Above-Ground Biomass Structure of a Chusquea tessellata Bamboo Pramo, Chingaza National Park, Cordillera Oriental, Colombia Author(s): Gerrit

J. Tol and Antoine M. Cleef Reviewed work(s): Source: Vegetatio, Vol. 115, No. 1 (Nov., 1994), pp. 29-39 Published by: Springer Stable URL: http://www.jstor.org/stable/20046500 . Accessed: 07/03/2012 20:16
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1994. Vegetatio 115: 29-39, 1994 Kluwer Academic Publishers.

29 Printed in Belgium.

Above-ground

biomass

structure

of a Chusquea

tessellata

bamboo

Chingaza National

Park, Cordillera Oriental, Colombia

of Amsterdam, Kruislaan

p?ramo,

Gerrit J. Toi & Antoine M. Cleef

Hugo de Vries-Laboratory, Department 1098 SM Amsterdam, The Netherlands
Accepted 10 February 1994

of Paleo/Actuo-ecology,

University

318,

Key words: Bunch-grass

p?ramo, Leaf Area

Index, Leaf distribution

Abstract The present study was carried out in the bamboo (Chusquea tessellata) p?ramo of 'Parque Natural Nacional de 1987 to April 1988. Above-ground biomass structure Chingaza', Eastern Cordillera, Colombia from December of bamboo p?ramo was quantified in 16 plots. These data are compared with previous results on above-ground biomass structure of bunch-grass (Calamagrostis spp.) p?ramos. The total (non-living and living) above-ground biomass of a Chusquea tessellata bamboo p?ramo was low (2,625 g DW- m~2) compared to bunch-grass p?ramo. Nevertheless, higher values of standing living biomass and litter are found in the bamboo p?ramo due to the leaf shed of the bamboo. The thick litter layer may inhibit germination and growth of nearby plants. Maximum biomass is found near the ground surface. Cumulative LAI (In transformed) and height in the bamboo vegetation are related parabolically for Chusquea tessellata and linearly for bunch-grass due to differences in leaf distribution. The mean bifacial LAI of living Chusquea tessellata leaves is 2.2 m2- m~2, whereas it is 2.5 m2- m-2
for all Poaceae.

Introduction The p?ramo is present between the upper forest line (about 3,300-3,700 m alt.) and the nival belt (4,800 Its vegeta 5,000 m alt.) of the Colombian Andes. tion is characterized by bunch-grasses (Calamagrostis
spp.) and caulescent stem-rosettes by dwarf (e.g. Espeletia shrubs. spp., In azon Asteraceae), accompanied

tessellata has the widest distribution of any species in the section Swallenochloa. In the Andes this species occurs from Venezuela to northwestern Bolivia. Alti tudinally, it ranges from 2,800 to 4,300 m alt., the broadest and highest range for the section (McClure 1973; Clark 1989). The Costa Rican Talamanca bam boo p?ramos are dominated by another dwarf-bamboo species: Chusquea subtessellata Hitchcock. In the Chingaza National Park (Eastern Cordillera, Colombia) Dierikx & Kaandorp (1986) mapped 21 p?ramo vegetation communities, nine of which includ ed Chusquea tessellata. Recently Vargas & Rivera (in press) also reported on a Chusquea tessellata commu nity from the Chingaza study area. Obviously, bamboo In plays an important role in this p?ramo ecosystem. the same park, close to 'Embalse de Chuza', about 60% of the area is covered with dwarf-bamboo vege tation and themean above-ground standing biomass of Chusquea tessellata was determined at 1,419 g DW m-2 (Aristizabal & Casta?eda 1983).

al wetter locations cushion-forming plants can be found (Bosman et al. 1993). Dwarf-bamboos (genus Chusquea; previously known as Swallenochloa', Clark 1989) are conspicuous in the vegetation of wetter zonal 1978, 1981). dwarf-bamboo species studied in the area is Chusquea tessellata Munro Chingaza study is one of the largest bamboo (Poaceae). Chusquea genera in the world with about 120 species, rang to Argentina, Chile and Uruguay, ing from Mexico p?ramo (Cleef The woody the Caribbean Islands and eastern coastal and southern Brazil (Soderstrometf?/. 1988; Clark 1989). Chusquea

30
73?50'

4?40

4?40

Legend
Contour line

Park limit Road Camp 4"30' 4?30' Pluviometncai Lake station

73?50' Fig. L Location of the study site (after maps 228, 229, 247 and 248 of IGAC, Santa F? de Bogot?).

Most phytomass analyses in the tropics have been carried out in lowland forests and grasslands and only a few in highland ecosystems (Smith & Young 1987). In ECO ANDES the framework of the Colombian-Dutch research programme (Van der Hammen et al 1983, 1984, 1989 and in press), the present study was car ried out as part of the in-depth studies of the high
Andean p?ramo ecosystem. Inventory and mapping

1987; Keizer 1989; Rossenaar & Hofstede 1992; Ver in press; Hofstede et al, in press) weij & Beekman, are needed for development of an ecologically sustain able management strategy for the p?ramo (Verweij & Budde 1992; Verweij & Kok 1992; Schmidt & Verweij 1992). Thus far the bunch-grass p?ramo ecosystem has been most studied. Earlier Toi & Cleef (1992) reported
on the nutrient status data of on a dwarf-bamboo above-ground p?ramo. p?ramo. biomass The results This struc are provides ture of the first the same

of the p?ramo vegetation (e.g. Cleef 1981; Cleef & 1984; Sturm & Rangel 1985; Franco-R. Hooghiemstra et al. 1986; Bekker & Cleef 1989; Salamanca et al. in
press; Kloosterman et al, in press) and p?ramo ecosys

dwarf-bamboo

compared with those of previous

p?ramos, using the same biomass

studies in bunch-grass
estimation methods,

tem studies (Lutz & Vader

1987; Beekman & Verweij

31 at about the same altitude (3,620 m) on the opposite dry site of the Colombian Cordillera Oriental (Lutz & Vader 1987; Beekman & Verweij 1987) and elsewhere in the tropical Andes.
Temperature Precipitation

f?C)

(mm)

Methods Site description Fieldwork was carried out from December 1987 to

April 1988 in the Chingaza National Park (Eastern Cordillera, Colombia), about 35 km E of Santa F? de Bogot? at 4o 40' N and 73? 48' W (Fig. 1). This National Park covers 58,350 square kilometres. It is administered by the 'Instituto Nacional de los Recur sos Naturales Renovables y del Ambiente' (INDER ENA) and the 'Empresa de Acueducto y Alcantarilla do de Bogota' (EAAB). The Chingaza p?ramo is of as it forms themain natural drinking water importance source for Santa F? de Bogot?. A number of small and large lakes of glacial origin are found in the study
area. Below 4,000 m elevation these lakes are usual

climate diagram for Barajas, 1974-1987 (4? 41' Fig. 2. Walter N and 73? 45' W topographic map 228 (edited by IGAC), data provided by EAAB).

Plot sampling Biomass technique

a number

ly bordered by Sphagnum bogs. The study site was selected, using a 1:10,000 vegetation map (Dierikx & 1986), on a slope 2 km NW of the shal Kaandorp low lake 'Laguna Seca', at 3,650 m altitude; 9? (sd? 2.7) slope and ? 100? SE aspect. The criterion for the site selection was a homogeneous vegetation belonging tessel to the Calamagrosti? bogotensis-Chusqueetum latae community (Franco-R. et al. 1986), consisting of dense bamboo clumps up to approximately 80 cm in height. station Barajas, 5.4 km The EAAB-meteorological N of Laguna Seca (Fig. 1), is situated at the same alti
tude as the study site, and climate conditions are quite

was estimated using a destructive sampling (Milner & Hughes 1968). Every month
of bamboo clumps were randomly select

ed from dense bamboo vegetation. The clumps were sampled by clipping all standing vegetation from one square metre of clump surface. Litter and bryophytes were were collected by hand. InDecember 1987 four clumps from January 1988 to April sampled; monthly 1988, four, five and three clumps were sampled respec tively. The material was separated into: Chusquea
tessellata stems and leaves, grasses (Calamagrostis

similar and representative for the study area (Fig. 2). The mean annual temperature is 6 ?C while at night the temperature may drop below zero (?4 ?C). The cli mate is relatively humid with about 1,850 mm annual precipitation and a distinct dry season from December to the Koppen system (Koppen to February. According & Geiger 1930), the climate can be classified as EH (>3,000m). The soils are poorly developed, usually showing hydromorphic characteristics, as a result of the humid climate, and lack podzolic features (Ingetec 1970; Sturm 1978). The soil can be classified as a histosol (soils having an organic horizon of 40 cm or more) according tem (FAO 1974). to the FAO/UNESCO soil classification sys

effusa and C. bogotensis), dwarf-shrubs, herbs, ground dwelling species (Arcytophyllum muticum, Pernettya litter and prostrata and Cardamine sp.), bryophytes,
lichens.

No distinction was made between bunch-grasses and other tall grasses because the latter were just marginally present. Subsamples of all monocotyledons and dicotyledons were further separated in non-living and living material to estimate their non-living: living biomass ratio. Bamboo leaves, which abscisced when gently pulled back just after clipping, were considered non-living. Plant material, consisting of 2/3 or more green material, was considered alive. All material was pre-dried in a greenhouse and subsequently ovendried in the Soil laboratory of (105 ?C, 24 h) and weighed the Instituto Geogr?fico Santa F? 'Agust?n Codazzi', de Bogot?.

32

Chasquea Chusquea

tessellata tessellata

stems leaves

Bunch-grass

Dwarf-shrubs Lichens Herbs

Ground covering plants

?j?i

Epiphytic bryophytes j?

Terrestric

bryophytes Litter
1000 dry mass (g.m )

1200

living
Fig. 3. Mean dry mass (g- m 2) of the different plant fractions

non-living
= 16), lines indicate s.d.

unseparated

(n

Vertical

structure

In February 1988 the vertical structure of the vegeta tion was determined within aim2 plot. The vegetation was harvested in 10 cm layers, according to ground sur face. The yield was divided into different groups, dried and weighed as described above. In this paper aerial biomass is defined as above-ground biomass minus
litter.

For Calamagrostis the effusa (incl. C. bogotensis) relation mass-LAI of Lutz & Vader (1987) was used: = 98.425 cm2 1 g Calamagrostis effusa living = 109.349 cm2 1 g Calamagrostis effusa non-living It was recorded when bamboo leaves were infected with the fungus Phyllachora chusqueae P. Henn. & Lind. (det. Dr H. van der Aa, CBS, Baarn) and/or dam
aged and/or non-living. Chusquea tessellata leaves

Leaf Area Index (LAI) The LAI of Chusquea tessellata and Calamagrostis C. bogotensis) was attained, using effusa (including plants from the plot on which the vertical structure was determined. The area of all the bamboo leaves of one layer, was measured by copying the outline of
the leaves onto paper with known area-weight relation.

were graded on the degree of infection when more than two spots of fungi were detected on a leaf. Damaged leaves showed traces of insect herbivory.

Results
biomass structure

Above-ground

The area of the leaves was determined by cutting the leaf outlines from the paper and weighing them. In this manner the LAI could be calculated. It is assumed
that the somewhat vertical Chusquea tessellata leaves

The mean above-ground biomass of the 16 plots was 2,625 g DW- m~2 (sd ? 486). Ground dwelling plants and herbs consisted mainly of living material. Seven
ty five percent of bunch-grass material was non-living

are photosynthetically active on both sides, so the LAI includes bifacial leaf area.

(Fig. 3), while 78% of Chusquea tessellata was alive. During 4 months of monitoring some senescence could be detected (Fig. 4). The percentage of non-living

33
100

height of 30 cm. Epiphytic bryophytes were found to about 50 cm on the bamboo stems. Most (57%) of themicrophyllous Chusquea tessel lata leaves in the bamboo clump, were found below 10 cm due to the numerous small Chusquea tessellata stems in the understorey. Above 10 cm the number of leaves decreased with height. The percentage of undamaged leaves decreased with increasing leaf length (Fig. 6). The proportion of undamaged leaves of the longest leaf length class (111-120 mm) was exaggerated because of the small number of leaves in the sample (n = 4). The percent ages of damaged and non-living leaves remained near ly constant, regardless of their length. Small leaves were seldom infected with fungus. Longer (i.e. older) leaves were more heavily infected with fungus.

Dec. '87

Jan. '88 month

Feb. '88

| W?????

Chusquea

tessellata

LAI
The LAI of non-living bunch-grass leaves was always higher than that of living bunch-grass leaves (Fig. 7). The total area of living Chusquea tessellata leaves was greater than that of living and that of non-living bunch grass, while the total area of non-living Chusquea tes sellata leaves had a smaller LAI. For Chusquea tessel lata cumulative LAI (In transformed) and height are related parabolically. For Calamagrostis effusa (incl. C. bogotensis) they are related linearly as a conse quence of the contrasting distribution of leaves in the bunch. The slopes for non-living and living LAI run ratio parallel indicating a constant non-living:living (Fig. 7). The LAI of living Chusquea tessellata leaves was 1.13 (m2 m-2). The dry mass (105 g) of living leaves within the plot inwhich the vertical structure was deter mined was nearly half of themean dry mass of all plots (206 g). Therefore, the LAI of this plot had to be multi plied by 1.96 to obtain a representative LAI value. This short cut was justified because biomass and LAI were = strongly correlated (r 0.974). The resultant mean LAI of Chusquea tessellata leaves was 2.2 m2 m-2 (sd ? 0.8). Similarly, the bunch-grass LAI (0.16) had to be multiplied by 1.80, resulting in a mean bunch-grass LAI of 0.29 m2 m"2 (sd ? 0.28). The mean LAI of the Poaceae in the Chingaza dwarf bamboo p?ramo was 2.5 m2m-2 (sd ? 1.05).

Bunch?grass

' ' 88) percentage non-living material (Dec. 87-April Fig. 4. Monthly = tessellata ? sd) from Chusquea 16) and bunch grass (n (mean (n= 14). (*p>0.1).

Chusquea February.
non-living

Bunch-grass
material

tessellata material reached itsmaximum showed peak percentages

in January.

in of

Sixty percent (854 g DW- m-2)

of non-woody vascular plants was

of all aerial biomass

alive. The contribu

tion of bamboo in biomass was related to the domi nance of Chusquea tessellata in the vegetation cov er. Seventy three percent of the aerial biomass con sisted of Chusquea tessellata', bryophytes accounted for 15% and bunch-grasses for 7%. All other groups to the aerial biomass. Litter, contributed marginally consisting almost entirely of non-living Chusquea tes sellata material, contributed up to 34% of the above ground biomass (i.e., 900 g DW- m~2). The amount of bryophyte biomass correlated negatively with the amount of litter (Fig. 5). Vertical structure The p?ramo vegetation was dominated visually by tessellata bamboo and to a lesser extent Chusquea effusa (incl. C. bogotensis). Except by Calamagrostis for some dwarf shrubs, other plants did not exceed 10 cm in height (Fig. 8). The total dry mass per layer decreased with increasing height above ground. The surface was densely covered with litter and ground bryophytes. Litter was found in bamboo clumps to a

0.6

litter (kg.m
Fig. 5. Dry mass of bryophytes (kg DWm-2) as a function

)
(kg DW- m~2) per plot (n 4 = 16).

of litter dry mass

10

20

30

40
leaf

50
length

60

70
(mm)

80

90

100 110 120

I undamaged m

non-living

?Up

infected[7T*]damaged

6. Mean

proportions

of Chusquea

tessellata

leaves expressed

as % of total on 1m2 arranged by leaf length classes

(mm) (n

2,098).

Discussion

and conclusions

Verweij
rosettes elsewhere

1987; Verweij & Beekman,

o?Espeletia in the are area rare they at our are

in press).
study site,

Stem

although associated

Studies of drier bunch-grass p?ramos indicate that the main part of the aerial biomass consists of Espele tia stem-rosettes (Lutz & Vader 1987; Beekman &

commonly

with bamboos (Cleef 1981; Franco et al 1986). our plots the aerial biomass consisted principally

In of

35
O.OOIn

-4

0.0005

(y=

-7.93X 2

1.67X

+9.16)X

10 -4

(y=

-7.13X

1.54x

+6.25)

10 -4

(y= -12.8x + 9.23) x 10 -,-1

0.2 0.4 0.6 0.8 1.0

(y= -12.6X + 8.02)

x 10

"4

height

(m)

?O
?

Bunch-grass
Bunch-grass

living
non-living

Chusquea Chusquea

tessellata

living

_g

tessellata non-living

Fig.

7.

Cumulative

LAI

(In transformed)

(m2m

2) of non-living

bunch-grass

and Chusquea

tessellata

leaves as a function

of the height

(m).

2 ; 73%). This Chusquea tessellata (1,263 g DW- m is comparable to the biomass value of Aristizabal & On average, Casta?eda (1983) (1,419 g DW-m"2). the total above-ground biomass of the bamboo type under study is lower than biomass values (Table 1). The tropicalpine bunch-grasslands are assumed to values extrapolated to 1 ha p?ramo of other biomass be low

er, taking account of the space between the clumps. However, this is also true for the bunch-grass p?ramo studies in the Colombian Andes, referred to inTable 1. The bamboo plot studied can be classified as mesic and is therefore comparable with the bunch
grass p?ramo plots referred to before. In fact, oth

the amount of living of living tissues. Consequently on when the samples are taken. material depends We suspect that below-ground biomass in the bam boo p?ramo is considerably greater than in the bunch grass p?ramo, taking into account the impressive and dense rhizomes of the Chusquea tessellata stands (see the also Soderstrom & Calder?n 1978). However, tes above: below-ground biomass ratio of Chusquea sellata (1:0.4; G. J. Toi unpubl.) falls, in this low bam in the range (1:0.4 to 1:1) found for boo vegetation, of non-seasonal cold environments, including species high tropical mountains (Smith & Klinger 1985), and of ratios found for bunch-grass p?ramo (Lutz & Vader 1987 (1:0.5); Beekman & Verweij 1987 (1:0.3)). bunch-grass p?ramo (Lutz & Vader Comparing with this bamboo p?ramo, the following points 1987) are of particular note (Fig. 8). Both p?ramo vegetation types are dominated by Poaceae of different growth forms. The dominant plants in the bunch-grass p?ramo is Calamagrostis effusa and in the bamboo p?ramo Chusquea tessellata. Total above-ground biomass of this bamboo p?ramo is lower than that of bunch-grass p?ramo, but living aerial biomass is higher. Sixty per cent of the aerial biomass of non-woody vascular plants is living in the bamboo p?ramo. Lower percentages

er bamboo p?ramo vegetation types in the study area apparently contain much higher levels of above-ground
biomass. Under favourable conditions Chusquea tes

sellata grows
these sites,

luxuriantly and forms dense groves. At

are wet and peaty. Water and nutri

soils

ents are readily collected from the surrounding slopes. Increased biomass values of Chusquea tessellata are found where total bases are higher (Toi & Cleef 1992). Apart from spatial variation, temporal variability is present in the bamboo p?ramo plots. At the end of the
dry season senescence occurs, reducing the proportion

36

Table Source

1. Biomass

figures Country

(g DW-m

2) of tropical alpine grasslands. Latitude Altitude Vegetation type Aerial biomass (g-m-2) bunch-grass paramo Litter (a) (b) Above-ground biomass (a+b)

(Study area) Cardozo & Colombia (Eastern Cordillera) Lutz & Vader Colombia (Eastern Cordillera) Beekman & Colombia (Eastern Cordillera) Toi & Cleef (this study) Colombia (Eastern Cordillera) Hofstede & Colombia (Central Cordillera) Smith & Klinger Venezuela (M?rida Andes) (1992) Ecuador (Eastern Cord.) Hnatiuk(1978) New Guinea

(degrees) (m)
4?33' N 3,300-3,400

(g-m 2) (g-m 2)
735-2,660

Schnetter(1976)

74?02'W

4?35' 74?04

N W

3,620

bunch-grass p?ramo

2,476

615

3,091

(1987)

4?35'

3,620

bunch-grass p?ramo

3,052

334

3,386

Verweij (1987)

74?04' W

4?40'

3,650

bamboo p?ramo

1,725

900

2,625

73?48' W

4?48'N 75?24'W

4,100

bunch-grass p?ramo

1,392

1,472

2,864

Witte (1993)

8?47'N 70?48' W 1?48' S 78?32'W 5?47' S

3,530-3,560

bunch-grass p?ramo

398-651*

(1985)
Ramsay

3,750 3,950 3,400 3,510 4,350

bunch-grass p?ramo tussock grassland

837 794 2,668 2,460 3,431

(Mt. Wilhelm)

145?01'E

329 369 122

2,997 2,829 3,553

* heavily grazed

of

living

aerial

biomass

occur

in bunch-grass

p?ramo

sellata

sheds

senescent

leaves,

which

become

heav

(Lutz & Vader 1987 (24%); Beekman & Verweij 1987; Verweij & Beekman, in press (44%); Hofstede et al, in a lower value of total (living press (33%)). Likewise, and non-living) Poaceae LAI (3.7) is found in the bam boo p?ramo compared with bunch-grass p?ramo (Lutz & Vader 1987 (12.8); Beekman & Verweij 1987 (4.7)). Living Poaceae LAI of the bamboo p?ramo (2.5) is similar or higher than that of tropicalpine bunch grass lands (Hnatiuk 1978 (1.6-2.4); Lutz & Vader 1987 (2.4); Beekman & Verweij 1987 (1.4); Hofstede et al, in press (2.4)). The LAI value of the bamboo p?ramo falls in the range of unifacial, living LAI values for the central European Alps and the Venezuelan Andes, (Vareschi 1953). respectively, 0.4-1.3 and 0.05-1.2 A main difference between the growth forms of
bamboo and bunch-grass is that bunch-grass keeps

ily infected with the fungus Phyllachora this bamboo species must Apparently,
mechanisms and/or strategies against

chusqueae. have other

radiation and

frost. In this connection it is interesting to note the characteristic 45? (upward) leaf inclination and the
xeromorphic leaves of Chusquea tessellata.

A result of the leaf shedding is the higher amount of litter in the bamboo p?ramo (900 g DW- m~2 (sd? 211); Table 1). Bamboo leaves seem to have a slower
turnover rate than bunch-grass leaves. Establishment

of other plant species ismore difficult on a thick layer of Chusquea litter and is thus strongly controlled by Chusquea litter fall (in agreement with Grime 1979).
The bunch-grasses the are co-dominant and can apparent tessella ly withstand co-occurrence of Chusquea

most

leaves attached to the tussock, of its non-living providing protection against strong solar radiation and freezing temperatures (Hnatiuk 1978). Chusquea tes

ta, possibly by infiltration with stolons in the manner described by Coughenour (1985). Under extreme wet climatic or edaphic conditions Calamagrostis bunch grass species are completely absent (Cleef 1981).

37

I c3

C/> cd "o <D Q_

> O ? "U c 13 O O

sz c/> -o c (O c/> ?? o X

-5 LU

O) c

O) C ? *> c S3 13 o

I CD g ?

(0 (0 CD (D ? ? CO CO O O

*"

DI

.00

38 Acknowledgments We would like to express our gratitude to the Ar?valo family in Santa F? de Bogot? who provided a home base during the first author's stay in Colombia. We thank the Instituto Geogr?fico 'Agust?n Codazzi' for office and soil laboratory facilities. We are indebted to for logistical help. INDERENA Dr. M. H. Ramirez-A. fieldwork in the Chingaza National kindly permitted of Santa F? de Bogot? (EAAB) Park. The Waterworks in the Chingaza Nation offered field accommodation al Park. We remember the 'amistad' of the EAAB
employees. Robert G. M. Theo C. M. and Brock, two Yvonne anonymous Widmer, review Hofstede years. 42-66 pp. Dis 79, J. Cramer, Vaduz. to grazing by large responses Coughenour, M. B. 1985. Graminoid herbivores: and interacting processes. expatiations, adaptations, sertationes Botanicae Ann. Missouri Bot. Gard. 72: 852-863. in the P?ramo Dierikx, L. & Kaandorp, M. 1986. Vegetation mapping of Chingaza M. Science Thesis, (Cundinamarca, Colombia). 67 pp. of Amsterdam. Hugo de Vries Laboratory, University Soil map of the world. Vol. 1. Legend. FAO, 1974. FAO-UNESCO Unesco, Franco-R., Paris. O. & Lozano-C, G. 1986. Estudios P., Rangel-Ch., en la Cordillera Oriental. II - Comunidades ecol?gicos vegetales de la Laguna de Chingaza de los alrededores Cundinamarca. 15:219-248. Caldasia John processes. strategies and vegetation & Sons, Chichester. R. J. 1978. The growth of tussock grasses on an equatorial islands. In: Troll, C. & and on two sub-antarctic high mountain relations between the southern Lauer, W. (eds.), Geoecological J. P. 1977. Plant A continuous record of the last 3.5 million

Grime,

Wiley Hnatiuk,

ers are thanked for comments on an earlier draft of this paper. Paul M. Ramsay kindly corrected the English
text.

159-190 pp. Erdwiss. temperate zone and the tropical mountains. 11, Franz Steiner Verlag, Wiesbaden. Forschung H. J. L. 1993. An evaluation R. G. M. & Witte, of the Hofstede, estimation use of the dry-weight-rank and the comparitive yield biomass in p?ramo ecosystems research. Caldasia 17: methods 11-14. Hofstede, E. J. & Sandoval-S., E. M. (in R. G. M., Chilito-P, de las macollas de la microestructura paramu press). Aspectos In: Cleef, nas; implicaciones y impacto del pastoreo. ecol?gicas mountain M. (eds.), Neotropical A. M. & Monasterio, ecosystem and global change. Tercer in relation to biodiversity functioning Mundo Editores, Sant? F? de Bogot?. del Proyecto Chingaza. EAAB, San Ingetec, 1970. Mapa Geol?gico

References
de la fitomasa a?rea J. 1983. Estimaci?n Aristizabal, H. & Casta?eda, en la cuenca del r?o Chuza, P?ramo de Chingaza, Cundinamarca. de Biolog?a, Facultad de Ciencias, Tesis de Grado, Departamento Santa F? de Bogot?. 36 pp. de Colombia, Universidad Nacional Beekman, A. & Verweij, P. A. 1987. Structure and nutrient status of in relation to soil and climate. a p?ramo bunch-grass vegetation M. Bekker, Science Amsterdam. Thesis, 66 pp. Hugo de Vries Laboratory, University of

del p?ramo de 1989. La vegetaci?n An?lisis de Tausa, Cundinamarca). 1-193 pp. 14, IGAC, Santa F? de Bogot?. Geogr?ficos P. C, Young, R. & Cleef, A. M. Bosman, A. F., van der Molen, a p?ramo cushion mire. of Turbera de Cacostros, 1993. Ecology Science 4: 633-640. Journal of Vegetation en el ecol?gicos Cardozo, H. & Schnetter, M. L. 1975. Estudios Laguna R. P. & Cleef, A. M. Verde (Municipio La biomasa de tres aso Colombia.III. de Calamagrostis effusa y la productividad vegetales Ruhl. en compara columbiensis (H.B.K.) Steud. y Paepalanthus 11: 85-91. de clorofila. Caldasia ci?n con la concentraci?n section Swallenochloa, of Chusquea Clark, L. G. 1989. Systematics and section Longifoli section Serpentes, section Verticillatae, P?ramo de Cruz Verde, ciaciones ae (Poaceae.Bambusoideae). 27, Systematic Botany Monographs Ann Arbor. 1-127 pp. of Michigan. University G. & Ellis, R. P. 1989. Natural hybridization Clark, L. G., Davidse, section Swallenochloa Evidence in bamboos: from Chusquea Nat. Geogr. Res. 5: 459-476. (Poaceae: Bambusoideae). Cleef, A. M. 1978. Characteristics relations. and its subantarctic relations Geoecological and the tropical mountains. of neotropical p?ramo vegetation In: Troll, C. & Lauer, W. (eds.), the southern temperate zone between Forschung of the Colom 61, J. Cramer, 11,

ta F? de Bogot?. Keizer, J. J. 1989. A Colombian bunch-grass p?ramo along a fire induced age gradient. M. Science Thesis, Hugo de Vries Labora 55 pp. of Amsterdam. tory, University Koppen, W. & Geiger, R. Borntraeger, Berlin. Kloosterman 1930. Handbuch der Klimatologie. Teil I.

S. & Cleef, A. M., in press. Mapa de la H., Salamanca, (Cordillera vegetaci?n del Parque Nacional Natural Los Nevados In: T. van der Hammen & A. G. dos San Central, Colombia). Transecto tos (eds.), La Cordillera Central Colombiana, Parque Studies on Tropical Los Nevados (Tercera parte y conclusiones). 4. J. de Ecosistemas Andean Ecosystems/Estudios Tropandinos

Cramer (Borntraeger), Berlin-Stuttgart. and nutrient status Lutz, R. & Vader, P. 1987. Biomass, productivity of a Colombian p?ramo. M. Science Thesis, Hugo bunch-grass 61 pp. of Amsterdam. de Vries Laboratory, University to the New of bamboos native F. A. 1973. Genera McClure, World (Gramineae: Bambusoideae). to Botany 9: 1-148. Smithsonian Contributions

for the measurement R. E. 1968. Methods Milner, C. & Hughes, no. 6, I.B.P. Handbook of the primary production of grassland. Oxford and Edinburgh. Scientific Publications, Blackwell Ramsay, the com of Ecuador: 1992. The p?ramo vegetation P. M. of tropical grasslands munity ecology, dynamics and productivity of Wales, Bangor. 274 pp. in the Andes. Ph.D. Thesis, University Rossenaar, A. J.G. A. & Hofstede, R. G. M. 1992. Effects of burning In: Balslev, in the p?ramo ecosystem. and grazing on root biomass under H. & Luteyn, J. L. (eds.), P?ramo. An Andean ecosystem human influence. 211-213 Press, London. pp. Academic O. in press. The A. M. & Rangel-Ch., S., Cleef, Salamanca, Los Neva of the Parque Natural Nacional p?ramo vegetation In: T. van der Hammen & dos (cordillera Central, Colombia).

365-390 pp. Erdwiss. Franz Steiner Verlag, Wiesbaden. of the p?ramos 1981. The vegetation Cleef, A. M. Botanicae bian Cordillera Oriental. Dissertationes 321 pp. H. Cleef, A. M. & Hooghiemstra, area of the high plain of Bogota. Vaduz. tional and climatic 1984. Present

of the vegetation In: Hooghiemstra, H., Vegeta history of the high plain of Bogot?, Colombia:

39
A. G. Studies dos Central Colombiana, Santos (eds.), La Cordillera (Tercera parte y conclusiones). Parque Los Nevados on Tropical Andean Ecosystems/Estudios de Ecosistemas O. & Rivera-O., D. in press. Comunidades Vargas-R., vegetales del Parque Nacional Natural Chingaza: sector I r?o La Playa-r?o M. (eds.), Neotropical Guatiqu?a. In: Cleef, A. M. & Monasterio, mountain in relation to biodiversity and ecosystem functioning global change. Tercer Mundo Editores, Sant? F? de Bogot?. Van der Hammen, T., Perez-R, A. & Pinto-E., P. (eds.) 1983. Stud ies on tropical Andean Vol. 1. La Cordillera Cen ecosystems. tral Colombiana. Transecto Parque Los Nevados (Introducci?n y datos iniciales). J. Cramer, Vaduz. 345 pp. Van der Hammen, T. & Ruiz, P. M. (eds.) 1984. Studies on tropical Andean Vol. 2. La Sierra Nevada de Santa Marta ecosystems. J. Cramer, Vaduz. Cumbre. (Colombia). Transecto Buritaca-La Van S. & Alvarez, V J. (eds.) 1989. Vol. 3. La Cordillera ecosystems. Central Colombiana. Transecto Parque Los Nevados (Segunda parte). J. Cramer, Vaduz. 600 pp. Van der Hammen, T. & dos Santos, A. G. (eds.) in press. Studies on tropical Andean Vol. 4. La Cordillera Central ecosystems. Colombiana. Transecto (Tercera parte y Parque Los Nevados Verweij, J. Cramer, Berlin, Stuttgart. conclusiones). P. A. & Budde, P. E. 1992. Burning and grazing gradients in Initial ordination analyses. In: Balslev, H. & p?ramo vegetation: under human Luteyn, J. L. (eds.), P?ramo. An Andean ecosystem 603 pp. der Hammen, T., Diaz-R, Studies on tropical Andean

Transecto

4. J. Cramer (Borntraeger), Berlin-Stuttgart. Tropandinos Schmidt, A. M. & Verweij, P. A. 1992. Forage intake and secondary in extensive livestock systems in p?ramo. In: Balslev, production under H. & Luteyn, J. L. (eds.), P?ramo. An Andean ecosystem human influence. 197-210 pp Academic Press, London. Smith, A. P. & Young, T. P. 1987. Tropical alpine plant ecology. Ann. Rev. Ecol. Smith, Syst. 18: 137-158. J.M. B. & Klinger, L. F. 1985. Above-ground: below-ground and their sig ratio in Venezuelan phytomass p?ramo vegetation 17: 189-198. nificance. Arctic and Alpine Research T. R. & Calder?n, C. E. (Poaceae: Bambusoideae): new species. Brittonia 30: 297-312. and Swal 1978. Chusquea and Generic relationships

Soderstrom, lenochloa

E. J. & Clark, L. G. 1988. Distribu Soderstrom, T. R., Judziewicz, tion patterns of neotropical bamboos. Neotropical distributions: Rio de Janeiro, 12-16 January of a 1987 Workshop, Proceedings Brasileira de Ciencias. 1987, Academia der andinen P?ramoregion. Sturm, H. 1978. Zur Oekologie Biogeo 121 pp. 14, Junk Publishers, The Hague. graphica O. 1985. Ecolog?a de los p?ramos andinos. Sturm, H. & Rangel-Ch., Una visi?n preliminar integrada. Instituto de Ciencias Naturales. de Historia Natural, Biblioteca Jos? Jer?nimo Triana 9, Santa F? de Bogot?. 292 pp. status of a Chusquea 1992. Nutrient Toi, G. J. & Cleef, A. M. tessellata bamboo p?ramo. In: Balslev, H. & Luteyn, J. L. (eds.), under human influence. An Andean 123? ecosystem 135 pp. Academic Press, London. de sociedades Vareschi, V 1953. Sobre las superficies de asimilaci?n de cordilleras Bol. Soc. vegetales tropicales y extratropicales. Venez. Cie?e. Nat. 14: 121-173. P?ramo. Museo

177-195 pp. Academic influence. Press, London. Verweij, P. A. & Kok, K. 1992. Effects of fire and grazing on Espele tia hartwegiana In: Balslev, H. & Luteyn, J. L. populations. under human influence. (eds.), P?ramo. An Andean ecosystem 215-229 Press, London. pp. Academic