 3913 3933 3926 3926 3950 4010 4014 3940 3940 3940 3940 3940 Long. 7117 7050 7103 7103 7050 7055 7043 7035 7035 7110 7110 7227 7225 7145 7145 7145 7145 7127 7127 7127 7126 7126 7126 7126 7126 7126 7137 7137 7137 7137 7137 7137 7137 7150 7150 7150 7150 7150 7150 7216 7216 7210 7205 7214 7217 7148 7148 7225 7235 7230 7310 7310 7310 7310 7310 10 540 1890 1800 200 200 300 1060 800 200 300 100 150 1000 960 880 820 850 850 850 1860 1850 1850 1800 1680 1600 1220 1220 1100 1070 1040 820 750 1500 1400 1240 1000 940 850 200 200 240 250 220 260 790 760 180 100 100 50 50 50 50 50
1

C.J. HEUSSER

Alt. (m)

Plant formation

Av. summer temp.

(c)
2 2-3 2-3 2 2 2 2 2 2 2 2 2 3-4 3-4 3-4 3-4 3-4 3-4 3-4 5-6 5-6 5-6 5-6 5 5 4-5 4-5 4 4 4 3-4 3-4 5 5 4-5 4 4 4 3 3 3 4 3 3-4 4 4 4 3 3 3 3 3 3 3 17,1 19,0 16,2 16,5 20,4 20.0 20,0 19.5 19.4 20.0 20.0 19.0 18.0 12.5 12.8 13.2 13.5 13.2 13.2 13.2 9.7 9.7 9.7 9.9 10.3 10.6 11.9 11.9 12.3 12.4 12.5 13.2 13.5 10.9 11.2 11.8 12.6 12.8 13.1 16.3 16.3 16.2 16.2 16.2 16.2 16.0 16.0 16.0 16.0 16.0 16.0 16.0 16.0 16.0 16.0

Av. annual precip. (mm) 60 200 1000 1000 300 400 400 600 600 800 800 1200 1200 2000 2000 2000 2000 1700 1700 1700 2000 2000 2000 1700 1700 1700 3000 3000 2200 2200 2200 2200 2200 4000 4000 3500 3500 3000 3000 2000 2000 2300 2300 2300 2300 4200 4200 2500 1300 1300 2000 2000 2000 2000 2000

ICE AGE VEGETATION AND CLIMATEOF SUBTROPICALCHILE TABLE II Station

117

(continued)
Location Lat. Long. 7310 7308 7307 50 150 100 3 3 3 Alt. (m) Plant formation Av. summer temp. ("C) 16.0 16.0 16.0 Av. annual precip. (ram) 2000 2500 2500

56 57 58

San Martin Paillaco Paillaco

3940 4000 4005

~Station locations by south latitude and west longitude in degrees and minutes, each as four consecutive numbers (see display of stations in Fig. 6); plant formations by number as follows: 1=subtropical thorn shrub succulent, 2=subtropical broad-sclerophyllous woodland (matorral), 3=deciduous beech forest, 4=evergreen rain forest, 5 = high andean beech forest, and 6 = andean tundra. Source of climatic data is Almeyda and S~ez (1958); vegetation formations are according to Schmithiisen (1956, 1960) with modification.

Trees

/ ~ S h r u b s

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Fig.7. Modern pollen spectra for near-surface sample stations in subtropical and adjacent temperate Chile (Table 2, Fig. 6).

nized in the pollen stratigraphy (Fig.8). Leading taxa throughout are Gramineae, chenopodsamaranths, and Tubuliflorae (Compositae); the principal arboreal components, Nothofagus dombeyi type and Prumnopitys andina, gain importance at depth (zones TT-5 to TT-2). Table 3 covers the lithology of the section; Table 4 summarizes the pollen assemblages and radiocarbon time control at and between zonal boundaries.

Zone TT-5, estimated 53,800-50,000 yr B.P.

Over a span of nearly four millennia, chenopods-amaranths increase from 10% to a maximum 66%, following an abundance of N. dombeyi type with Gramineae and Tubuliflorae. This increase parallels an apparent drying trend and lowering of lake level, whereby the peripherally exposed lake bed became invaded by a representative halophytic element (Atriplex, Salicornia, Suaeda). Cheno-

118
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ICE AGE VEGETATION AND CLIMATE OF SUBTROPICALCHILE TABLE 11I

I19

Lithology of sediments in 10.7-m section from Laguna de Tagua Tagua Depth (m) Description
0 1.8 2.3 3.4 6.6 1.7 2.2 3.3 6.5 7.3 Dark ()live gray (5 Y 3/2)" silty clay Olive yellow (5 Y 6/6) silty clay Dark brown (10 YR 3/3) clay, variously indurated Dark olive green (5 Y 3/2) clay gyttja Dark grayish brown (10 YR 4/2) to dark brown (10 YR 3/3) peaty gyttja Gray (10 YR 5/1) to very dark grayish brown ( 10 YR 3/2) silty clay, bottoming in sandy silty clay

for some seven millennia. A drop in lake level is implied, which sustained species of this group on the exposed lake bed. Percentages of arboreal taxa, beech and podocarp in particular, are rather irregular and not so striking as in zone TT-4. Climate, returning to a setting developed over the course of early millennia (zone TT-5), seems to have been relatively dry and less favorable to tree populations.

7.4 10.7

"Munsell Soil Color Charts (1975).

pods-amaranths are identified in the modern pollen rain of the northern part of subtropical Chile (Fig.7), where Atriplex (for example, A. chilense, A. repandum, and A. philippii) are common on saline soils (Navas, 1976). The significant percentages of N. dombeyi type (maximum 29%) imply at least in the beginning the immediate presence of beech, probably as thinly stocked woodland, which later became decimated as drying advanced.

Zone TT-4, estimated 50,000-35,500 yr B.P. During this extended interval, N. dombeyi type is coupled with Prumnopitys andina and the Gramineae. Comparatively mesic conditions and higher lake levels seem apparent, albeit with some fluctuation, as indicated by the variable percentages of chenopods amaranths. Beech woodland enriched by podocarp in the local vegetation is evident from the data, representing displacement of these trees northward and to a lower altitude compared with today. Amounts of Gramineae and Tubuliflorae running frequently >30% and as much as 44%, respectively, also imply much openness. Coincident are Acaena and Rumex (also in zone TT-5). Species of these genera evidently migrated to the lowland from the andean cordillera and higher latitudes, where, for example, A. magellanica and R. crispissimus, exist at present (Haumann, 1918; Moore, 1983). Zone TT-3, estimated 35,500-28,500 yr B.P. Chenopods-amaranths at peak values (maximum 81% at 33,300 yr B.P.) dominate the record

Zone TT-2, 28,500-10,000 yr B.P. For the entire record, quantities of N. dombeyi type and Prumnopitys andina are highest, 32% and 33%, respectively, before 14,500 yr B.P. in subzone TT-2b. These quantities, in concert with increased amounts of Gramineae, Tubuliflorae, and Acaena, contrast minimal amounts of chenopods-amaranths. Conditions supporting optimal beech podocarp woodland expansion were thus in effect for 14 millennia. Climate was evidently cooler with increased precipitation/humidity and reduced evaporation to an extent not shown previously. Modern pollen stations in the Andes (Figs.6 and 7, Table 2) with comparable amounts of beech and podocarp infer much greater precipitalion than at present. The lake, under a regime of high input, presumably greater than the interval of zone TT-4, must have attained its greatest size. After 14,500 yr B.P. in subzone TT-2a, as chenopods amaranths and Gramineae increased under the warming/drying trend of the late-glacial, percentages of N. dombeyi type and Prumnopitys andina overall declined. Zone TT-1, 10,000 0 yr B.P. Holocene sediments contain maximal chenopods-amaranths (>50%) in subzones TT-Ie (before 9000 yr B.P.), TT-Ic (between approximately 6000-2500 yr B.P.), and TT-Ia (after about 200 yr B.P.). Subzones TT-ld and TT-lb (9000 6000 and 2500 200 yr B.P., respectively), identify alternating, corresponding peaks of Gramineae (> 25%), Gunnera chilensis, Umbelliferae, and Pteris chilensis. Over the course of zone TT-1, arboreal taxa become minimal; only subzone TT-Ib exhibits, along with Ephedra andina, an increase of N. dombeyi type. Poor depiction or absence of broad sclerophyllous woodland taxa in the latest record (for example, Cryptocarya, Quillaja, Lithraea) is

120 TABLE IV Pollen assemblage and age stratigraphic data for Laguna de Tagua Tagua a Pollen assemblage zone TT-Ia (0-0.2 m) TT-lb (0.2-0.6 m) TT-lc (0.6-1.5 m) TT-ld (1.5-1.9 m) TT-le (1.9-2.3 m) TT-2a (2.3-2.9 m) Pollen assemblage Age (14C yr B.P.)

C.J. HEUSSER

. Chenopodiaceae-AmaranthaceaeTubuliflorae ............................

ca 200

Gramineae-Nothofagus-EphedraGunnera-Umbelliferae
............................ Cbenopodiaceae-Amaranthaceae ............................ ca 2600 2830 + 120 (1.0 m, RL-1961) ca 6000 6130 + 250 (1.6 m, RL-1962) ca 9000 9100 + 360 (2.0 m, RL-1952) ca 10,000 9860__+320 (2.2 m, RL-1953) 11,170 + 320 (2.3 m, RL-1954) ! 2,970 + 390 (2.4 m, RL-1955) ca 14,500 14,500_+ 350 (2.9 m, QL- 1666) 21,500 ___ 650 (4.0 m, QL-1667) 28,100+ 1400 (5.0 m, QL-1668) ca 28,500 29,800 + 1000 (6.0 m, QL- 1669) 33,300 _ 1400 (6.8 m, QL-1670) ca 35,500 37,000 + 2000 (8.0 m, QL- 1671) > 43,000 (9.0 m, QL-1672) ca 50,000 > 45,000 (10.7 m, QL-1674) ca 53,800

Gramineae-GunneraUmbelliferae ............................ Chenopodiaceae-Amaranthaceae ............................

Gramineae-PrumnopitysNothofagusChenopodiaceae-Amaranthaceae

............................ TT-2b (2.9-5.3 m)

Prumnopitys-NothofagusGramineae-Tubuliflorae

TT-3 (5.3-7.5 m)

............................ Chenopodiaceae-AmaranthaceaeGramineae-Tubuliftorae-

Nothofagus
............................ TT-4 (7.5-10 m)

Nothofagus-PrumnopitysGramineae-TubulifloraeChenopodiaceae-Amaranthaceae ............................ Chenopodiaceae-Amaranthaceae-

TI'-5 (10-10.7 m)

Nothofagus-GramineaeTubuliftorae ............................

aDepths of radiocarbon-dated levels in zones la-e and 2a are from the short section (Fig. 9) and have been adjusted by means of the pollen stratigraphy.

a t t r i b u t a b l e to l i m i t e d p o l l e n p r o d u c t i o n d u e to insect p o l l i n a t e d a d a p t a t i o n s . W h e r e a s t h e a q u a t i c / s e m i a q u a t i c C y p e r a c e a e a n d Myriophyllum a r e b e t t e r r e p r e s e n t e d in z o n e s T T - 5 to T T - 2 , Typha

angustifolia, f o l l o w i n g p e a k Utricularia in s u b z o n e T T - l e , o c c u r s a l m o s t e x c l u s i v e l y in z o n e T T - 1 , c o n t r i b u t e d , as l a k e levels fell, b y p l a n t s c o n s t i t u t ing m a r s h y g r o u n d . D r y , s u b t r o p i c a l c l i m a t e

ICE AGE VEGETATION AND CLIMATE OF SUBTROPICAL CHILE

121

prevailing over the past 10,000 yr of zone TT-I became pronounced during subzones TT-le and TT-lc, when amounts of chenopods-amaranths increased, while shorelines of the lake are presumed to have been lower.

Loss on ignition
Maximum amounts of ignition loss in zone TT-3 at depth (33-38%) result from optimal net organic accumulation from lake biota roughly 35,000 yr ago. Earlier in zone TT-4, lower values imply less than optimal conditions for accumulation. In zone TT-5, quantities were low during the early stage of lacustrine deposition and, as climate became subtropical, were lowest in subzone 2a and zone 1.

Charcoal
Amounts of charcoal (gm 2 cm 2 X 103) increase in zones TT-5 and TT-3. But it is not until later in upper subzone 2a, near the close of the late-glacial, and subsequently in zone 1 that charcoal became abundant. Values appear to increase in phase with Gramineae (out of phase with chenopods-amaranths), suggesting a relationship with rising lake levels, which may favor charcoal preservation. Some oxidative loss of charcoal was possible, however, at times when the lake was seasonally dry. Charcoal in upper zone 2a is presumed to result from burning by paleo-Indians, who are first known to have been active about the lake 11,380 yr ago (Montan6, 1968). Large concentrations of charcoal in the late-glacial and Holocene alternate with maxima of chenopods-amaranths. These amounts suggest that human use of fire as a means to hunt game occurred at corresponding times of high lake level, when climate more favorable to habitation attracted hunters.

records. Trends set by the principals (Prumnopitys, Nothofagus, Gramineae, chenopods-amaranths, and Tubuliflorae) show only slight variation. Radiocarbon dates provide the basis for chronological control used in the late-glacial and Holocene zonation of the 10.7-m section (Table 4). The dates, unfortunately with large errors, are considered reliable, except perhaps the uppermost of 2830_+120 yr B.P. at 1.0m and 6130+_250 yr B.P. at 1.8 m in the section, as these sample levels possibly contain modern roots. For comparison is a date of 6130_+115 yr B.P. obtained from sediments 1.0 m below the top of the lacustrine sequence at the site of the drainage canal (Montan6, 1969). Deeper in the 3.2-m section, the date of 11,170+ 320 yr B.P. at 2.5 m, agrees closely with a date of 11,380+_320 yr B.P. from 2.4 m at the canal (Montan~, 1968).

Estimation of lake level fluctuations from microfossils

Frequencies of aquatic seed plant pollen (consisting principally of Typha angustifolia and Cyperaceae; Fig. 8), algal remains (Pediastrum and Botryococcus), dinoflagellate cysts (cf Peridinium), and massulae of water fern (Azollafiliculoides) are shown in relation to pollen of principal tree, shrub, and herb taxa (Fig.10). For reconstructing past lake levels, algae and dinoflagellates have poor indicator value, owing to the diversified conditions under which they are found (Hutchinson, 1967). Of the remaining microfossils, Typha angustifolia and Azolla filiculoides are characteristic of developing shallow water. In conjunction with the chenopods-amaranths, inhabitants of mud flats during times of low water, these groups serve to estimate lake level. While the chenopods-amaranths range widely in Chile, Typha is temperate-subtropical; Azolla grows throughout below an altitude of 2000 m (Looser, 1961). Diminution in size of Laguna de Tagua Tagua, following origin of the lake, is inferred by increase in chenopods-amaranths. This is corroborated by the shoaling conditions, under which quantities of Azolla were deposited in upper zone TT-5 (Fig. 10). Later, after an interval of relatively high water, lake levels appear to have been lower when

Fossil record of the 3.2-m section

The section, from below 50 cm of cultivated soil and thus truncated at the top, embraces subzones TT-2b TT-lc (Fig.9). Its pollen assemblages and zonation correspond in the main with the upper portion represented by the 10.7~m section (Fig.8); variable kinds and amounts of pollen produced/ preserved locally account for differences in the

122

c.J. HEUSSER

T r e e s ~ /

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/Aquatics&

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Laguna de Tagua Tagua

Fig.9. Diagram of pollen and spore frequency (trees, shrubs and herbs, and aquatics and cryptogams) and radiocarbon chronology, subdivided by pollen assemblage zones, for the 3.2-m section from Laguna de Tagua Tagua. Disturbed upper 0.5 m of the section not shown.

Trees.hru.&her,s

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Fig. 10. Frequency diagram of the pollen of leading trees, shrubs, and herbs, pollen and microfossils of aquatics, and radiocarbon chronology, subdivided by pollen assemblage zones, including estimated changes in lake level, for the 10.7-m section from Laguna de Tagua Tagua.

ICE AGE VEGETATION A N D CLIMATE OF SUBTROPICAL CHILE

123

chenopods-amaranths increased and Azolla reappeared in zone TT-3. For the entire record, as implied by the minimal amounts of indicator taxa, maximum levels apparently were reached in zone TT-2b, which includes the time of the last glacial maximum. Close to 14,500 yr B.P., on evidence provided by greater amounts of Cyperaceae and Azolla, in addition to increasing quantities of chenopods amaranths, lake levels, while fluctuating, began to drop. The sum of data indicates that Laguna de Tagua Tagua for the entire period of record was at minimum size with a considerable part given over to peripheral marsh during subzone TT-lc. Fossil asemblages in the zone consist of peak amounts of chenopods-amaranths and seed plants (mostly Typha angustiJolia, an indicator of brackish/alkaline water), while lacustrine Pediastrum, Botryococcus, dinoflagellates, and Azolla are poorly represented. Ultimately, late in the record, former lake levels were regained to some extent (aquatic microfossils in their variety return in the record), after which, just prior to drainage, the lake again appears to have been decreasing in size. This interpretation of the history of Laguna de Tagua Tagua closely follows the account given by Varela (1976). The microfossil data thus, in the main, complement the geomorphic and macrofossil evidence previously used to reconstruct the sequence of past events. Ages employed by Varela (1976), however, based on two radiocarbon dates covering the top 2 m of sediments (Montan6, 1968, 1969), apply only to the Holocene and late-glacial, whereas at depth, the chronology has been estimated. In addition, the three intervals of erosion noted at the canal exposure were not observed in the core from the central part of the lake, although corresponding times of comparatively low water, when erosion occurred on the exposed margins, are recognizable.

the basin. It is instructive, therefore, to examine extra-local taxa in profiles generated by omitting chenopods-amaranths. Principal taxa are southern beech, podocarp, grass, and composite, which represent > 90% of remaining pollen (Fig. 11). Southern beech is dominant in the Pleistocene through much of the lower part of the section. Beech declines upward after reaching a peak in zone TT-2 and follows trends not unlike those in the diagram of total pollen (Fig.8). Trends of podocarp are likewise comparable in both diagrams, revealing greater emphasis, when chenopods-amaranths are excluded, of an increase of podocarp in zone TT-3. Grass and composite, more or less of equal frequency at depth, are of greatest importance in zone TT-1 during the Holocene. Fluctuations, emphasizing the interplay of these nonarboreal types, show grass the dominant early in upper zone TT-2 and later in upper zone TT-I. On the assumption that the chenopods-amaranths reflect episodes of relatively xeric conditions in the record, it seems likely that more mesic beech podocarp communities at these times were less extensive and, therefore, less pollen productive. However, in the absence of pollen influx, which, given the limited time control, was not possible to calculate reliably, frequency data provide no insight as to numbers of these taxa.
Discussion

Summary pollen diagram of extra-local taxa

Overrepresentation by chenopods amaranths (Fig.8), apparently having invaded the exposed bottom of Laguna de Tagua Tagua when water levels were low, mask performances of plants in communities established beyond the periphery of

Climate and vegetation of subtropical Chile for the last >40,000 yr of the Pleistocene, as interpreted from Laguna de Tagua Tagua stratigraphic data, are clearly contrasted with the setting in effect over the past 10,000 yr of the Holocene. Changes in the ice age landscape were considerable, broadly adhering to the directional pattern recognizable in contiguous parts of the andean cordillera of Chile-Argentina. Few data from elsewhere in southern South America for the time span covered by the lake record establish climatic parameters or explanatory mechanisms for the shifting environmental episodes. Implications regarding temperature and precipitation during Pleistocene episodes of southern beech-podocarp woodland (zones 4 and 2; Fig.8),

124

C. J, HEUSSER

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Fig.ll. Frequency profiles for the 10.7-m section of Laguna de Tagua Tagua of major tree taxa (southern beech and podocarp) and shrub and herb taxa (grass and composite) after omitting chenopods-amaranths from pollen sums.

when precipitation/evaporation ratios were apparently highest, are from estimates derived from the near-surface pollen stations (Table 2, Figs.6 and 7). Average summer temperature and annual precipitation at stations where percentages of Nothofagus and Prumnopitys appear comparable to percentages reached in subzone TT-2b (stations 14-20 and 29-33; Fig.7) are, respectively, close to 13C and 2000 mm. These data, representing the nearest modern analogs and applicable to the last glacial maximum, imply a temperature depression of about 7C and an increase of 1200mm precipitation (present-day estimated values for Laguna de Tagua Tagua are 20C and 800 mm; Table 2). Instances of Nothofagus in peak amounts and Prurnnopitys minimal or unrecorded (compare modern high montane vegetation stations 37 and 38; Fig.7, Table 2) indicate possible temperature depression of > 7C and precipitation greater by a factor of around four. Thus, it seems reasonable to believe that when Prumnopitys became prominent in upper subzone TT-2b, climate was neither as cold nor as wet/snowy, as it was when the proportion of Nothofagus earlier was greater. Explanations to account for fluctuating levels of moisture in subtropical Chile during the ice age are contained in models of atmospheric circulation for the Southern Hemisphere (see references in Heusset, 1989a). The models favor a central mechanism involving oscillations of the polar front, including variable intensity of zonal circulation in the belt of

southern westerly winds. Opinions differ, however, regarding direction of wind movement and sources of moisture. One postulate contends that expansion of the westerlies during the full-glacial was poleward with moisture at Tagua Tagua coming from a subtropical source (Markgraf, 1989). The other maintains that the westerly wind system was strengthened equatorward with precipitation coming from storm systems advancing farther northward than today (Heusser, 1989b). For subtropical Chile, more mesic conditions during the Pleistocene are evident from related studies, albeit with no time control, of the distribution of extinct and relict biota (Kummerow et al., 1961; Paskoff, 1970; Troncoso et al., 1980; Villagrfin and Armesto, 1980; P~rez and Villagrfin, 1985). Westerly circulation in the circumpolar vortex is thought to have extended an estimated 5 of latitude into the subtropics (Caviedes and Paskoff, 1975; Lauer and Frankenberg, 1984), spreading cooler, wetter conditions northward from the south of Chile. Shifting of frontal systems was both poleward and equatorward, in keeping with accompanying changes in the subtropical anticyclone, as explained by Lauer and Frankenberg (1984). The anticyclone lay over interior South America during the last glacial maximum, having shifted its position eastward from over the Pacific Ocean. Thus, more effective continental high pressure, blocking the movement of humid air directed northward along the Chilean coast, is

ICE AGE VEGETATION AND CLIMATE OF SUBTROPICAL CHILE

]25

inferred during drier millennia of the Pleistocene. Glacier advances recorded in the subtropical Argentine Andes (Bengochea et al., 1987; Espiztia, 1989), located at a somewhat lower latitude than Laguna de Tagua Tagua, probably occurred during the corresponding, more humid intervals dated 50,000-35,500 and 28,500-14,500 yr B.P. at the laguna (zones TT-4 and TT-2b: Fig.10). The latest advance (Harcones), with travertine ages on drift of > 10,000 and at least 23,000 yr B.P., falls within the last interval; the advance previous (Penitentes), tentatively bracketed by travertine dates between 38,000-23,000 yr B.P., appears related to the one earlier. Snowlines, rising from west to east during times of glacier advance, indicate glacial nourishment by moisture from the Pacific. This moisture source, as it influenced late Pleistocene high water at Laguna de Tagua Tagua, may also account for high lake level recorded east of the Andes in Argentina. Salina del Bebedero at 3320'S enlarged between 17,500 13,200 yr B.P. (Gonzfilez, 1981; Gonzfilez et al., 1981; M. A. Gonzfilez, pers. comm., 1982), and Laguna Carl Laufquen near 41S was greater in size between 19,500-15,000 yr B.P. (Galloway et al., 1988), Northward extension of frontal systems related to the belt of southern westerly winds, an hypothesis to explain intervals of greater precipitation in otherwise drier latitudes during the Pleistocene, is indicated by fossil evidence from other geographic sectors. Cushion bog species, adapted to wet, southernmost coastal Chile, became established on Isla Grande de Chilo~ (42S) between 27,000-18,000 yr B.P. (Villagrfin, 1988). This migration followed the movement northward of the zone of maximum precipitation, which at present is positioned at approximately 46S. Cushion bogs today are scattered on the tops of the Coastal Mountains as far north as the Cordillera Pelada at 40S. The bogs form disjunctive communities, remaining in unique climatic lacunae since the Pleistocene, when cooler, wetter conditions favoured their spread northward (Heusser, 1982). Late-glacial climate at Laguna de Tagua Tagua after 14,500 yr B.P., as chenopods-amaranths began to multiply (subzone TT-2a; Fig.8), was

characterized by an overall increase of temperature and intervals of desiccation. Communities containing Prumnopitys, not only about the laguna but also at least as far south as 3930'S (Heusser, 1984), remained at low altitudes until approximately the close of the late-glacial. Prumnopitys over its range is subject to winter-wet, summer-dry climate, suggesting that its prominence during the late-glacial, as Nothofagus began to decline, followed in response to developing Mediterraneantype climate. Climate during the Holocene, as at present, was apparently subject to interaction between Pacific cyclonic and anticyclonic atmospheric centers. The premise that the polar front lay considerably south of Eaguna de Tagua Tagua during the early part of the Holocene is supported by data from Rucafiancu at 3930'S which show relatively warm and dry conditions at 10,000-8000 yr B.P. (Heusser, 1984). Prominences of chenopods-amaranths (subzones TT-le, TT-Ic, and TT-la; Fig.8), reflect greater aridity and possibly higher temperatures, resulting from control by the Pacific anticyclone. Conversely, peaks of the Gramineae (subzones TTld and TT-lb; Fig.8), indicate times of somewhat higher humidity and increased incidence of cyclonic storms of the westerlies. The implication from chenopods-amaranths in near-surface pollen station data (station 1, for example; Fig.7, Table 2) is that annual precipitation during the maximum of this assemblage (subzone TT-Ic, approximately 6000 2500 yr B.P.) may have dropped to levels close to 100 mm. This may represent the cumulative effect of drying, beginning in the late-glacial, which emphasizes the growing domination of the Pacific anticyclone regionally. The last approximately 2500 yr of record cover the final episode of peak Gramineae, including small but significant amounts of Nothofagus and Ephedra, and the latest rise of chenopods amaranths (subzones TT-I b and TT-la; Fig.8). Climate is estimated to have been cooler and more humid (summer temperatures possibly 1 2C below the present and precipitation greater by >400 mm). This apparently preceded a reversal of trend, and, over the period, led to increased subtropical conditions. In conclusion, the sequence of late Quaternary

126

C.J. HEUSSER Caviedes, C. and Paskoff, R., 1975. Quaternary glaciations in the Andes of north-central Chile. J. Glaciol., 14: 155-170. Clapperton, C. M., 1983. The glaciation of the Andes. Quat. Sci. Rev., 2: 83-155. Cobos, D. R. and Boninsegna, J. A., 1983. Fluctuations of some glaciers in the upper Atuel River basin, Mendoza, Argentina. In: Quaternary of South America and Antarctic Peninsula. Balkema, Rotterdam, 1 pp, 61-82. Corte, A. E. and Espizfla, L. E., 1981. Inventario de glaciares de la cuenca del Rio Mendoza. Inst. Argent. Nivol. Glaciol., Mendoza, 62 pp. Darwin, C., 1958. Voyage of the Beagle. Bantam, New York, 439 pp. Donoso, Z. C., 1975. Distribuci6n ecol6gica de las especies de Nothofagus en la zona mesom6rfica. Univ, Chile Fac. Cienc. For. Bol. T6c., 33: 1-21. Donoso, Z. C., 1978. Dendrologia. Arboles y arbustos de Chile. Univ. Chile Fac. Cienc. For. Manual, 2, 142 pp. Espizfla, L. E., 1986. Fluctuations of the Rio del Plomo glaciers. Geogr. Ann., 68: 317-327. Espizfla, L. E., 1989. Glaciaciones Pleistoc~nicas en la Quebrada de los Harcones y Rio de las Cuevas, Mendoza, Reptiblica Argentina. Thesis. Univ. San Juan, San Juan, 266 pp. Galloway, R. W., Markgraf, V. and Bradbury, P., 1988. Dating shorelines of lakes in Patagonia, Argentina. J. South Am. Earth Sci., 1: 195-198. Gay, C., 1833. Tagua Tagua. Ann. Sci. Nat., 28: 369. Gonzfilez, M. A., 1981. Evidencias paleoclim/~ticas en la Salina del Bebedero (San Luis). In: Actas 8th Cong. Geol. Argent. San Luis, 3, pp. 411-438. Gonzfilez, M. A., Musacchio, E. A., Garcia, A., Pascual, R. and Corte, A. E., 1981. Las lineas de costa Holoceno de la Salina del Bebedero (San Luis, Argentina). Implicancias paleoambientales se sus microf6siles. In: Actas 8th Congr. Geol. Argent. San Luis, 3, pp. 617-628. Haumann, L., 1918. La v6g&ation des hautes cordill~res de Mendoza. An. Soc. Cient. Argent., 86: 121-188; 225-348. Heusser, C. J., 1971. Pollen and Spores of Chile. Modern Types of the Pteridophyta, Gymnospermae, and Angiospermae, Univ. Arizona Press, Tucson, Ariz., 167 pp. Heusser, C. J., 1982. Palynology of cushion bogs of the Cordillera Pelada, Province of Valdivia, Chile. Quat. Res., 17: 71-92. Heusser, C. J., 1983. Quaternary pollen record from Laguna de Tagua Tagua, Chile. Science, 219: 1429-1432. Heusser, C. J., 1984. Late-glacial-Holocene climate of the lake district of Chile. Quat. Res., 22: 77-90. Heusser, C. J., 1987. Quaternary vegetation of southern South America. In: Quaternary of South America and Antarctic Peninsula. Balkema, Rotterdam, 5, pp. 197-221. Heusser, C. J., 1989a. Polar perspective of late Quaternary climates in the Southern Hemisphere. Quat. Res., 32: 60-71. Heusser, C. J., 1988b. Southern westerlies during the last glacial maximum. Quat. Res., 31: 423-425. Heusser, L. E. and Stock, C. E., 1984. Preparation techniques for concentrating pollen from marine sediments and other sediments with low pollen density. Palynology, 8: 225-227. Hutchinson, G. E., 1967. A Treatise on Limnology. II. Introduction to Lake Biology and the Limnoplankton. Wiley, New York, N.Y., 1115 pp.

vegetation replacements observed at Laguna de Tagua Tagua is ascribed to a predominating, variable storm pattern, regulated by the varying strength of westerly wind circulation. The pattern, in effect, is a modification of existing climate controls, whereby changes in seasonal precipitation, caused by interaction between polar and tropical maritime air masses, involved a fluctuating polar front. Vegetation replacement of broad sclerophyllous woodland occurred as a response to Pleistocene climate, when higher precipitation/evaporation ratios, resulting from greater storm activity under conditions of reduced insolation, characterized intervals of beech-podocarp woodland in subtropical Chile.

Acknowledgments
Thanks are expressed to the late Carlos Mufioz P., who in 1963 first called my attention to Laguna de Tagua Tagua, and to Eugenio Sierra R., who assisted with the first section taken from the lake bed that year. Thanks are also extended to L. E. Heusser, A. Hauser Y., M. Mufioz S., and S. Moreira E. for assistance during subsequent field excursions; M. T. Cafias P., Servicio Nacional de Geologia y Mineria, for field transportation; M. Stuiver and C. Tucek for radiocarbon dating; L. E. Heusser and E. Stock for laboratory processing of samples; and B. van Geel for critical review of the manuscript. Supported by National Science Foundation grants ATM-8115551, ATM-8308021, and ATM-8617154 to New York University.

References
Almeyda, A. E. and Sfiez, S. F., 1958. Rec0Pilaci6n de datos climhticos de Chile y mapas sin6pticos respectivos. Minist. Agric., Santiago, 195 pp. Armesto, J. J. and Martinez, J. A., 1978. Relations between vegetation structure and slope aspect in the Mediterranean region of Chile. J. Ecol., 66: 881-889. Bengochea, L., Porter, S. C. and Schwarcz, H. P., 1987. Pleistocene glaciation across the high Andes of Chile and Argentina. In: Progr. with Abstracts, Int. Union Quat. Res., 7th Int. Congr., Ottawa, 1987, 127. Briiggen, M. J., 1950. Fundamentos de la Geologia de Chile. Inst. Geogr. Mil., Santiago, 374 pp. Casamiquela, G. R., Montan6, M. J., and Santana, A. R., 1967. Convivencia del hombre con el mastodonte en Chile Central. Not. Mens. Mus. Nac. Hist. Nat., 132: 1-6.

ICE AGE VEGETATIONAND CLIMATEOF SUBTROPICALCHILE Kummerow, J., Matte, V. and Schlegel, F., 1961. Zum Problem der Nebelw/ilder an der zentralchilenischen Kfiste. Ber. Dtsch. Bot. Ges., 74: 135-145. Lauer, W. and Frankenberg, P., 1984. Late glacial glaciation and the development of climate in southern South America. In: J. C. Vogel (Editor), Late Cainozoic Palaeoclimates of the Southern Hemisphere. Balkema, Rotterdam, pp. 104-114. Lliboutry, L., I956. Nieves y Glaciares de Chile. Fundamentos de Glaciologia. Univ. Chile, Santiago, 471 pp. Looser, G., 1961. Los pteriddfitos o helechos de Chile. Revi. Univ., 46: 213-262. MacPhail, D. D., 1973. The geomorphology of the Rio Tend lahar, central Chile. Geogr. Rev., 58: 517-532. Markgraf, V., 1989. Reply to C. J. Heusser's "Southern westerlies during the last glacial maximum." Quat. Res., 31: 426 432. Markgraf, V. and D'Antoni, H. L., 1978. Pollen Flora of Argentina. Univ. Arizona Press, Tucson, Ariz., 208 pp. Marticorena, C. and Quezada, M., 1985. Cat~ilogo de la flora vascular de Chile. Gayana, 42:1 157. Mapa Geol6gico de Chile, 1982. Hoja No. 3. Serv. Nac. Geol. Miner., Inst. Geogr. Mil., Santiago. Miller, A., 1976. The climate of Chile. In: W. Schwerdtfeger (Editor), World Survey of Climatology. 12. Climates of Central and South America. Elsevier, Amsterdam, pp. 113 145. Montan& J. C., 1968. Paleo-lndian remains from Laguna de Tagua Tagua, central Chile. Science, 161:1137-1138. Montand, J. C., 1969. Fechado del nivel superior de Tagua Tagua. Not. Mens. Mus. Nac. Hist. Nat., 14:9 10. Mooney, H. A., 1977. Convergent Evolution in Chile and California Mediterranean Climate Ecosystems. Hutchinson and Ross, Stroudsburg, Pa., 240 pp. Munsell Soil Color Charts, 1975. Determination of soil color. Munsell Color, Baltimore, MD., 32 pp. Moore, D. M., 1983. Flora of Tierra del Fuego. Nelson, Oswestry, 396 pp. Navas, L. E., 1973. Flora de la Cuenca de Santiago de Chile. I. Pteridophyta, Gymnospermae, Monocotyledoneae. Univ. Chile, Santiago, 301 pp. Navas, L. E., 1976. Flora de la Cuenca de Santiago de Chile. 2. Dicotyledoneae, Archichlamydeae. Univ. Chile, Santiago, 559 pp. Oberdorfcr, E., 1960. Pflanzensoziologische Studien in Chile. Cramer, Weinheim, 208 pp. Oliver, S. C., 1926. Lista preliminar de los mamiferos fosiles de Chile. Rev. Chil. Hist. Nat., 30: 144-156. Paskoff, R., 1970. Recherches Gfiomorphologique darts le Chili Semi-aride. Biscaye, Bordeaux, 420 pp. P6rez, C. and Villagr~in, C., 1985. Distribuci6n de abundancias de especies en bosques relictos de la zona mediterrfinea de Chile. Rev. Chil. Hist. Nat., 58: 157-170.

127 Pino, Q. M. and Varela, B. J., 1977. Aplicacidn del m6todo de dataci6n por hidrataci6n de obsidiana al sitio arqueol6gico de Laguna de Taguatagua. Actas 7th Congr. Arqueol. Chile, I: 25 52. Reiche, C., 1907. Grundzfige der Pflanzenverbreitung in Chile. Engelmann, Leipzig, 374 pp. Rodriguez, R. R., Matthei, S. O. and Quezada, M. M., 1983. Flora Arbdrea de Chile. Univ. Concepci6n, Concepcidn. 408 pp. Santana-Aguilar, M. R., 1973. La glaciation quaternaire darts les Andes de Rancagua (Chili Central). Bull. Assoc. Geogr. Fr., I973:406 407; 473-483. Schmithfisen, J., 1954. Waldgesellschaften des n6rdlichen Mittelchile. Vegetatio, 5-6:479 486. Schmithfisen, J., 1956. Die r/iumliche Ordnung der chilenischen Vegetation. Bonn. Geogr. Abh., 17: 1-86. Schmithfisen, J., 1960. Die Nadelh61zer in den Waldgesellschaften der sfidlichen Anden. Vegetatio, 9:313 327. Segerstrom, K., Castillo, U. O., and Falcon, M. E., 1964. Quaternary mudflow deposits near Santiago, Chile. U.S. Geol. Surv. Prof. Pap. 475 D: D144 D148. Sufirez, J. A., I983. Rasgos del modelado glaciario en la Quebrada Benjamin Matienzo, Andes Centrales, Cordillera Principal, Mendoza, Argentina. Contribucidn al Projecto de Palinologia IANIGLA. INCA, Mendoza, 40 pp. Thrower, N. J. W. and Bradbury, D. E., 1977. Chile-California Mediterranean Shrub Atlas. A Comparative Analysis. Hutchinson and Ross, Stroudsburg, Pa.. 256 pp. Troncoso, A., Villagran, C. and Mufioz, M.. I980. Una nuewt hip6tesis acerca del origen y edad del bosque de Fray Jorge (Coquimbo, Chile). Bol, Mus. Nac. Hist. Nat., 37: 117-152. Varela, B. J., 1976. Geologia del Cuaternario de Taguatagua (Provincia de O'Higgins). Actas Primer Congr. Geol. Chil.. pp. DSI-D113. Villagrtin, M. C., 1980. Vegetationsgeschichtliche und pflanzensoziologische Untersuchungen im Vicente P6rez Rosales Nationalpark (Chile). Diss. Bot., 54: I- 165. Villagrfin, C., 1988. Expansion of Magellanic moorland during the late Pleisotcene: palynological evidence from northern Isla de ChilD& Chile. Quat. Res., 30:304 314. Villagr~_n, C. and Armesto, J. J., 1980. Relaciones floristicas entre las communidades relictuales del Notre Chico y la zona central con el bosque del sur de Chile. Bol. Mus. Nac. Hist. Nat., 37: 87-101. Wingenroth, M. and Heusser, C. J., 1984. Polen en la Alta Cordillera. IAN1GLA-CONICET, Mendoza, 195 pp. Wyman, J., 1855. Description of a portion of the lower jaw and tooth of the Mastodon Andium: also, of a tooth and fragment of a femur of a Mastodon from Chile. In: U.S. Naval Astronomic Expedition to the South Hemisphere. 2: 275 28 I. Zeil, W., 1964. Geologie yon Chile. Borntraeger, Berlin, 233 pp.