Upper Pleistocene-Holocene climate vegetation of the Sabana Bogota - van der Hammen & González 1960

Upper pleistocene vegetation

holocene climate

and

of

“Sabana

the

South

(Colombia,

and

de Bogota”

America)

by

Th.

van

der

Hammen

& E.

Gonzalez

Contents

Pag. Abstract

262

1.

Introduction

2.

Geographical position

and

3.

Actual

"Sabana

4.

Actual

263

climate

of

the of

vegetation

the

geological history d<>

Bogotá"

"Sabana

de

of

the

"Saban: -

.

and

Bogotá"

and

surroundings

.

.

264

268 271 272

Weinmannietum

tonientosae

272

Iliëto

Eugenietoso

273

Valleetiun

jorullcnsis

Alnetum

274

Quercetum Dwarf-forest

275

of

Vaccinion

the

Andean

floribundi

276

Páramo

and

related

associations

Bscallonietum Meadows

and

276

276

Polylepietum

277

"rosulicaulon"

of

the

Andean

Páramo

277

Espeletietum

278

Acacnetum

278

5.

Tree-line

6.

Recent

7.

The

8.

Description

and

of

9.

Description

and

snowline

279 282

pollen-deposition

boring,

General

the

cores

the

observations

general

The

separate

10.

The

temperature

11.

The

radiocarbon

Conclusions

and

the

sediments

species

and

of

the

diagram

interpretations

diagram

dates

298

300

precipitation and

298

299

curves

and

283 284

interpretation

The

References

Bogota" .

surroundings

Andean-forest

12.

de

their

curves

interpretation

301 309 313

315

Abstract

The and

intervals

of

of

for

them

in

the

later.

above

level,

sea

From

at

the

phases was

of

for

temperature the

Würm

1300

some

Radiocarbon Holocene curve

and

for

the

the

North

and

the

have

been

at

that

than

for

and

It

families

content,

pollen

be

and

recognized,

the

results

West

of

that

the

at

many

will

be

metres

2500

Greenwich.

interglacial periods

shows

also

Pleistocene

its

approximately

of

74°—74½° and

sediments,

could

being analysed altitude

an

glacial

America.

affected

glacial periods

the

were

interglacials interpluvials. of

the

8° C

±

(fig.

5).

lower

and

now

of

tree-line, changes

calculated

were

less

is

core

lake

of

genera

equator,

deduced

this

glacials

Drenthe

stadial).

The

lowest

stadial),

Then

more

for it

the

the

together

important

of

part

followed

follows

of

the

surface

seemed

interglacials

long interstadials, The

Pleistocene

Emiliani

knowledge

and

that

prove

today,

than

the

showed

altitude an

and

precipitation

annual

Temperatures during

snow-line

the

Sabana

the

de

and

North

diagram

of

the of

even

considered

by

Gross for

correlate

America, to

present

the and

study

also

using

correspond

high-glacial the

tree-line differ-

greater

and

are

be

the

respectively temperature

surprisingly

well

with

Europe (fig. 6). the

older

phases

principally to

the

Würm-glacial the

to

Moreover

ages.

interstadial,

interglacial,

the

to

seems

I—II

Interpleniglacial),

conclusions

section those

Bogotá corresponds

justified

and

Riss

Riss-Würm called

to

water

ocean

fully

the

the

correspond

Europe

of

by

of

parts

Würm-glacial really

upper

nomenclature.

(=

dates

later

published by With

the

of

fluctuations

glacial

metres

lies

were

(fig. 5).

ence

that

real

the

be

and

pluvials,

the

of

Bogotá

of

analysed

principally

core

Bogotá

species,

rest

de

may

Europe

metre

de

seventy

North

it

2436

a

Sabana

The

time. Sabana

4½° —5°

as

the

About

diagram

time

same

Curves of

first

of

metres

from

cm.

The

the

equally

changes

age

10—15

published

tropics

32

uppermost

Holocene

end

the

Riss

II

the of

(=

(subdivided

with

alpine Hiss

I

Warthe

by

two

Holocene. summarized

in

paragraph

12.

1.

The

Sabana

Eastern

considered lived

de

the

on

cording

bottom before

the

(South

of

the

Sabana

Bochica, the

through

away

of

became

once

Chibcha-heroe

ran

lake.

arrival

which

America),

old

an

mountains in the

plain surrounded by

upland

an

Colombia

flat

skinned

which

Sabana,

of

Sabana

which

to

the white

the

as

is

Bogotá

Cordillera

INTRODUCTION

Even

lake. a

day

had

legend

This

opened

present,

always

been who

Chibeha-indians,

Spaniards,

the a

the

has

also

the

to

way

of

outlet

legend,

a

tells of

waters

the

ac-

that the

Tequendama

waterfall. When

analysing

Sabana-formation,

consisted almost

thick,

So

pollen. and

the

Pliocene

material.

the

ous

At

section,

At

de

Bogotá,

this

point

and Dr.

Dr.

of

Department the

this

technical

We

also

several

providing

Schlumberger any

to

the

Geological

of

of

a

from which

case

the

of the

of

of

pollen-analysis and

the at

equator,

the

the

Lopez

upper 33

to

Dr.

to

Casas,

(¡eológico

metres

of

of

the

metres.

2560

Hubach

Enrique

Geológico Nacional have

facilitated the

former

Nacional,

chief of for

the

help

his

boring.

our

and

Colombia

lake-sediments,

Pleistocene

altitude

an

director,

present Jaime

Servicio

the

the

continu-

taking

the

Upper

thanks

our

express

the

in

drilling equipment

Institute,

base

from

cores

out,

reached.

was

the

Dr.

to

core-barrels of

to

Silva,

express

present,

was

carried

former directors of the Servicio

and

study,

and

algae

pollen-diagram

was

the aid

surface

the

of

would like

execution

wish

of

boring

with

Holocene

North

hydrogeology

of

of

clays,

the

Fernando Paba

realization

in

we

special

a

the

the results

Benjamin Alvarado,

facies

metres

Pleistocene

collected continuous

get carefully

below

red-mottled

±y>>°

lake

continuous

in of

series

young

of

world. to

department

comprising

rich

very

in

developed

complete

University-City,

metres

present

now

Sabana

to

tertiary

on

the

203

thick,

the

hundreds

whole Pleistocene until recent,

purpose

the

Hydrogeological

We

and

this

of 1957 in

cores.

resting

of

For

all

a

the

how

was

lake-sediments, complete,

very

from

samples

several

formation,

sediments,

in the

unique

a

making

throughout

the problem

beginning

of

this

of

here

bore-hole

some

that

entirely that

Pliocene

even

almost

But this

seemed

possibility

Upper

would be

of

it

probably

giving the

provisionally realized

we

thanks

to

other

were

so

Shell-Cóndor

the

in

for

Bogotá

accessories.

kind

to

make

the

electric

log

without

cost.

Financial support

national The

Geophysical C14

Copenhagen Mr.

all

B. the

given

by

glaciology

the

commission

of

the

Inter-

year.

analyses

laboratory,

Juan

prepared

was

were

carried

through

Perico, samples,

the

laboratory and

did

out

gratis,

assistant much

Dr.

by

intermediary of

of

Dr.

the

valuable

H.

Johs

Tauber

palcobotanieal work

of

the

Iversen.

on

all

section,

our

field

excursions. The

collaborators

Mr. Roberto

of

the

Jaramillo and Mr.

Instituto

de

Ciencias,

Jorge Hernández,

especially

contributed

my

essentially

friends to

the

264

study,

present

Mr.

Erwin

altitude and

studies

the

in

specimens

of

National

Kraus

the

have

all

these

supplied

friends finish

to

The

"Sabana de

Colombia. 5°

Its

north

flat

several

THE

surrounded

Biirgl,

towns

all

at

the

by

be

to

Bogotá black

was

a

lies

principally and

red

in

the

them

of

1400

not

drained

the

by

,

0

4y 2

which

on

of

capital closed

the

basin,

river

Bogotá via

an

Greenwich. 2

km

completely

south-west,

at

Cordillera

between of

the

Bogotá,

almost

plains

Eastern

West

74%°

area

result

there

of

well

still

the

Lower

the

and

Alicachin-El

geological

known

remain

history.

(Hubach,

several

deposited,

with

sedimentation

is

the

of

1957;

structural

already

is

to

or

(Guadalupe

of

suffered

the

retreated.

sea

and

finally

a

On

age.

of

top

this

the

except

formation presents

shales and

slight

upheaval

clays,

sedimentation

sandstones

the beginning

at

movements took

the

of

frequently

formation).

Guaduas

Fresh

in

resulting

in-

sedimentation

(grey

a

de

(Villeta

sandstone

Maestrichtian age,

The

continental facies

Paleocene,

the

Maestrichtian

Paleocene.

geosyncline

with

Sabana

layers

of

amount

("plaencrs")

also

actual

the Lower Cretaceous

sandstone

the

closes

Lower

formation,

During

some

Cretaceous

Upper

the

Tertiary

Andean Geosycline.

Guaduas

the

sandstone forms

Cacho

mottled

the Lower

with

a

Lower

clays

a

the

This

formation

in

place

of

the

sandstones

foraminifera of

beginning

rests

of

layers, the

may

on

be

top

probably

Upper

the

Bogotá

formation,

In

the

principally

middle

part

of

the base present, representing In

of

the

Tunjuelo valley, Bogotá

the

and Lower

beginning

Oligocene,

and

Oligocene

of the Middle

that

part

south

formation

Middle Eocene age,

Eocene

marks the

Upper

of

formation consists

sandstones.

sandstone).

sandstone of

several coal

sandstone

Bogotá

containing

(Lenguazaque

Usme

locally

clay

the basal

age.

grained sandstone

thick basal

sandstone with

Eocene

and

coarse

Eocene

the

Bogotá,

consists of

At

Marta.

would

HISTORY

approximately

an

rather

1957),

Tierna-sandstone

transitional

Thus

formation

series

the

which

Paleocene and

of

of

the

of

guide

sandstone).

The

the

is

region

and

were

marine

the Maestrichtian,

(Cacho

of

a

Santa we

several upland

is

the

the

logically

Hammen.,

sandstone

part,

coals).

beginning

of

the

Tierna

uppermost

of

shales

sandstone called

sandstone

of

and

an

in

sandstone and siliceous rocks

respectively

The

is the

Cretaceous

During

and

creases,

of

part of the Bast

marine

formation).

has

and

outlet

de

help

the

on

our

resolved.

the

During'

Nevada

was

thanks.

one

(fig. 1)

mountains,

and

waterfall.

der

Van

of

BOGOTT

between 74°

Bogotá

observations,

whose

above sea-level

represents

single

situation

geology

1957;

de

Bogotá

a

the

pollen

recent

GEOLOGICAL

situated, amongst

are

sides

with

present

problems

soft

Sierra

DE

largest

metres

and

Sabana

Charquito-Tequendama

Although

2560

Sabana de

tributaries,

The

is the

Bogotá"

the

important The

“SABANA

his

or

without

sincere

our

geographical position

of

of

mountains,

the

institutions

study,

of the equator,

part

country.

its

in

companion

and

approximately

in

collaborating

collection

based

snow

GEOGRAPHICAL POSITION AND

altitude of

The

collections,

the

data,

many

different

in

this

OF

and

plant

facilitating

Herbarium.

snow-limit

been able

2.

of

and

inestimable mountain To

the recent

determinating

vegetation

recent

of

a

and

shale

age.

A

Oligocene.

the

Eastern

OF

DEPARTMEN THE

THE AND

BOGOTA,

OF DE

THE

LOCATION SABANA THE

INDCATING CUNDIAMRC, MAPS

PARK

UNIVERSITY THE IN X

BORING

265

an ex s see 1b

fig.

266

which

Geosyneline movements:

At that

began

time the

This

the final upheaval Sabana

the

the

may

do

we

A in

know

not

what

and clay,

loam

of

base

the

found

were

lower

did

suffer

not

Probably

of

mainly

in

lake

formation

do

inundations

by

with

below,

of

part

It

the

the

and

but

the

lake wet

or

in

the

the

the

Selling, much

still

was

suffered

The

its

formation

often

are

the

the

present.

Sabana formation It

Sabana.

that the of

metres

but

sediments,

of

inundations,

lower

with the inundation of

of

Near

(Berry)

faults

and

gravel,

consists

thick.

These

seasons.

today

occur

small

Holocene,

uppermost

marsh

kaolin.

Tilatá formation.

Tilatá

sand,

Cordillera

depths, showing

the

Only

of

Cordillera

given

the

Sabana,

proper

and

is

Bogotá

consists

Eastern

metres

different

de

became

the

Sabana

deposited

clays

notwithstanding

of

parts

lake

the as

of

during

improvement

the

climate

The

Sabana.

will be shown

clays coincides,

the

final

glaciation.

the

its

that

formation,

Pleistocene,

ash

truly flat part 400

to

at

of

deposits

be clear

of

200

beginning

last

will

last part river

still

of the lake

contact

the

now

the

during

drainage,

artificial

is

consist

not

Several

indicate

to

cipaconensis

but

imporrtance,

intervals.

at

the

of the

deposition

any

present

are

that

indicating

the whole Pleistocene and

what

marsh

a

of

Humiria

that

It

it.

volcanic

diatomite,

seems

the

surround

of

Sabana

into the

debouching

fruits

it

sediments,

Peat-layers partly

valleys

1957),

folding

during

deposited

was

the older

fossil

after

principal upheaval

the

possibly

in

So

day.

to

and Then

altitude in

same

seems

through

sometimes

(Hubach,

than

Sabana

valley.

the

for their

and

mountains which

and

fact

responsible

the

Miocene.,

Pliocene.

the

syncline

East-West

Pliocene

formation

the

closed

This

the

is.

deposited

was

the base of

near

Cordillera. feature

of

approximately

at

orogenetic

Hammen, 1959).

features of

during

beginning

and

deep occur

tectonic

it

the

During

formation

Eastern

section

geological 2.

fig.

the

major

some

wide, plains

folding

the

der

(Van

completed

in

suffered

Cordillera,

morphological

was

probably

place

upland

of

part be

probably

structural and

composed,

a

Eastern

the

and

development

took

(but smaller)

central

there

was

is

upheaval

principal

develop.

to

similar

actually

slight

a

that

in

Pliocene

the

of

course

and

tributaries,

the of

great quantities

in

deposited

and

Quarternary

rock

the

eroded

were

form

of

and

clay

the

probably the

by

Bogotá

sand

in

the

Sabana-lake. Thus the

end

subject

to

the

out

thickness

of

the

The

the

of

we

Sabana

of

amount to

equally

may

of

the

Thus

are

Sabana

As

now.

mountains

of

is

over

draw

the

somewhat

Sabana

Sabana flat

it

is

higher

were

for

important

in the past,

were

that

area

is

(the

to

say

area,

it

conclusion

less

than

200

may

the

flat part 2500

would

that

at

our

carried

we

The

2

but

constant, contains not

are

estimated

500

km

.

not

Sabana

be

1400

km

has 3

420 km

thick,

very 80

at

km

3

.

3

.

catchment-area of

of

the

Bogotá

the Sabana and the somewhat

km

form

the

metres

is

valleys

approximately

approximately this

the

higher

there

is

Bogotá

formation

somewhat

is

de

part of

deposited

sediments

erosion,

the

the

the

material

sedimentation

were

the

surrounding

they

these

of

part

sediments in

valley-plains).,

material

Thus

flat

metres.

300

exposed its

that

higher

sediments

The

total

area

higher of

of

the

quantity

river less

mountains than

calculation.

of

area

material.

the

the

Pliocene

know how much

average

and So

the

following

The

an

naturally

of

2

.

a

Extending layer

mountains

higher

at

of

200

which

the end

of

the

500

metres

surround the

km

3

thick.

the

Pliocene

2

FIG.-

268

than

If

nowadays.

average

We will

thousand years.

3.

the

sedimentation

sedimentation would

of

rate

ACTUAL

CLIMATE

2

1

was

in

em

OF

THE

million

century,

a

that these figures

below,

see

time

be

are

SABANA

years, 20

or

near

very

the

in

one

to the truth.

AND

BOGOTร

DE

then cm

SURROUNDINGS

The of

Sabana

2560

m.

order

In

it

is

department

of

changes,

de

These

Bogotรก

be

to

lies

factors

two

able

necessary

414

o

lat.

to

appreciate

to

have

a

the

look

The

valley, of

the In

to

the

order

relatively

the

Bogotรก";

Cundinamarca,

tation,

of

centre

tropical

East

to

at

know

we

but

department

in

the

first

Eastern

spread

wind),

of

at

the

and

equator

an

altitude

Holocene

climatic

at

climate.

Pleistocene the

and

actual

climate

of

the

entire

Cundinamarca.

Fig.

"Sabana de

N.

determine its

West

high

Plains the

plotted

because

it

mountains

relation

of

see

by

the

part of

up

to

high

the

4000

plain

metres

of

the

Magdalena

tropical

and then part

Orientales).

of

than

variable we

occupied

comprises

(Llanos

more

nevertheless

is

3

average 100

local

temperature

points

in

conditions

that the average

fig. of

3. each

clearly

and

altitude

The

points

place forms

in are

(precipia

straight

269

line.

This

last

Eastern

gives

The

metres

this

from

rises

interested We

Bogotá

of

the

in

two

fig.

seasons

The

wet

temperatures

months.

This

is

us

the

following

wet

4a).

take

consult

may

quote

average

is

night the

the Cordillera

of

this

much

Eidt

of both

the

gives

from

a

two

way

4540

±

The

m.

snow-line

in

the

that

the

2

lowers

temperature

general

change

in

local

/

°

C

3

conditions

due

to

the

the

following

June-September

In

meet.

is

by

the

zone

zone

the

sun

are

the

sun

of

westward

dry period

moves

four or

may

the

and

the

of

anyone

Sabana

with two

dry

higher

The

during

radiation

de

seasons

October-November.

the

during the

sometimes

(night-frost

the

of in

the

winds

caused

occur:

two

zone

occurs

are

one

one

seasons.

"The

the trade

winds

wet

where

frequently

principally

from

passes

maximum rainfall

this;

dry seasons).

situated

this

this

(1952),

system

42

explanation is

On

Eidt.

that ground

overcast

no

during

the

discuss

to

months and

dry

fact

Eidt

by to

Eidt.

report

April-May

during

there

if

picture

going

of

here

fall

lower

mainly

when

detail

study the

extensely

according

seasons

fall in

of

treated

differentiated, alternating

that while

periods

the other

The may

at

the

may be

(of Cundinamarea)

hemispheres

such

ward,

the

(provided

was

into

mentioned

facts,

are

greater

strong rains which

year,

0° C

with

department

far

too

"Sabana de Bogotá"

department

in

that

curve,

FIG.

on

well

quite

Cundinamarca

climate

would

It

Koppen.

wet

one

climatology

classified

(see

corresponds

change).

not

who

conclude

may

100

every

temperature of

theoretical

a

altitude

Cordillera.

We

do

line

mentioned

tropic

when

to

the

variable.

the

by

the

sun

The

convergence,

other

moves

every north-

southward".

equivalent eastward

be much

alternating

from

more

the

dry

Sabana.

pronounced

and

wet

stations

Westwards

than the other

the one,

270

and

eastwards

the

December-February

dry

period

be

may

much

more

pro-

nounced. While the it

low,

total

increases

The

annual

rainfall

yearly

both

towards

precipitation

the on

the

on

NW

with

local

the

region The

to

the

maxima

towards of

and

of

These

difference Bast

of

the

great

of

the

(relatively and

slopes

with

the

therefore

steeply

a

on

Sabana:

dry

a

forests

Quercetum

is In

of

almost

slopes

of

600—900

down

Going

increases

to

mm,

from

2850

(relatively

the of

west

is

that

temperature

humid

of

the

mm

in

this

forests,

the on

(xerophitie

range,

covered

main

the

between maxima or

hill range

a

region

where

the

much greater rain fall.

a

continuously

fact

with

rainshadow of

there is

difference exists

higher

temperature-differences).

the

to

This

the

part

regions,

places

these

in

driest

contrast

effect".

clear with

the

warmer

many

"greenhouse

vegetation

Bogotá).

precipitation

Sabana lies

it

toward

are

is

Bogotá

relatively

SE.

4b.

(near

mm

annual

de

is

Bogotá"

the

4b).

Mosquera, etc.).

country slopes

clouds,

(fig.

border of

western

vegetation

1390 the

NW,

Sasaima

west

till

up

the

de

towards

Sabana

the

FIG.

Sabana

"Sabana

and

slopes

the and

with

of

fog-

for

reason

the

the hills

Weinmannietum lower

cloud-forests,

minima)

with

not

so

271

4.

ACTUAL

VEGETATION AND

the

Although well

quite

specific real

known,

Cuatrecasas

(1934).

observations

are

which

he

work,

which

study

Cuatrecasas

studied the

of

reality

of

fact,

and

history,

of

in

if

the

the

Eastern

the

but

the

We

associations of

the

and

in

this

present

were

importance

Roberto

Ciencias)

Cordillera

the

which

great

his

himself,

in

use

Mr.

by de

Instituto

Eastern

will

is

by

short time

admire profoundly

data.

surveyed

published

states

Those

are

Cordillera

only

were

considers

one

nevertheless

were

from

(both

BOGOTT

DE

Cuatrecasas

as

definitions. which

of

studies

cannot

one

wealth

a

and

distinguish

flora

Nevertheless,

gives

names

vegetation

may

the

"observations",

Hernández

Jorge One

in

“SABANA

vegetation

matter

a

Cuatrecasas,

study

Mr.

his

realized

by

As

THE

of

content

sociological

complete.

not

OF

SURROUNDINGS

not

for

Jaramillo, the author.

following

major

vegetation-belts.

Snow 4500 Meadows and

"resulicaulon" of

m

the

andean "páramo" 3200—3300 Dwarf-forest

of

the

m

andean

"páramo" Andean

3000—3200

m

2400 —2500

m

forest ±

Subandean forest

forest

Tropical

Of

these

will

and

be

the

mannion and forest

of

more

flat is

Weinmannietum,

of

part

the

special

a

of

are

detail.

in

Sabana

for

importance andean

The

which include the

the Quercion,

the

three

uppermost

discussed

de

association,

of

majority Bogotá, is

as

the

the

Wein-

the associations. related

although

the

study,

present

includes

forest

The

to

the

determined

edaphically

Alnetum. The

and and

(meadows It

locally the

has

belt

certain

kept

in

It

is

valleys of

associations, more

for

areas

near

the

that

are

covered tree-line.

the

the

the andean that

according

Vaeeinion

to

in

the

may

of

the

considerably

the belt

highest

Espeletion.

these

belts

miero-climatical conditions. up

go

includes

The

includes

paramo)

altitudinal limits the

to

andean paramo),

Weinmannion.

higher

in

may

Thus wind-

note,

the

that

there

following

are

short

many

local

descriptions

variations

only

some

in of

the the

mentioned.

"páramo" cold

of

slopes.

importance

term

the

or

and

important

The used

mind

considerably

Weinmannion and

of

(dwarf-forest

"derivations"

"rosulicaulon" of be

to

differ

protected

are

higher

next

Vaccinion,

higher with

used

parts

here

cannot

of the

meadows

and

be

strictly

mountains.

resulicaulon,

In

defined; the

or

it

present

with

the

is

popularly

study

these

dwarf-forest

272

ANDEAN-FOREST

Weinmannietum tomentosae

On

the

"Sabana

of

slopes

de

the

Bogotá"

mountains

and

further

(Cuatrecasas,

which

East

in

form the

the

1934) eastern

border

the

Cordillera,

most

of

the

common

is with climax forest Weinmannia original vegetation relatively high a tomentosa L.f.

parts

of

destroyed

by

2800—3100 The a

as

the

a

it

still may

of

be

precipitation

and

climax-vegetation

man,

intact

in

but over

June-September)

have,

14° C.

and

two

There wet

as

far

an

lower

partly

altitude

could be

as

The

mm.

dry

two

are

stations

at

the or

of

areas.

650—900

approximately

entirely

places

large

On

species.

was

various

rather

Weinmannietum of

temperature lies between 9° and March

dominant

frequently

original

covered with

slopes

total annual

the

influence

the

m

and

constant

slopes

stations

(April-May

checked,

average year

(December-

and

October-

November). Cuatrecasas Weinmannietum

mentions near

amongst

others

the

following

species

Guasca :

Weinmannia tomentosa

Hypericum

goyanesii

Hesperomeles

obtusifolia

Hypericum

mexicanum

Hes peromeles

goudotiana

Drimys Clethra

Hypericum

laricifolium

granatensis

Hypericum

brathys

chrysoleuca

Berberis

discolor

Oreopanax Clusia

sp.div.

Tibouchina Bucketia Vallea

My

rica

goudotii

parviflora

Pernettya pentlandii

grossa

Ribes

columbianum

Rubus

floribundus

stipularis

Rubus

loxensis

trianae

Rubus

nubigenus

Gynoxis

glutinosa

Baccharis

floribundum

Diplostephium

rosmarinifolium

aff.

Salpichroa

diffusa

Muehlembeckia tamnifolia

Miconia eleaoides

Passiflora

Miconia

Gaultheria coccinea

Miconia

summa

ligustrina

Hypericum Rapanea Senecio

hartwegii

ferruginea lanatus

lanata

Gaultheria

lanigera

Diplostephium Valeriana Bomarea

tomentosa

Viburnum

suratence

Blechnum

Vaccinium

floribundum

Alsophila

moritzianum sp.

Gaultheria anastomosans

Lycopodium

Cavendishia

Peperomia

cordifolia

phyllicoides

oblongifolia

complanatum blanda

Macleania nitida

Pilea

Ceratostema

Cortaderia colombiana

parvifolium

aff.

jamesoniana

tessalata

Cavendishia guascensis

Chusquea

Miconia

Tillandsia recurvata

salicifolia

Tillandsia turneri

Bejaria

aestuans

Bejaria

resinosa

Guzmania sp.

Aragoa

abietina

Pleurothallis sp.

Arcytophyllum

nitidum

Perilochroa lindenianum

Gaylussacia

buxi folia

Hymenophyllum

sp.

Aetanthus mutisii Gaiadendron tagua

from

the

273

At

or

that altitude different and

several

or

the

in

general

the

Cuatrecasas the

with

plant,

the

Stevia lucida, Solanum

Rapanea

to

It

nevertheless

seems

vegetation

of

Sabana.

the

lower

Probably

of

frequency

have been

may

less

Iliëto

The

was

very

the

as

Miconia

climax

the

Bogotá,

near

most

Vallea

constant

stipularis

floribundum,

Durantha

mutisii,

Eupatorium.

Other

Miconia,

Lantana,

considers

present,

Weinmannia tomentosa, Bomarea

div.,

sp.

this

vegetation

of

part

near

as

frondea, near

very

dominant species

Ilex

kunthiana

Alnus

and

Piper

Eugenia

following

Verbesina

Miconia Senecio

Sabana at

Together list

of

(J.

sp.

crassiramia

pyramidalis

reclinata americanus

is a

occupied few

with

Mr.

were

Physalis

etc.

nepalense

Muehlembeckia

Plantago

Peperomia Baccharis Rubus

sp.

sp.

Eupatorium

fastigiatum

sp.

floribunda

bogotensis

Cynanchum Borreria

sp.

anthospermoides

Castilleja

palaefolia

Solanum

sp.

Nertera

sp.

caripense

(Gomozia)

completely

of

Hernández

peruviana

buxifolium

Oxalis

Jorge

found:

melanochoranthum sp.

been

Aragoa

almost

remnants

plants.

Cestrum

Salvia

in

inedited)

today

places

Centrum

Alchemilla

the

Cordia

have

may

Valeriana,

Hernández,

Polygonum

bogotense

Polymnia

bogotanum

Ribes,

with

changes

stipularis,

foliosa

plants

surrounding

small

Vallea stipularis

sp.

Hesperomeles

the

and

Croton

Drimys,

the

certainty

identical,

being

Vallea

jorullensis,

and

like

almost

not

are:

jorullensis

Symplocos

Alnus

with

mountains

differences

only

Nevertheless

following observations

The

if

the

absent.

or

Suba.

reconstruct of

slopes

Valleetum Eugenietoso

especially

Moreover the

the

species

to

similar,

Bocconia

other

grassland.

Subdominant is:

the

with

plants;

frequent

flat

truly

cultivated

the

in

Baccharis

ferruginea,

v.

difficult

parts

it

Piper,

while

frequent,

more

be of

several

Palicaurea,

lanata,

by

both

majority:

Clusia,

incisum,

vegetations,

frutescens.

lanata

the

cordifolia,

bogotanum,

to

Weinmannietum tomentosae, the

Oreopanax

Cuatrecasas

etc.

Cordia

of

Often

important species,

of 2650—2750 meters,

with

jorullensis

the

species,

Bocconia

forming

Cavendishia

aquilinum

also

angustifolia,

P.

Croton

ferruginea,

destroyed

vegetations

climax.

a

the

species

Alnus are:.

forest

secondary

of

one

interesting

is

altitude

an

Piper croccatum,

species

Pteridium

from

and

cornifolium,

important

An

ones,

become

may

Cordia lanata”,

following

low

in these

frequent

very

Ericaceae.

lineariflora

and

partly

is

vegetation

shrub

represent the original climax vegetation.

are

secondary

of

The

jorullensis

describes

"Consoeietas

Palicaurea

Miconia

the

metres

man.

do not

Alnus of

species

by

past)

of

places

and

vegetation

2600—2800

in

species

other

on

of

altitude

an

(recently

sp.

forest we

are

made

274

sp.

Polypodium

angustifolium

tristis

Polypodium

murorum

Hydrocotyle Salprichoa Bomarea

lanceolatum

Polypodium

sp.

Senecio

formosus

Dryopteris

Prunus

serotina

paleacea

Rubus

floribundus

Solanum

nigrum

Rubus

guianensis

Rhamnus

goudotianus

Rubus

glaucus

The

differences of

principally

much

Bogotá" or

of

annual

13°—14°

the

On

Alnetum of In

the

dominant The

"Sabana

are

de

found Alnus at The

a

the

flat

the

flat

and

are

of

part

zone

the

flat

the

which

Northern

Alnus

the

an

the

been

very

wet

common

But

well

list

of

in

even

plants

preserved

few

the

Weinmannietum,

higher.

the

In

Sierra

racemosa

Prunus

serotina sp.

peruviana vulgaris

Rubus

floribundus

Rubus

nubigenus

Rubus

glaucus anthospermoides maritimum sp. revoluta

sp.

about

stream to

the

is

the

parts

of

remnants of

up

altitudes

to

Nevada

the this

del

Cocuy

of we

m.

was

made

Alnetum North

Mr.

by of

Jorge

Hernandez and

Funza.

granatensis

Nertera

cf.

fastigiatum

Cuphea

Stachys

(old

similar

lower

the

on

to-day

aphanoides

Baccharis

is

ferruginea

var.

Lachemilla

Geranium

de

scarcer,

Hemisphere.

jorullensis

seasons.

altitude of 3500

buxifolium

Lythrum

Sabana

very

Lachemilla

Borreria

much

Sabana

is

Eupatorium

Prunella

"Sabana

the

indedited)

part of

develops

the

of

of

part

Solanum

Jussiaea

may

rather

association.

occurs

jorullensis

Epilobium

general

general

Miconia reclinata

Cestrum

Eugenia

in

mm.

Hernández,

vegetation

possibly

following

in

but

left.

and

metres,

in

places

have

must

(J.

Bogotá"

this

jorullensis

Alnus

Alnus

of

Alnetum

association

author

jorullensis

temperate

species

In

Vallea

limit,

Weinmannia is in

rainfall about 700—900

inundated during

Sabana,

3100

the

due

absent.

peat-forming

a

and

temperature of

flooded

easily

wet,

etc.),

Weinmannietum may be

Ilex,

forest

Weinmannietum.

Alnetum

beds

the

species

the

to

up

dominants,

average

and

C,

with

conditions.

sub-dominant

be completely

even

The

the

become

they

may

association

higher, practically

species

scarce

this

and edaphic

dominant and

The occur

soil

to

Oxalis

nigrum

Verbena

sp.

hydropiperoides

Polygonum Calceolaria Galium

Stellaria Carex

mutisii

sp.

palustris

trianae

cuspidata

spp.

Cortaderia sp.

Scirpus

californicus

Cynanchum

tenellum

Pol podium y

angusti folium

Blechnum

sp.

Dryopteris

sp.

Equisetum

bogotense

the

275

Quercetum A

Quercetum

between

Sabana

the

flanks

of

de

lower

annual

higher

Also

of to

in

in

occurs

3600

many

favorable

(see

Thus

it

of

forced

are

East the

on

appears the

Here

almost

heavy

altitudes

at

immediately

limit

regions.

vapour

a

to

more

with

Sabana,

air

from

streams

rise

of

west

to

the

continuous cloudiness

and

to

up

of

the

be of

of

(North

found up

to

at

Quercetum

such

frequent

clouds

such

and

altitudes and

altitude

an

be

may

higher

attendant

the

Quercetum

above

possibly

rainfall

higher

as

Cundinamarca)

or

due

average

"greenhouse

3).

that

see

instead

Boyacá

may

occurrence

paragraph

both

as

and

conditions,

continuous

more

case,

The

due

we

Quercetum

in

but

western

warmer

water

of

department places,

local

temperatures

with

resulting

Cordillera

Cordillera

absent,

the

forms

Eastern

the

precipitation.

the

metres.

effect"

a

Sabana,

completely

towards

slopes

saturated

the

is

the

In

metres.

which

range

steep

regions,

altitudes of

it

in

present

3600

±

Bogotá

hill

the

rather

their

be

may

2500 and

±

of

one

a

the

clouding.

The and

Quercetum

factors

principal

Weinmannietum is soil

the

higher

a

to

seems

be

for

the

annual of

Weinmannietum

no

a

and

influence

this

found

are

of

occurrence

precipitation in

especially

on

sandstones. The

millo,

following

J.

more

Hernández, of

departments

important

and

Boyacá

genera

the authors

and

in

and

the

species

lindenii

Podocarpus

Quercus

humboldtii

Pilea

R.

limit

Jaraof

the

(scarce)

Rhamnus

Siphocampylus

Gunnera

Piper

Weinmannia

Miconia

Bocconia

Berberis

Panopsis

Gaiadendron

Cedrela

Clethra

Hedyosmum

Palicaurea

Rapanea

Persea

Myrica

etc.

list

of

species,

altitude

in

Quercus

the

lindenii

and

Quereetum

genera

on

(dominant!)

Vaccinium

Clusia

by

sp.

Carica

Vallea

the

Soralea

lepidota

Cinchona

A

found

near

Santander.

Quercus

Tibouchina

were

Quercetum

the

made

Alto

de

between

Onzaga

(sp.div.)

Weinmannia tomentosa

fimbriata

Vernonia karstenii suaveolens

Panopsis Eugenia

Rapanea

guyanensis

Diplostephium

Rapanea

ferruginea

Berberis

Scheflera

Symplocos

Macleanea Gaiadendron Palicaurea

Rhamnus

punctatum

and

Cavendishia Clethra

stipularis

3060

(Boyacá),

Miconia

(2

Oreopanax

spec.)

3450 is

metres

of

the following:

276

Gaultheria

Psychotria mutisii

Persea

Brunellia

Viburnum

Podocarpus

oleifolius

Symbolanthus

Compositae

indet.

Bomarea

Myrsinaceae

Hedyosmum

bonplandianum

indet.

indet,

Myrtaceae

Passiflora

Blechnum

Cestrum

Elaphoglossum

Aethanthus

Polypodium

glaucophyllum

Thernstroemia

Polypodium

sp.

From

this list

have many is

in

present not

cetum, de

it

will be

genera and the

and

Bogotá

in

Quercetum ;

single

a

clear that the

species

while

but is

Quercus

found

Vaccinion

Near the tree-line

Cuatrecasas

with the

Vaccinium

floribundum

OF

the

THE

is

at

the

dwarf forest

species

Senecio

floribundi

Hypericum

summa

Tibouchina

etc.

J.

by

dominant

Miconia

in

Associations

Hernández

similar

or

more,

and

or

This

present.

from

small

m.

the

Eastern

trees).

small

of

this

the

associations

wind

in

grossa

found

are

protected

at

type

authors

laricifolium anastomosans

salicifolia

Miconia

hartwegii

or

be

lanatus

Miconia

walls

Quer-

Sabana

Weinmannia tomentosa

obtusifolia

identical

may

3200—3300

at

(shrubs

Gaultheria

metres

the

the

associations

eleaoides

Almost

(various species)

dominant of

PARAMO

and end

m

Vaccinion

dominant

following

related

of

3000—3200

chrysoleuca

3600

the Weinmannietum

Weinmannietum of

Miconia

Hypericum

the

ANDEAN

and

floribundi

describes

(1934)

Cordillera,

Clethra

in

and

Weinmannia

Quercus

transitional belt

a

begin

may

Hesperomeles

Even

surroundings.

DWARF-FOREST

irregular belt

Quercetum

common.

altitudes "Páramo

altitudes

Palacio",

up

natural

3500—3600

between de

at

behind

valleys,

may

m

to

rockfound

contain

as

species: Diplostephium

summa

Hypericum Vaccinium

sp.

Valeriana arborea

sp.

Rib

floribundum

es

Berberis

Aragoa

Weinmannia is

absent.

practically

Polylepietum A at

special type

times be list a

of

dwarf-forest is

altitudes rather far above

of

a

true

species

Polylepietum

Guantiva".

high forest., was on

made

with

by

moraine

the

Polylepietum,

the local forest-limit.

R. soil

trees

up

to

Jaramillo, at

an

7 —10 J.

This m

found often in patches association

and

Hernández

altitude

of

3400

and m

may

The

more.

the

in

some-

following

authors

of

"Páramo

de

277

Polylepis

(dominant)

boyacensis

Hypericum

mexicanum

Hypericum

sp.

Syphocampylus Gaylussacia Bidens

rubifolia

Senecio

Befaría

Arcythophyllum

Bomarea

Clethra Vallea

columnae

buxifolia

Chusquea Espeletia

verrositae

Aragoa

Espeletia

glandulosa

Weinmannia

Niphogeton

Miconia

Halenia

stipularis

Cerastium

Bucquetia Berberis

Paepalanthus

Vaccinium

Ranunculus

Geranium

Ilex

Eryngium

Gynoxis Macleanea

Excremis

rupestris

Hesperomeles

Rubus

Symplocos

Blechnum

Pernettia prostrata

Jamesonia

Gaultheria

Elaphoglossum

Tibouchina In as

the

real

a

Nevada

Sierra

forest,

4000

to

up

del

Cocuy

found

we

a

Polylepietum,

developed

metres.

Escallonietum

the

In dwarf'

altitude of J.

2.5

3300

m

the

The

following from

(dominant)

fastigiatum

de

(Páramo

soil

peaty

on

authors

myrtilloides

Eupatorium

Visitador"

high,

metres.

Hernandez and

Escallonia

del

"Ciénaga

forest,

(marsh

list

of

this

species

K.

by

of an

Jaramillo,

Halenia Cortaderia

strutheolaefolium

Cyperaceae

Hypericum

sp.

Geranium

Galium

Xyris

Paepalanthus

Cerastium

andicola

Lycopodium Festuca

Puya Gentiana

In

(2

spec.)

places

with open

water:

Marsilia

Potamogeton

MEADOWS

From

3200

areas

abundant this

made

at

Niphogeton

(codominant)

Hypericum

large

was

type

found,

was

Escallonietum:

Senecio subruncinatus

Vicia

another

Guantiva)

vegetation)

belt,

or

are

AND

3300

m

occupied

occurence

besides

“BOSULICAULON”

the

of

upwards by

the

associations

different

grasses,

(in

OF

species

small

shrubs

ANDEAN

Eastern in

of

THE

Cordillera

general

Espeletia.

are

still

PÁRAMO

near

characterized In

the

abundant,

lower

like

Bogotá) by part

the of

Hypericum,

278

Gaultheria, in

the

become

Vaccinium,

Pernettya,

higher the

the

parts

grasses,

Arcythophyllum, specially

Diplostephium, and

Calamagrostis

etc.;

Chusquea,

but soon

dominants.

Espeletietum Quatrecasas peletietum

(1934)

(3300—3460

gives

the

following

list

from

calamagrostiosum

argenteae

argentea

Senecio

Espeletia

grandiflora

Halenia

anastomosans

Pernettya

Arcytophyllum

aristatum

Lupinus

paniculatus

Geranium Azorella

glaucescens

multiceps

multifida

tenera

Paepalanthus

Cosmos

chrysanthemifolius

Acaena

Ranunculus

candoleii

de

Festuca

ensifolius

bogs of

tracts

are

with

Palacio",

caespitosum

abundant in In

ground.

many

the

peat-bogs,

the

humboldtianum

Espelction

higher the

Calamag rostis

following

this

plants

Sphagnum

belt

in

the

covers

"Páramo

abundant:

are

Bomarea

Festuca

Lachemilla

Geranium

Ranunculus

Halenia

Paepalanthus

Gentiana

Jamesonia

Bartschia

Blechnum

“Valeriana”

Valeriana

stenophylla

Elaphoglossum Polypodium

sp.

Rhizocephalum Although same

and

belt,

of

parts

Stachys

Chusquea

the

effusa

sp.

Paspalum

very

the

ocisapunga

Calamagrostis

humile

Peat

large

Cerastium

sessiliflora

Paepalanthus

ruiziana

Orthosanth us

hybridus

Eryngium

pauana

Rhynchospora

piloselloides

Hypochaeris

karstenii

cylindrostachya

Halenia

peruvianas

Rhizocephalum

caulolepis

glutinosa

Lobelia

Erigeron

Es-

gallioides

Jamesonia

mutisiana

Castratella

his

Guasca"

pedicularioides

Elaphoglossum

pentlandii

Niphogeton

de

asclepiadea

Alchemilla

laricifolium

Gaultheria

from

formosus

Bartschia

phyllicoides

Diplostephium Hypericum

species "Páramo

m).

Espeletia

Gaultheria

of

the

Sphagnum no

Isoetes

was

found,

spores

of

this

plant

are

very

common

in

sites.

Acaenetum

A

species de del de

very is

Onzaga, Cocuy Santa

curious

found in

association, páramos

Páramo de up

to

Marta

an

up

Acaena

the

an

of

more

altitude of

cylindrostachya

Quercetum

Guantiva, etc.).

altitude to

with

above

We also than

4300

4000 m.

as

forest-limit

found it m,

and

in in

in

the

dominant

Boyaeá

(Alto

the Sierra Nevada the

Sierra

Nevada

279

A.

Although limit

upper

be

it

in

lacking The

in

some

of

cylindrostachya

the

this

Boyacá),

association

Acaenetum

at

Acaena

cylindrostachya

Acaena

elongata

Orthosanthus

the

with

of

majority Alto

The

cases.

Onzaga

and

humus,

Nevertheless

following

(department

of

Senecio

chimboracensis nitidum

Clethra Gaultheria

coartata

Weinmannia

(dwarf-form)

Pernettya Hypericum Lycopodium

Lachemilla

Festuca

(Castilleja)

Calamagrostis

(Befaria)

Jamesonia

Sometimes

Espeletia

become

may

and

composition,

Espeletietum

in

that in

But

acaenosum.

the

association

the

general

without

abundant,

more

case

Espeletias

are

the

changing

could

be

called

scarce,

an

comple-

or

lacking. Exact

3300

indicates

and

Ecuador

for

absolute

with

234

rain

at

closely (fig. 3)

A.M.

2.9°

Under upper,

most into

zone

tree-line

account

protected Bogotá

higher

at east

slopes

approximately more.

cipitation

On and

(Quercetum!).

the

3300

tne

limit dwarf

de

metres,

frequently

but

the

in

de

on

Onzaga

clouded),

belt.

the

For or

it

90. de

tem-

same

Ecuador)

Cundinamarca

region.

our

or

this

an

understand

high

more

limit

are

here

upper-

not

taken

dwarf forest in small wind

de

lies

at

±

flanks

lies

the

at

high

de

Eastern

metres.

tree-line

(wet!)

at

Sabana

the

(in

3250

the

apparently

forest-limit

of

Palacio

(dry!)

eastern

(with

In mm,

"Páramo

(in

m

will

we

metres

surroundings

Cocuy

the

the

3—5

Páramo

Bogotá)

1071

the

for

m)

15° C.

temperature for

limit")

±

forest

In

3600

relation

a

in

and

P.M. at

of

and

observation

between

(3600

author,

same

193-1)

SNOW-LINE

forest

of

tree-line

the

8.°°

at

high

too

forest

metres.

Sabana

de

the

4° C

AND

of

Sierra Nevada

Alto

to

oscillation

varies between 80

days

be

to

isolated stations

the

daily

;

temperature

average

seems

and

The

1.5° C

minima of

than

higher

Cuatrecasas oscillate

humidity

of

in

temperatures which

according

m,

altitudes

absolute

m)

8

the

(quoted

altitude-temperature

closed,

etc.

of

is,

"altitudinal

(or

3200—3300

of

(3600

TREE-LINE

forest

valleys

lies

Cordillera, western

the

3650

our

m

m) ¡

during

6.5° C

4450

at

principally of

of

with

5.

the

1958

April

corresponds but

16° C

precipitation

altitude

an

7.°°

at

perature

in

of

for

Espinosa

monthly

(4150

The relative

days.

made

Palacio",

2.9° C

máxima

the annual

We

average

and

m)

not available

are

Eastern Cordillera.

in the

(3600

Cotopaxi

data

meteorological

metres

6.5° C

at

to

3450 metres.

Carex

or

little

dunging.

the

de

Espeletia

tely

the

seems

Geranium

(dominant)

Paepalanthus

specific

above

páramos

Acaenetum

very

and

grazing

the

on

approximately

of

soil

rocky

on

mexicanum

Hypericum

the

the

Bogotá,

Miconia

Arcythophyllum

Excremis

in

made

was

altitude

an

found

mostly

be related with cattle

to

natural

a

in

present

region.

seems

this

be

may

Weinmannietum around

Acaenetum is cases

presents

list

of

On

was

the

found

3500 metres annual

3500—3600

pre-

metres

280

In

the

line lies

Central Cordillera 3800

at

del

the Nevado

of

nounced

dry

personal

hundreds

principally 1°

Temperature

2°

Humidity

Another almost in

tation

on

and

highest

the

lowest

taken

not

those

into

and

is

and

In

those

its

be

forest-limit).

the

following

If

year.

regions

way

depends

marked

dry

the

seasons,

the

overcast).

this

as

(forest of

factor dwarf

or

forest

3900( —3800)

at

semi-desert

or

of

vege-

and

distribution reaches

tree-line reaches is

precipitation

tree-line

is

metres,

conditions

are

the tree-line, between

precipitation

the

season,

annual

distribution

:

Andes lies

annual

its

lack

tree-line;

precipitation

the

and

continuous

importance,

differences in altitude annual

on

the

(desert

metres

be

due to

the

dry

the fact

predominance

that

show

season,

("greenhouse

is

(or

will

We

of

of

low

reduced

of

this

maximum its

and

highest

there

its lowest

to

having

areas

higher

which

higher

a

annual

average

skies prevents

overcast

effect")

relatively

more

at

two

pronounced

The

combination of

one

or

be

"humid

dryness"

two

are

level

than

annual

tempe-

heat losses the

compensates

surface forest

by

here

climate". or

is

the

present,

the

I

o

first and

of

Quercus

are

has

not

apparently and

case

in

2°

conditions

(cloud-forest!) second

the

case

a

association). of

altitude 3900

(to

(=t

m

the

metres,

cloud

forest

(Quercetum

or

and

the

minimum altitude

in

the

"Sabana

etc.)

800(700 —900)

annual

mm

de

of

Bogotá"

precipitation

The

major

a

dren

and

in

quantity

seasons

the

contrary

of

and

annual

of

following,

will

be

a

precipitation,

more

with

indicated

the absence

continuous low

the

term

with

the

overcast,

"humidity" term

or

"relative

"dry climate". species

("eselero a

In

types.

(Weinmannietum

3200

species and

dry stations).

pronounced

indicated

Several

3800

forest

conditions)

difference between

maximum

at

certain

immigrant

young

association)

related

the

dry

the

area),

where

country

different forest

related

a

(or

take

the

relatively

a

by

associations)

(actual

will

occur

found several

the tree-line

a

páramo

the

marked

maximum

Quercetum

related

of

of

local

only

extensive

below

dry

places

indicated

Weinmannietum

and

has

it

more

Where

regions

reached

the

that

precipitation

above

in

may

no

radiation

(Quercus

a

even

may

be

insolation.

present

may

still

pro-

Kraus,

m).

In such

ground

lower

but

all

Local

no

annual

in the Colombian

the

pronounced

precipitation

by

clear

far

depend

These variations

ratures.

will be

valleys

it

account).

during

(3200—3000

of

3200( —3000)

at

there

two

or

in

tops

(3800 —3900 m).

limit one

wind,

tree-line

values,

two

precipitation values

is

could express

The

it

"greenhouse-effect"

year;

effect

windswept

One

forest

may

tree-

slopes a

(Erwin

metres

indications that

dwarf forest

without

round,

year 3900

at

the

(altitude)

wind-protected

forest

the

On

1934).

up.

(amount

the

factor

equal

all

the

humid climate,

more

factors:

two

on

a

Cuatrecasas,

even

are

of

patches

data mentioned above

during

of

rains lies

and there

higher

metres

the

it

tree-line

Small

metres.

of

From

is

the

communication) ;

4000

to

up

del Tolima;

where

Huila,

season,

with

places

on

(Nevado

metres

of

the

humid

meso-microfilas")

smaller leaf surface

forest

are

characterized

and several

species

of

("eselero micro-nanofilas").

by the

a

greater

relatively

leaf-

dry

281

The Kraus the

on

snow-ine in

(personal

flank

west

of

between

change

any

the

de

Laguna Pan

de

and

his observations

give

the West

on

la

Cordillera

flank

of

del

1943.

Nevado

Pico

la

de

de

de

1943

4520

■1520

4520

4500

4650

of

additional measurements

the

metres.

4570

The

In

If

metres.

de the

(or

we

take

m,

at

mentions 1943

flank

Euiz

the SW

on

was

The

at

Tolima) 3800

4320

1948

4720

and

by

in 1959

3900)

on

in

4150

the

to

be

a

the

on

Nevado

same

following

on

E

de on

year

the

the

has

altitude

an

conclude that

Cundinamarca

of

where

figure.

the tree-line lower.

considerably

found

on

the NE

4150

flank

(Quatre-

m.

1934).

Isabel

W

extreme

the

(in

may

flank

flank in Santa

the

observations:

was

W

Sierra

average

trustworthy

most

Cordillera,

del Tolima and

m

m

the

the

approximately

ago,

we

the

the tree-line. in

This

is

Sumapaz

data,

of

an

in

representing

C,

the snow-limit may lie

the Nevado

on

line 0°

central part

the

Central

with

us

these

probably

1959.

decades

some

seems

of the

metres,

flank at

all

in

the

is

exercised

Kraus

by

for

del

1948

position

us

by

given

for

metres

provided

and

m

authors

have

the

on

the Nevado

account

the

average of

flank

found

was

of

the

the

on

Nevado

del

the

is

of

these

Cordillera

on

two

4350

on

the W

4100 metres,

figures

for

Central

observations

m;

flank of

and in

is

the

the tree-line

on

4100

West-flank of

4150

only

the Nevado del Huila

1944 also at

metres.

The

the

Central for

average

1943

on

this flank lies according

to

made

in 1934 and

(tree-line

at

m.

group of the

the

East

Nevado

Quatrecasas

m.

is

with the at

into

4500

found at

(based

It

that

snow-line

4200

at

Nevados

del

and

data obtained

altitude

Nevados

In 1940 the snow-line

flank

4600

4200.

at

3900!)

and

approximately

the

the

cannot

influence

all the

surroundings)

In 1943 the snow-line

In

1938

which

lost its snow-cap

4550

at

of the

Kraus

know

we

and

even

Erwin

4200

of

1938

which

snow-line

Bogotá

region

3800

Mr.

W

4550

m

between

important

theoretical

Moreover

theoretical

casas

1948

4650

by

altitude and average temperature (fig. 3)

Cundinamarca),

(Sabana

4325

and

made

were

fluctuation

between

Cocuy

metres.

4560

at

m

snow-line

climatic

few decades any

a

is

at

1946

m

therefore took the average

relation of

lies

4390

m,

of

minor

a

del

a

4425

4325

Nevada

of

made

were

4660

retirement

course

of

he

glaciers).

4780

4600

Plaza

la

of

SE

(and

4740

4640

Azúcar

The

4550

Nevado

same

snow-line

area

effect

We

without

Sierra),

the

the observations

1942

Plaza

following

Pan

All

la

occasions

Cocuy.

Castillo

Northern

Cerros

the

several

4690

del

The

of

1938

Sierra

Azúcar

del

de

Erwin

higher.

or

on

in

places

here in tabular form.

the

m

metres

(Laguna

other

general retiring

a

4500

at

4520

Cocuy On

Ritacuva Cerros

lies

measured

Nevado

1938 and

found different values We

Eastern

the

comttmnication)

interesting

m.

advances,

that

Espeletia-limit It and

seems

may

on

this

probable be

last

mentioned

the Nevado del to

us

dependent

figure

that the snow-line on

corresponds

very

Tolima, given by Quatrecasas

small-range

continuously

solar

cycles.

closely (1934),

retires

The

and

average

282

extension of upper

From

the above

the

By between

that

and 4150

a

As

tree-line of The

with

still

a

3900

line may

The

each

caught

was

and

pollen

during

in

general

Van

glass slides

with

of

the

one

same

Sabana

de

that

Bogotá,

reason

The

la

Viga",

of

one

general,

the

same

interesting

most

the

to

a

east

station

of

Bogotá.

of

3450 It

the

These

shows

a

the

Podocarpus

0.4%

23.5

less

9.9%

extense

be

a

site

in

the

of

forest

results

As

of

including

we

Pleistocene

of

the

the

almost

are

of

origin

forest

much

by

higher

destroyed, by

diagram, all the

man.

than

vegetation.

human action

"Páramo

del

Alto

de

percents:

51.5

%

48.5

%

%

0.2%

approximately

páramo-area.

found

the

total

calculating

the

influenced in

weeks

the

0.2%

plants

Total

at

not

Recent

two

or

%

5.7

Rapanea

%,

and

human of

Bogotá

0.1%

Drimys

23.5

one

Summing

general

natural,

de

4.3%

Myrica

interesting

where,

0.6%

Urticaceae

forest-plants,

of

is in

following

3.6%

Weinmannia

most

say

and

theoretical

(1960).

pollen-spectra

altitude

Alnus

Hedyosmum

of

Sahana

spectra

destruction

and the

metres

Bocconia

The

This

Perico

stations

meadows

the

with

Miconia

pollen

may

metres,

the line,

to

the

&

fossil

Gramínea e

Total

4550

meet,

stations.

these

for

by

site

in

pollen

Gramineae

spectrum,

at

snow-line

Hammen

extense

the percentage of

correspond

is

in

or

done

the

by

Cordillera,

we

metres.

correspond

different

obtained.

were

of

glycerine jelly, exposed

each

as

way

influenced

heavily

in

der

six

at

year year

Central

POLLEN-DEPOSITION

by

whole

"Sabana de

metres.

atmospheric

of

treated

pollen-spectra

rather

would

on

a

deposition

percentages

For

is

4150

the

influence,

snow-line

a

forest-limit and 4000

of

of the

same

would

forest-limit

RECENT

deposition

recent

to

snow-line of

a

approximately

at

surroundings,

pollen

to

highest

the

the

fluctuates

the snow-line

surroundings

on

coresponds

metres

greater humidity,

lay

depends

with

regions

conclusions.

following

group of Nevados

(central

certain

minimal extension.

"humidity", the

(in

metres

of

years

the

lesser

or

tree-line

6.

and

greater

metres

theoretically

the

draw

now

4550

3200

then coincide in

during

may

metres

the

tree-line of

a

we

influence of

flank).

would

snow

visible

approximately

Bogotá") West

the eternal

Bspeletia-limit,

250

will

part

a

see

of

this

100

spectrum

percentage metres

below, the

%

above

almost

Hedyosmum the

almost

pollen

the

are

of

closed

identical

diagram.

50

as

tree-line spectra

%of

high in are

as an

to

283

Another in

the

of

centre

Alnus of

the

the

trees, is

forest Even

with

with

forest,

is

ation, In

the

midst

the A

tree

still

rather

scape

at

78

of

%

3330

of

In

is

of

1

to

was

inches In

inch

of

from

in 2.3

the to

was

these

The

collect

to

of

veget-

(56%

of in

Alto

Culture

com-

influence and

which

Viga",

the

partly

so

partially culture

de la

%.

12 in

this

Gramineae), a

the

found

by

of

pollcngrains

wind.

be

found

only

surrounded

metres,

forest

to

distance)

km

was

of

(Acacia,

trees

(only

3.5

the

del

and

is it

land-

showed

From

30

few

and

of

the

is

above and

SEDIMENTS

boring

sea-level.

the

from

a

to

a

dia-

a

metre

mud.

In

down

metres

first

the

1;

1.5

or

metres

the

with

continuously

2.5

212

from

to

water

of

fig. with

1

every of

depth

down

distance

in

Cores

up

use

out

carried

was

indicated

drawn

metres

metres

coring

THE

site

part

recovered,

will

spectra

second

a

hole,

with

core-barrel

of

metres

four

core

the

the a

of

C

14

uppermost

hole

2.5

Besides

deep.

samples

a

and

metres,

excavated

was

at

the

continuous

collected,

were

of

obtain

moreover

2

boring-site, of

series which

X

samples

the

upper-

charcoal. of

cores,

the

but

the

to

more

coring, exact

real

amount

interval

of

cores or

of

the

were

compressed,

less half

of their

compression samples

for

was

in

the

original

core-

length.

measured,

in

taken

pollen-analysis

cores.

real

5.5

AND

without

dry,

pollen

recent

diagram.

exact

metres

was

from

4

loss

no

sample

uppermost

Only

the

first

The

done

these

from

of

continuously,

a

analysis,

the time of

know

to

at

equipment,

C14

each

Por

was

metres.

consisted

during

order

the

station

pollen

at

small

by

CORES

the

2560

taken

out,

analysis,

There

barrel,

this

diameter.

wide,

three

1957

section

32.2

carried

for

pollen

most

of

were

complete

order

metres

for

forest of

Eugenia)

in

present

vegetation

2880

site

drawn

equipment.

Coring

inner

material 2

the

patch

the

trees

Vallea,

culture

(16 %)

the

patch

THE

approximately

"Portadrill"

a

2

a

depth

a

boring

this

the

80%,

nearest

the

of

interpretation

BORING,

mining

approximately. way

the

beginning

small

elevation

this

in

natural

the "Pรกramo

near

(Ilex,

hardly

show

far

very

conclusions

for

THE

the

a

meter

the

below

7.

with

metres

from

dominating

production

meadows,

altitude

a

in

producing

surround this

spectrum

The

Alnus,

Apparently

(in

pollen

%.

the

at

which

station

Gramineae.

pollen

Weinmannia

strong

highly

the

placed

was

Gramineae.

Several used

of

an

station

another pollen

at

at

10

Sabana,

transported

not

are

apparently

be

Cita,

pollen

from

Weinmannietum.

the

percent.

the

the

of

Gramineae

trees

percent

%

in

culture

the

surround

high

La

near

destroyed

is

the

and

%,

rather

station

mon

8

hills which

few

station

the

by

Gramineae.

low

almost

Universitaria",

Eucalyptus)

of

very

by

of

38

influence

a

the

culture meadows

the

80%

by

represented

"Ciudad

in

only

the

in

this

almost

evidently

an

represented

are

is

greater

Suba,

near

of

visible

from

the spectrum

by

Although

surrounded

completely

influence

clearly

shown

Subachoque.

near

forest,

is

fact

interesting

very

Alnetum

metres, the

interval

2.3 and

metres, 15

uppermost

cm

32.2

used 10

(see

cm

from

5.5

metres

pollen

(or

in

to

(first

diagram)

some

32.2

cases

was a

7.5

little

or

15

less)

cm

from

metres.

boring;

C. U.

X.)

have

yet

been

284

and

analysed,

continuation

will

The brick.

a

contact

yellowish

From

m

and

depth

a

The

intermittent

in

certain

and

of

From

Diatoms

with

algae of

of

this

main

the

in

as

graph the

habitat

of

and

m

No.

I,

37;

C.U.

X.„

1:

PI. No.

3.00

—

like

algae

the

sediments

these

clays

of

the

went

means

to-day

occur

"inundation-clays". is

lake-sediments with

clay-gyttjas)

depths

moment

by

on

still

(5.70,

present.

rather

a

fine

and Pediastrum) ,

This

present.

In

179b;

of

those

present

28.60

this

high

plant-detritus

12.90,

peat

deepest is

sand

algae

nor

of

percentage

we

18.45,

contains

18.75,

little

a

X.,

No.

a

few

183;

PI.

I,

7:

No.

PI.

(Pl.

37;

C. U.

2:

I,

PI.

X.,

at

also

So

we

m

peaty, A

m.

small

a

(32.20 m) 0.60

uppermost rest

have

to

few

layer

ash.

C. II. X.

the

the

partly

24.00

volcanic

boring

spores,

THE

the

them

of

of

layer.

peat

contains conclude

m

some

that

lake-deposit.

a

the

I,

diagram represent here

No.

shortly,

be

and

given

species,

first

give

about or

genus

records

a

the

photoaspect,

family.

1—7)

C. U.

I,

SPECIES

will

details

conditions

present,

layer

while

Isoetes.

is

layer

describe

will

C. U. X.,

is

contains

clay-gyttjas)

the

m.

30.50—30.85

point

Isoetes

(or

above

29.00

present;

mentioned in

addition,

lake-clays

layer

from

diatomite

the

to

clayey

species

and

humic

and

samples,

ecological

4:

series

DESCRIPTION OF

the

C. U.

I,

of

at

up

(which

higher than the peat

fine-sand

the

as

black

a

Quercus PI.

call

series

is

layer

28.30,

much

increasing

each.

from

but

pollen,

dried

seasons

various

type

24.85,

contain

pollengrains,

of

is

particles

and

important part

lake

(or

continuous

a

same

downwards

grained

part of

many

fossil

m

some

8.

As

charcoal

sedimentation

will

We

peat

a

31.00

do not

an

from

m.

occasionally.

wet

lake-clays

intercalated; m

fine

layer

and

algae

is

31.00

of

at

content

found

diatomite

layer

to

clay

were

the

present at

the

found

29.00 a

From a

of

again,

From and

But

Botryococcum

m

24.00—29.00

were

level

consists

extends

and

clay,

this

This

absent.

are

present

At

m.

dark

m).

22.25—24.00

algae

found

continuous

a

of

and

sand

rather

1.65—2.00

with

grey

of

a

layer.

veins,

abundant

an

Sabana

during

mainly

are

form

horizon.

m

(especially

contain

only

Sabana.

the

22.25

fill of

0.85

from

clay

gray

succession

clay.

the

that m

brown

yellowish

a

which

were

m,

3.45

are

19.65

m

of

under-lying

especially

humic

a

blackish

is,

diatoms

pollen;

From

is

3.45

to

algae

19.05 and

elay;

the and

analysed

pollen-diagram).

consist

depth

a

the

occur

the

man-made

recent

until

with

3.05

to

Pediastrum,

of

3.45

sediments

content

be

sedimentary

of

sediments

yellowish

depth

inundations

parts

Prom These

of

diagram

will

Y.)

the

of

very

mentioned above

the

by

with

m

explanation

represented of

of

a

veins,

particles

humic

sediments

Botryococcum below

The

part.

C. U.

column

natural

present

2.65

from

strongly

a

is

clay,

2.35—2.65

Then

present.

of

layer

brown

charcoal

m;

description

uppermost

sharp

grey

The

that

to

boring;

stratigraphies!

a

the

reddish

0.85 —2.35

3.45

is

m

this

with

clay

rather

of

(second

short

a

the

(see

0.20

top

Below

grey

corresponds

section

give

now

C. U. X.

boring

diagram

this

later.

published We

the

of

5:

X.,

No.

C. U.

A-150.

183;

X.,

PI.

No.

I,

3:

179b;

C. U. PI.

I,

X., 6:

285

scabrate

Tricolporate, be

but

distinguished,

ponds

the

to

of

type in

Dominant

C. U.

8:

I,

Vesiculate,

often

(55 —70 micron; The

and

the

large

of

another

the

is

in

the

more

C. U.

10:

II, No.

X.,

one

in

in

and

duction

±

but

13:

around There

tried out

and

the

CU. X.,

psilate,

the

least

samples.

small

3500

type),

is

Other

metres.

C. U.

No.

X.,

PI.

88;

II,

12:

(Some small irregular

grooves

is

climber

Quer-

and

temperate

woody

a

found

equally

in

the

belts.

tropical

of

the

Woinmannictum

The

pollen

pro-

No.

(Pl.

184-A;

25—40

±

II,

PI.

13—14) 14:

II,

micron.;

C.

pores

No.

U.X.,

with

184-A.

annulus;

"granulate"

pores.

two

are

is

important two

species,

M.

species

by

shrubs,

or

related

and

parviflora means

of

M.

We

pubescens.

with-

but

size-statistics,

common

both

in

PL

II,

15:

Periporate, wide.

the

Quercetum

and

the

X.,

No.

Weinmannictum

associations.

Styloceras C.U.

No.

X.,

reticule,

Number of

Styloceras

pores

±

40—50

II, II,

micron.

15—16) 16:

C. U.

The

muri

are

one

179b or

granula

two

12—25.

±

laurifolium

(Pl. PI.

150;

(Willd.)

11. B. K.

is

of

tree

a

the

andean

forest

general. Bocconia

PL

II,

17:

Periporate, frutescens

L.

is

Ill,

(Pl.

II,

17)

PV4aII.

reticulate, a

very

±

30—40 mieron.

common

tree

Juglans PI.

20:

other

the

success.

Trees

in

of

I,

146—153

(anomophilous).

differentiate the

to

the

PI.

micron; No.

but

type

10—12)

11:

H. B. K.

Hedyosmum

of

great

very

Triporate, zone

II,

II,

30—48 micron.

bonplandianum

associations

II,

(Pl.

PI.

88;

Myrica PI.

in

at

82

(±

C. U. X.,

grains

small

pollengrain).

Boyacá;

is

No.

X.,

(?), Clávate.

the

Hedyosmum cetum

large

to

up

distinguished,

relatively

87.

of

area

may

corres-

different associations.

C. U.

Inaperturate in

1:

159.

be

one

samples

common

Boyacá,

Hedyosmum PI.

the

No.

may

is

relatively

(with pollen

in

Quercetum in

the

Don

D.

types

I,

cloud-forest.

X.,

types

There

dominant in

general

oleifolius

found

are

and

8)

is

C. U.

Different

impossible.

I,

PI.

I, 8—9)

9:

I,

was

PI.

small type

element

species

micron.

type

Podocarpus an

55—82

±

difficult.

representing

(Pl.

PI.

184-A;

determination

the specific

9).

No.

X.,

Different

micron.

lindenii A. D. C.

Quercetum,

Podocarpus PL

28—48

determination is

specific

Quercus

the

±

(inord.).

the

18:

C. U.

X.,

No.

(Pl.

201;

Number of

shrub

or

III,

PI.

of

pores

±

6.

the andean forest

Bocconia

in

general.

18—20)

Ill,

19:

C. U.

X.,

No.

201;

PI.

Ill,

PV9aII.

Periporate

(heteropolar),

psilate,

±

35—55 micron.

Pores

on

the

equator

286

(like

stephanoporate

a

Juglans,

species

of

andean

forest.

and

grain)

on

which

trees,

21:

Ill,

No.

C.U.X., micro

Tricolporate,

related

(intra ?)

found

in

above

the

associations

the

of

a

very

in

the

Certain

micron.

tree

common

very

Eastern

Cordillera.

species

be

may

of

the

Weinmannietum

Several found

species

also

are

considerably

even

"tree-line".

Rapanea PI.

several

are

common

21)

11—15

reticulate,

is

the

Quercetum.

III,

not

177.

Weinmannia tomentosa L. f. and

(Pl.

There

hemispheres.

apparently

Weinmannia PI.

of the

one

are

22:

Ill,

Stephanocolpate (Sometimes

No.

C.U. X.,

(4

III,

(Pl. PI.

124;

furrows),

22—23) C. U.

23:

Ill,

±25—30

psilate,

X.,

No.

132.

Polar

micron.

large.

area

tricolpate).

Several

which

species,

trees

are

shrubs,

or

in

frequent

are

the

andean

forest.

Symplocos PI.

24:

Ill,

C. U.

foveolate

Tricolporate, view

(in

polar

not

always

the

grains

(-psilate),

or

small

25:

Ill,

Tetrads,

of

trees

CU.

29:

the andean

IV,

C.U.

No.

27:

micron.

Furrows

Small

short

very

transversal

furrows,

(Pl.

III, PI.

132;

a

relatively

IV,

26:

IV,

45—50 micron.

±

Pl.

25;

26)

C. U.

No.

X.,

The separate

70

grains

in

tree, especially

common

monoporate.

are

the

higher

forest.

C.U.

No.

X.,

No.

X.,

PI.

A-154;

(Pl.

183;

IV,

IV, 27—31)

PI.

30:

28:

IV,

C. U.

C. U. X.,

No.

X.,

No. PI.

184-A;

PL

A-154; 31:

IV,

IV,

C. U.

X.,

184-A.

Tricolpate, ferentiate

on

clávate,

base

and shrubs

Trees

siderably

higher

of

±

the

of

the

than

micron.

30—40

PI.

pollen. andean

the

forest.

32:

IV,

C.

Heterocolpate, There

are

which

belt), species

grow

not

possible

the

majority of

basis

to of

19

the

the

in

the

Certain

difficult

are

type

species

of

Ilex be

may

dif-

to

kunthiana.

found

con-

in

the

andean

the

pollen

diameter

Melastomataceae of

of

X.,

our

of

grains We

Miconia

(in

(and

basis

of

"tree-line".

Melastomataceae.

species

C. U.

forest

the

on

above

the

33:

IV,

183.

No.

micron.

differentiated

other

all

species

is

IV, 32—33)

PI.

13—19

considerably

micron

the other

species be

(Pl.

183;

±

differentiate

their size:

than

while

psilate,

many

even

No.

U.X.,

cannot

The 28

IV,

"tree-line".

Miconia PI.

less

40—45

triporate).

andean forest.

Ilex PI.

24)

the

No.

X.,

reticulate

granatensis L. f.,

Drimys of

±

be

to

seem

Drimys

zone

III,

visible.

Shrubs

PI.

(Pl.

142.