Plant communities of montane secondary vegetation in Cordillera de Talamanca - Kappelle et al. 1994

Phytocoenologia

Berlin-Stuttgart, December 5, 1994

Plant communities of montane secondary vegetation in the Cordillera de Talamanca, Costa Rica

by Maarten KAPPELLE, Henk P. VAN VELZEN and Welmoed H. WIJTZES, Amsterdam with 3 photos, 5 figures and 5 tables

Abstract. A phytosociological analysis of the secondary vegetation found in the montane vegetation belt of the Costa Rican Cordillera de Talamanca was carried out. Releves were made in randomly-stratified clustered sample plots distributed over homogeneous patches of secondary vegetation. A TWINSPAN classification, refined by Braun-Blanquet procedures, revealed twelve secondary plant communities (six grasslands, two shrublands and four forests), which were grouped into a lower montane (ca. 2300 m alt.) and an upper montane (ca. 2800 malt.) set of six communities eaGh. These communities have been described by characterizing their structure, composition, ecology (environmental aspects) and distribution. Finally some differences and similarities between the communities, as well as their position along a possible successional gradient, have been discussed.

Resumen. Se hizo un analisis fitosociologico de la vegetacion secundaria encontrada en el piso montano de la Cordillera de Talamanca en Costa Rica. Levantamientos fueron realizados en parcelas eligidas al azar, agrupadas, y distribuidas sobre vegetacion secundaria homogena. Una clasificacion con el programa TWINSPAN, refinada con procedimientos propuestos por Braun-Blanquet, revelo doce comunidades vegetales (seis tipos de pastizales, dos tipos de matorrales, y cuatro tipos de bosque). Estas resultaron ser agrupados en dos bloques de seis comunidades, un bloque representando la faja montano-baja (alrededor de 2300 m.s.n.m.), y el otro la faja montano-alta (alrededor de 2800 m.s.n.m.). Estas comunidades fueron descritas caracterizando su estructura, composicion, ecologia (aspectos ambientales), y distribucion. Finalmente, unas diferencias y similitudes entre las comunidades fueron discutidos, tanto como la posicion de ellas a lo largo de un posible gradiente sucesional.

Introduction Recently the structure and floristic composition of a variety of primary plant communities inhabiting diverse neotropical montane ecosystems have been studied in detail (among others CLEEF eta~. 1984, KAPPELLE et al. 1989, VAN DER HAMMEN et al. 1989, HEANEY & PROCTOR 1990, VAN VELZEN 1992, WOLF 1993a and 1993b). However, secondary plant communities that establish in neotropical upland regions following human intervention (including clearing, burning and grazing) have received less attention (CUATRECASAS 1934, SUGDEN 1983, CLEEF et al. 1984, MONASTERIO et al. 1987, STUTZ DE ORTEGA 1990). Today such communities are widely distributed, every day 0340-269X/94/0022-0449 $ 9.25 Š 1994 Gebriider Borntraeger, D-14129 Berlin ¡ D-70176 Stuttgart

450

M. Kappelle et al.

occupying larger areas (SALAMANCA 1991, VAN VELZEN & WIJTZES 1990, VERWEIJ & BEUKEMA 1992). They are believed to remain impoverished for a long period of time before a retarded recovery process makes them return to a stage comparable to the original montane mature forest (KAPPELLE 1993 and in prep.).

In order to get a first insight in the structure and floristic composition of these man-induced communities a phytosociological study was carried out from 1989 through 1990 in the severely degraded montane forest belt of the Costa Rican Cordillera de Talamanca. Descriptions of the original natural plant communities (Chusquea-Quercus forest types) available for this mountain region served as a measure of comparison (KAPPELLE 1991; KAPPELLE et al. 1989 and in prep.). /

Study area

The study area is located in the Los Santos Forest Reserve (Fig. 1) in the western part of the Cordillera de Talamanca, which is made up of intrusive and volcanic Tertiary rocks, alternated with marine sediments (WEYL 1980, CASTILLO 1984, KAPPELLE et al. 1989, VAN UFFELEN 1991). Pleistocene glaciations have left their traces, such as fossil periglacial phenomena on the Cerro de la Muerte (3491 malt.) at the nearby Buenavista massif (HASTENRATH 1973).

Soils developed in volcanic ashes sustain montane mature Quercus forests and secondary vegetation at cleared sites, being medium-textured, moderately fertile and very acid. They have been excessively drained (VASQUEZ 1983,

Pacific Ocean 0

9 100 km

I 8o

o

86

0

85

Amistad Biosphere Reserve

0

84

Fig. 1. Location of the study area showing the 62000 ha Los Santos Forest Reserve and the bordering Amistad Biosphere Reserve in Los Santos Forest Reserve eastern-central Costa Rica.

Temperature ( C)o

Precipitation (mm) 11 .2

{2648) 400 300 200

50

100

40

80

30

60

20

40

10

20

0

0

-10 -20

Fig. 2. Walter climate diagram from O jo de Agua in the Costa Rican Cordillera de Talamanca at 5 km NW of the sample sites.

m a

m

Ojo de Agua (2960 m)

a

5

0

n d

(1963-1985)

Photo 1. Deforestatio n along the Panamerican Highway near Ojo de Agua, showing patches of upper montane secondary grasslands, shrublands and forests in the foreground, bordering large tracts of different Quercus forest types in the background .

452

M. Kappelle et a!.

VAN UFFELEN 1991). The area has a humid to perhumid, temperate to cold climate with a short dry season Qanuary through April). Fig. 2 shows a climate diagram after WALTER (1985) for the nearby location of Ojo de Agua (2960 m alt.). Here the average annual temperature is 11.2 째C, while average rainfall is 2648 mm per year (Instituto Meteorologico Nacional1988, HERRERA 1986). About 160m lower, at ca. 2800 m a.s.l., the mean annual temperature reaches 12 째C, while at about 2300 m alt. in the lower montane belt the mean annual temperature fluctuates around 14.5 oc (HERRERA 1986 ).

Diurnal fog during the afternoons is considerable (cloud forest), especially during the rainy season (KAPPELLE 1992).

Sample sites are situated in the 62 000 ha Los Santos Forest Reserve, which has been highly deforested during the last decades (9째35'40" N, 83째44'30" W).

Since 1982 this protected area, created in 1975 (MEZA & BONILLA 1990), has served as a buffer zone to the 612 570 ha UNESCO-declared Amistad Biosphere Reserve (RESERVA DE LA BIOSFERA LA AMISTAD 1990; Fig. 1). Continuous clearing of upper montane oak forest stands, initiated in the 1940's with the construction of the Panamerican Highway, followed by burning and grazing has generated a highly heterogeneous pattern of vegetation types (Photo 1). Today tracts of pristine lower and upper montane ChusqueaQuercus forests present within the borders of the Los Santos Forest Reserve between elevations of 2200 and 3000 m alt. alternate with patches of secondary vegetation of different ages, including grasslands, shrublands, and open and closed forests forming a mosaic (KAPPELLE 1993, KAPELLE et al. 1989, VAN VELZEN & WIJTZES 1990). The best example of such a vegetational mosaic is found in the upper part of the Rfo Savegre watershed in the vicinity of San Gerardo (lower montane, 2200-2400 malt.) and Jaboncillos (upper montane, 2700-2900 malt.) in the Copey District of the Dota Canton in the Province of San Jose.

Methods Site selection took place on basis of a preliminary interpretation of aerial photographs on a scale of 1 : 60 000 dating from 1969 (SPREUWENBERG 1989) and a rapid reconnaissance survey of the study area in early 1989 (VAN VELZEN & WIJTZES 1990), as recommended by ITC standard procedures (VAN GILS & VAN WIJNGAARDEN 1984 ). During the rainy season of 1989 phytosociological releves were conducted in 164 randomly-stratified, mainly clustered sample plots, following the Braun-Blanquet approach (BRAUN-BLANQUET 1965, ZoNNEVELD 1979). These plots were established in secondary vegetation stands occurring in both the lower montane (78 releves) and the upper montane (86 releves) rain forest belt. Information on the former landuse history of different selected plots was collected from informal conversations held with several nearby-living farmers. Plot sizes were fixed on 4 to 25 m 2 for speciespoor grasslands, 100 m 2 for shrublands, and 150 to 500 m 2 for species-rich forests, as CLEEF et al. (1984) successfully applied in similar vegetation types occurring in the Colombian Andes.

Montane secondary vegetation in the Cordillera de Talamanca

453

In these plots all terrestrial vascular plant species were sampled and their growth form recorded. Their quantitative occurrence was assessed by estimating the relative aerial crown or shoot cover projection using the well-known Braun-Blanquet scale (WESTHOFF & VANDER MAAREL 1973, MUELLER-DOMBOIS & ELLENBERG 1974, BRAUN-BLANQUET 1965, CLEEF et al. 1984, KAPPELLE et al. 1989). A total of 1950 specimens was collected, representing 388 vascular plant taxa (81 ferns and fern-allies, 1 conifer, 49 monocots and 257 dicots). These specimens were identified by different specialists and when fertile stored at the Costa Rican National Herbarium (CR) and at the herbarium of the Dutch University of Utrecht (U). A complete checklist of the vascular plant taxa identified at species level, including authorities, has been presented

by VAN VELZEN et al. (in press). Taxonomic nomenclature follows basically TRYON & TRYON (1982) and LELLINGER (1989) for pteridophyte taxa, and STANDLEY (1937-1938), WOODSON & SCHERY (1943-1980), BURGER (19711990) and KAPPELLE et al. (1991) for gymnosperm and angiosperm taxa. Terrestrial bryophytes and lichens were also collected for a presence-absence analysis, but they have not been taken into account in the present vegetation classification, since identification down to the species level has not been finished yet. In addition to the floristic data, the average height and cover of the tree, shrub, herb and bryophyte layers were measured, while the dominance of certain growth forms (short grasses and graminoids, forbs, bamboos, shrubs and trees) was recorded for a basic analysis of the vegetation physiognomy and structure (BEARD 1944, 1955; ErTEN 1968; UNESCO 1973; VAN GILS & VAN WIJNGAARDEN 1984 ). Additional environmental information on topography (altitude, aspect), physiography (terrain form, slope), soil (depth, layering, color, pH) and human impact (clearing, burning, charcoal production and cattle grazing) was gathered on-site.

As a first step in the field data analysis a rough vegetation table including both montane rain forest belts was constructed following classic BraunBlanquet phytosociologica! procedures (WESTHOFF & VAN DER MAAREL 1973, MUELLER-DOMBOIS & ELLENBERG 1974, BRAUN-BLANQUET 1965).

This table was subsequently treated with the assistence of the multivariate cluster analysis programme TWINSPAN (HILL 1979; ]ONGMAN, TER BRAAK & VAN TONGEREN 1987) in order to reach a first approximation of the possible secondary plant communities in the area. For this purpose species cover-abundances were converted from percentages as estimated in the field into nine cover classes (Table 1), using an adapted form of the logarith-

Table 1. Conversion table defining cover classes for multivariate data analysis, as modified after GAUCH (1982). aerial cover valu e(%) estimated cover class

0

<1

2

2-3

4-7

8-15

16-31

32-63

64-100

3

4

5

6

7

8

454

M. Kappelle et a!.

mic octave-scaling technique proposed by GAUCH (1982) and successfully applied by VAN VELZEN & WIJTZES 1990). This division into cover classes corresponded with eight TWINSPAN cut levels (0, 1, 2, 4, 8, 16, 32, 64).

Thus a first computer analysis was conducted, showing a clear dichotomy which reflected a main separation of the two altitudinally different data bases. Therefore a second TWINSPAN classification was carried out for the lower and upper montane rain forest belt separately. On basis of these computer outcomes two series of 30 representative, homogeneous releves were selected and once again treated with the TWINSPAN programme. The final computer tables have been refined by hand, applying current phytosociological techniques (MUELLER-DOMBOIS & ELLENBERG 1974) as recently again recommended by BARKMAN (1990).

Classification of the vegetation was followed by description of the plant communities on basis of their characteristic species and ecological species groups. Since little is known about the vegetation types occurring in the Costa Rican montane highlands the present communities are still in a provisional form and official syntaxonomical names in accordance with the Code of Phytosociological Nomenclature by BARKMAN et al. (1986) have thus been avoided.

Differences among the communities in structure - on basis of diagrams after CLEEF et al. (1984) -, vascular plant diversity and growth form spectrum have been analyzed and discussed. Additionally, a schematic vegetation profile diagram has been drawn for both montane belts, showing in an illustrative way the possible successional sere from actively-used grasslands and recently-abandoned shrublands to recovering secondary forests.

Results Classification of montane secondary plant communities In the present vegetation analysis twelve secondary plant communities have been identified for the Costa Rican Cordillera de Talamanca. They are equally distributed over the lower (ca. 2300 m alt.) and upper (ca. 2800 m alt.) montane vegetation belt. Fig. 3 shows, by means of two TWINSPAN dendrograms, how these communities are related to each other. These hierarchical vegetation classifications show a clear decrease in recent human impact for both vegetation belts. The floristic composition of the communities as well as their characteristic and dominating species are presented in two vegetation tables, representing the lower and upper part of the montane belt respectively (Table 2 and 3). These tables show, that both the lower and upper montane communities consist of 13 ecological species groups.

The whole gamut of secondary plant communities studied for the Costa Rican montane rain forest belt can be described as a Monochaetum neglectum - Rubus eriocarpus vegetation complex. The lower and upper montane communities belonging to this type are grouped as follows:

c:r (") '""I 0

"I'j

.......

'1 OJ

OJ

~

:0 p,

z

(1)

z

no...

"'

8

(1)

:0 0...

(/J

1.3. Thelypteris rudis - Carex jamesonii 1-'

OJ

:0 p,

'1

c I

~3

., .,"

...,

(")

,0

(')~

CD to

.....

(i..o

"

~~

(/J

-"' (1)

(/J

::r 0'

-·(JQ

0

1.2. Geranium guatemalense- Oenothera epilobifolia

1-'

~. :0 V'J s-~ 0

0

(/J (/J

05~

;::;:

1.1 . Orthrosanthus chimboracensis - Cheilanthes notholaenoides - - · - - - - - ,

<0

~ 3 ~ ~3,.,

-·

;J "

[

[

t" 0

8

.,3

tTl

2.2. Wercklea lutea - Cornus disci flora

....

1---.o.--w--tv-->->- -

~

OJ

"

(1)

(JQ

c.

(/J

~

(I)

OJ 1-'

a g_ :0

::1

p,

::r (/J , c

1. 3. Ageratina subcordata - Muehlenbeckia tam'li folia

~1'..

(I)

.,

(1)

0

~

c::

N

[

1.4. Pteridium aquilinum- Polypodium macrolepis

0' I

...'"'

"'"'tTl ;u 8 z o-3

.,

c:r" <1>"-<: :0

L

.,:0

-·o

"'

:0

"

(")

1.2. Halenia rhyacophylla - Lolium perenne

OJ

(I)

"' (I)

0

1-'

e:...o0

(1)

w

(/J (/J

~ 3 c

___

1.1. Bromus sp. - Gnaphalium americanum

,

<0

- · "0

.,"'

....

::1

0...

~ ~-

(1)

.,<"'. ..

0

?' c t"rJ"O

(JQ

~

2.1 . Freziera candicans - Quercus seemannii

o...o >--l"' .,.,-.,:0" (1)

::;: tTl ;u

1.4. Galium mexicanum- Vaccinium consanguineum

I[

2.1. Fuchsia arborescens - Abatia parviflora 2.2. Quercus costaricensis - Quercus copeyensis

~

tTl

Table 2. Lower montane secondary pla nt commun ities, Costa Rica. plant community

1.1.

1.2.

1.3.

1. 4.

2 .1.

plot number (01 to 30)

00000 00001 11111 11112 22222 22223 12345 67890 12345 67890 12345 67890

(J1 """" a--

2. 2. ! 1.1.

Orthrosanthus chimboracensis-

grassland

75766 43334 54555 45322 33344 44335 00000 55500 00505 00500 55505 00550

! ! ! 1.2 . ! ! ! ! 1. 3. ! ! ! 1. 4. !

topsoil pH (15 em below level)

65655 65655 55665 65556 43334 43343

! 2 . 1.

Freziera candicans Quercus seemannii forest

total number of vascular species

11111 12112 12211 12112 33334 46454 38045 51383 82948 50450 29819 77755

! 2.2.

Wercklea lutea -

altitude (x 10m alt. )

aspect (exposition) slope angle (degrees)

22222 22222 22222 22222 22222 22222 33333 33333 33333 33333 33333 32333 97829 86665 14122 63484 14775 33334 EENSS NNNNN NNSSW ENEEN SNNNN NSWWW w EEE EWW E E EEEEE W

Cheilanthes notolaenoides Geranium guatemalense -

Oenothera epilobifolia grassland

Thelypteris rudis Carex jamesonii grassland Galium mexicanum -Va ccinium consanguineum shrubland

Cornus disciflora forest TWINSPAN division plant species (001 to 134) arranged in sociological groups (I to XIII)

II

III

00000 00000 00000 00111

00000 00000 11111 00000

00000 00000 11111 11111 00000 11111 00000 11111 11111 00011 00001 00111 11111

- - --- ----- ----- ----- - -------- ----- ----- --------- - -------2--- ----- ----1 --- - - - - - --

TWINS PAN division

growth form

plant family

1100 1100 1100 1100

herb fern shrub herb

Iridaceae Adiantaceae Melastomataceae Valerianaceae

herb herb fern herb shrub herb herb herb fern herb fern herb

Onagraceae Geraniaceae Hymenophyllaceae Phytolaccaceae Melastornataceae

shrub herb climber herb

Rosaceae Asteraceae Rubiaceae Scrophulariaceae

001 002 003 004

Pityrogramma chrysoconia Miconia sp . 1 Valeriana prionophylla

677-2 -3--2 ---32 ---3-

005 006 007 008 009 010 011 012 013 014 015 016

Oenothera epilobifolia Geranium guatemalense Hymenophyllwn sp.

Phytolacca rugosa Leandra aff.melanodesma Lachemilla spp.

Hydrocotyle sp.

Plantago australis Thelypteris rudis Carex spp .

Eriosorus flexuosus Viola scandens

-2--- -3545 --------- 242 - 3 ----1-21- --------- 43--- --------1 -4461 2-243 ----2 34 - 3- -33------ 45255 233------ --344 422 - - - - -- ---23 33-33 ---- - -- - - - -3465 ----- -23------ -3--4

-- - -- ----- -1-1-

1101 1101 1101 1101 1101 1101 1101 1101 1101 1101 1101 1101

017 018 019 020

Rubus spp .

-5- 4- 753-2 35225 44345 23223 -321----- 21--- 123-- ---1- ----1---43233 ----- - - -- -3--- --1-- ----- ---44 --- -- - ----

1111 1111 1111 1111

Orthrosanthus chimboracensis

Hieracium irazuense

Galium mexicanum Castilleja talamancensis

------ --- ~-------- ---- --------- -----3-- - -----

------- --------------

-- -- - -- - -- ------------- - --- ----- -2-1-

-----

-- ---

Rosaceae

Apiaceae Plantaginaceae Thelypteridaceae Cyperaceae Adiantaceae

Violaceae

~ ~

.t) "0

~ (b'

~

~

IV

v

VI

VII

Pennisetum clandestinum

X

XI

herb herb herb shrub shrub tree herb fern herb herb herb shrub

56457 45334 23-56 55467 -13232-22 5555- -2--- 37257 ---12 -4--2---- 4----3---1---

1010 1010 1010 1010

shrub fern climber herb

55677 -3356 33-22 Ageratina subcordata 24254 Archibaccharis aff. panamensis -4--5 Buddleja ni tida 33--Gnaphalium attenuatum 32-33 Cheilantes notholaenoides 2-353 Gnaphalium americanum -2--Hypochaeris radicata ----Sal vi a i odochroa ---1Solanum rudepannum -----

67888 45444 22223 67433 23--3 43--6 ---22

033 034 035 036

Monochaetum neglectum Blechnum occidentale Muehlenbeckia tamnifolia Oxalis spiralis

Holcus lanatus

Conyza bonariensis

465-7 3-35---------

----2 --------5----

88888 45443 55333 34556 36436 -62-4 --------121-3-32---------

87757 54523 232-2 4452--23-

- - -----31 -1--11123 -11-1 3--13 -32-- --------3 ----1 2-2-- ------3-----1 -------2-

---------11--2-1-3----1------------------

Poaceae Poaceae

Asteraceae Asteraceae Asteraceae Loganiaceae

Asteraceae Adiantaceae Asteraceae

Asteraceae Lamiaceae

Solanaceae

31211 -3211---1 1------1-

1001 1001 1001 1001 1001

fern climber tree fern fern

042 043 044 045

Monnina crepinii Pteridium aguilinum

-13--2------1----

31223 6-464 55574 11111

-3-33 -1--2--------

0111 0111 0111 0111

tree fern tree herb

----- -1--- ----- ----2 ---1- -1--1 --------4 -2--1

0110 0110

tree herb

048 Smilax kunthii 049 Quercus seemannii

----- ----- 2-1-- -2-32 11226 -3--2 ----- ----- ----- 46-4- 45756 72---

0100 0100

climber tree

050 Tradescantia commelinoides 051 Senecio multivenius 052 Polypodium skinerii

----- ---22 ----3 -----1613 ----- ----2 ----- -1--- -1-1----- ---2- ----- ----- ----- 3----

0011 0011 0011

herb tree fern

----- ----- ----- -3-8- 27344 --------- ----- ----- -2--- 537-3 1-------- ----- -1--- 44335 11-1----55333 ----- ----- ----- ----- 12111 --- ------ ----- ----- ----- 32-13 -3------- ----- ----- ----- 24-4- 4---224-4 --------- ----- ----- 3-2-3 --------- ----- ----- ----- 132-1 ------------- -1111 --1------ ----- ----- ----- -12-3 --------- -----21-2 --------- ----- -- --- ----- -33-- --------- ----- ----- -11-- ~-------- ----- ----- ----- -1-2- --------- ----- ----- ----- --1-1 -----

0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000 0000

tree Ericaceae fern-ally Lycopodiaceae tree Theaceae Theaceae tree herb Rubiaceae Melastomataceae shrub Fagaceae tree Caprifoliaceae tree tree Aqui foliaceae tree Myrsinaceae Lomariopsidaceae fern fern-ally Lycopodiaceae tree Lauraceae shrub Melastomataceae tree Rutaceae Styracaceae tree herb Bromeliaceae

Vaccinium consanguineum

Gnaphalium elegans

053 054 055 056 057 058 059 060 061 062 063 064 065 066 067 068 069

Comarostaphylis arbutoides Lycopodium thuyoides Cleyera theaeoides Freziera candicans Relbunium hypocarpium Leandra subseriata Quercus copeyensis Viburnum costaricanum Ilex valerii Ardisia costaricensis Elaphoglossum aff. engelii Lycopodium clavatum/contiguum Phoebe cinnamomifolia Miconia glaberrima Zanthoxylum scheryi Styrax argenteus Vriesea williamsii

----- ----7 ----- -----

0

::>

.,...,p...

(1Q

11--1 ----1 -------1---1-

-2-54 525-5 56465 11---

"'0"" '<

2-443 ---2- -22-3 ----2 ----- ----- ----5 1-2-3 ----- ----- ----4 --------2 --3-- ----- ------44442 ----------

.,::>g

Blechnaceae Polygonaceae Oxalidaceae

Adiantum andicola Pseudobaccharis trinervis Weinmannia pinnata Elaphoglossum conspersum Elaphoglossum engelii

----2 ---------1--1

s:::

0

Melastomataceae

037 038 039 040 041

VIII 046 Myrsine coriacea 047 Bidens pilosa IX

-----

1110 1110 1110 1110 1110 1110 1110 1110 1110 1110 1110 1110

021 022 023 024 025 026 027 028 029 030 0 31 032

Adiantaceae

Asteraceae Cunoniaceae

<

"

"~ M

a¡ ::> s¡

g..

Lomariopsidaceae Lomariopsidaceae

"()

Polygalaceae

0..., p...

Dennstaedtiaceae Ericaceae Asteraceae

Myrsinaceae

Asteraceae Smilacaceae Fagaceae

.,...,~ p...

";j

;;;-

.,a::> ., ()

Commelinaceae Asteraceae Polypodiaceae

-!>\Jl '-1

....

Table 2. (cont.)

V1

00

1. 2.

1. 3.

1. 4.

2 .1.

2. 2.

plant community

1.1.

plot number (01 to 30)

00000 00001 11111 11112 22222 22223 12345 67890 12345 67890 12345 67890

XII

070 071 072 073 074 075 076 077 078 079 080 081

Chusquea tomentosa Oreopanax xalapensis Clusia palmana Bomarea acutifolia Myrsine pellucidopunctata Ilex pallida Meliosma glabrata Macleania rupestris Ilex discolor Oreopanax capitatus Polypodium macrolepis Phlebodium aureum

XIII 082 Polypodium plebejum 083 084 085 086 087 088 089 090 091 092 093 094 095 096 097 098 099 100 101 102 103 104 105 106 107 108

Fuchsia arborescens Fuchsia microphylla Peperomia tetraphylla Polystichum fournieri Passiflora sexflora Bocconia frutescens Cornus disciflora Polypodium plesiosorum Saurauia veraguasensis Tournefortia johnstonii Wercklea lutea Begonia udisilvestris Cestrum racemosum Peperomia palmana Anthurium concinnatum Cyrtomium macrosorum Liabum tonduzii Smilax subpubescens Palicourea brenesii Phyllanthus niruri Hansteinia ventricosa Oco tea laet evirens Jun gia ferruginea Billia hippocastanum Persea rigens Piper 1 acunosum

--- - - - ---- --1-- ---4- 77755 ----- ----- --2-- ----- 33215 ---- - ---- - ---- - -- --- 1-311 ----- ----- ----- ----- 111-1 ----- ----- ----- ----- 223-1 ----- ----- ----- -- --- -2------ -- --- ----3 ----- --- -- ---- - ----- ----2 ----- 2------------- - -- -- ---- - --- -- - ---- -1------ - ----- ----- ----- ----2 ---- - ---- - - ---- -- --- -- --1

8 - 657 23454 3311-1211 -3--2 23-- --1-2 1--1-4--3---1----1---

000 1 0001 0001 0001 0001 0001 0001 0001 0001 0001 0001 0001

bamboo tree tree climber tree tree tree shrub tree tree fern fern

Poaceae.

Araliaceae Clusiaceae Alstroemeriaceae Myrsinaceae Aqui foliaceae Sabiaceae

--- -- ----- ----- -----

- 1116 - 1--4 --1-2 ------ ---12-2 1-123 1131--------3 ----2 -----

------ -----------

---- -

-- --- - --- - - ------- - --- -- --- ------ --- - - --------- ----- ---- ----- --- -- - -- ---- -- --------- -- --- ------- -- -- --- -- ---

-- --- ------ --- -- ---

0010 0010 0010 0010 0010 0010 0010 0010 0010 0010 0010 0010 0010 0010 0010 0010 0010 0010 0010 0010 0010 00 10 00 10 0010 0010 0010 00 10

fern tree shrub herb fern climber tree tree fern tree tree

--- -- -- ------ - --- -- -- -- ----- --------- ----- -----

33111 12434 21-11-1144122 12332 22311 55443 43264-245 34-45 33777 11212 -123 11-11 1-111 1 - 111 17--4 2-3112--1 -1432 - 1121 - 11-5 -2-12 -1 -14 -211- 1 - 12

Polypodiaceae Onagraceae Onagraceae Piperaceae Dryopteridaceae Passif loraceae Papaveraceae Cornaceae Polypodiaceae Saurauiaceae Boraginaceae Malvaceae Begon iaceae Solanaceae Piperaceae Araceae Dryo pteridaceae Asteraceae Smilacaceae Rubiaceae Euphorbi aceae Acanthaceae Lauraceae Asteraceae Hippocastanaceae Lauraceae Pipe raceae

---- ------- --- --------- ------

- --1----1 --- -- - -------

---------- --- ----2 -- --- - ------ -- --- --

311- -32---2 ---2---2-2 - --1---- ---- -- ----1----- -

-- --1 ----2 -- -------- ----- -------- -- --------- ---

---- - -- --- -- ------ - ----- - ---- ----- - -- --

---- - -- --- -- --- -----

-----

----- --------- - - -- -- --- --- --

-----

-- -- -

tree herb shrub herb herb fern c limber climber shrub tree herb tree c limber tree tree shrub

Ericaceae Aquifoliaceae Araliaceae Polypodiaceae Polypodiaceae

~

.,~ '0 '0

";:;" M

~

109 110 111 112 113 114 115 116 117 118 119 120 121 122 123 124 125 126 127 128 129 130 131 132 133 134

Geonoma hoffmanniana Leandra costaricensis

Campyloneurum angustifolium Elaphoglossum erinaceum Hoftmannia josetina Centropogon costaricae Passiflora membranacea

Asplenium auritum Miconia sp. 2

Ardisia palmana Didymaea alsinoides

Ardisia glandulosomarginata TUrpinia occidentalis Arenaria lanuginosa Campyloneurum aff. pittieri Pteris paucinervata Chamaedorea sp.

Botrychium cicutarium Urtica leptophylla

----- -- - --- - - - -- ------ - ----- - -- --- -- - - -- --- --- -- - ---- - -------- - - ----- --- ------- - -- -- - ---------

--- --

----- - ----

-------------

- ----

--------- - -- -- ----- ---------

Valeriana scandens Asplenium moritzianum

Cissus microcarpa Asplenium harpeodes Hedyosmum mexicanum

-- --- ----- -----

Phenax rugosus

-----

Prestoea allenii

- - - - - -1 - 11 --552 - - - - - - - 111 31--12--11--11--11---- - - - 4--13--1 --- - - 1--1-33--32------ -1-2-1-1-1--3 - -- -- -1--1 ----- -1--1 --23--21--12----- --11--11----- --4 -2 --3-4 --3-4

0010 0010 0010 0010 0010 0010 0010 0010 0010 0010 0010 0010 0010 0010 0010 0010 0010 0010 0010 0010 0010 0010 0010 0010 0010 0010

palm shrub fern fern shrub herb climber fern shrub tree herb tree tree herb fern fern palm fern herb herb fern climber fern tree shrub palm

Arecaceae Melastomataceae Polypodiaceae Lomariopsidaceae Rubiaceae Campanulaceae Passifloraceae Aspleniaceae Melastomataceae Myrsinaceae Rubiaceae Myrsinaceae Staphyleaceae Caryophyllaceae Polypodiaceae Pteridaceae Arecaceae Ophioglossaceae Urticaceae Valerianaceae Aspleniaceae Vitaceae Aspleniaceae Chloranthaceae Urticaceae Arecaceae

;;: 0

::0

c::0 t1)

"' t1) (")

0

::0

,.,...,p_.

'<

< t1)

(JQ (i)

g

c;· ::0

s·

5(i)

()

...,0

s

if ., p_. (i)

~ ;:;;-

plant species found in only one plot

Aegiphila odontophylla

plot cover growth plant nr. class form family

28 27 27 30 Blechnum polypodioides 21 Campyloneurum amphostenon 02 Cardamine tlaccida 07 Cirsium subcoriaceum 02 Cystopteris tragilis 28 Clusia minor 28 Drymis granadensis 26 Elaphoglossum hottmannii 24 Elaphoglossum squamipes 25 Erigeron karwinskianus 04 Eugenia austinsmithii 25 Garrya lauritolia 27

Alfaroa costaricensis Ardisia compressa Ardisia revoluta

4 4 2 4 6 3 6 4 1 1 1 1 1 3 3 1

tree tree tree tree fern fern herb herb fern tree tree fern fern herb tree tree

! ! !

Verbenaceae !

Juglandaceae !

Myrsinaceae !

Myrsinaceae !

Blechnaceae Polypodi aceae!

Brassicaceae ! !

Asteraceae Woodsiaceae !

Clusiaceae !

Winteraceae !

Lomariopsid . !

Lomariopsid. ! !

Asteraceae Myrtaceae !

Garryaceae !

plant species found in only one plot

plot cover nr. class

growth form

plant family

Hemichaena fruticosa

01

shrub shrub tree herb herb tree climber shrub tree shrub tree shrub tree tree ·tree

Scrophular.

Rosaceae Magnoliaceae Orchidaceae Orchidaceae Myrtaceae Asteraceae Rubiaceae Myrsinaceae Asteraceae Rubiaceae Asteraceae Styracaceae Asteraceae Hydrophylla.

.,s::0 (")

Hesperomeles heterophylla 05 Magnolia sororum 25 Malaxis hastilabia

Maxillaria biolleyi Myrcianthes tragrans Oligactis volubilis Palicourea angustitolia Parathesis glabra Roldana heterogama

Rondeletia buddleoides Senecio copeyensis

Styrax glabrescens Verbesina oerstediana Wigandia urens

23 22 30 27 27 23 26 27 25 23 26 02

7 4 2 1 1 6 2 1 3 1 4 1 1 1 5

'"'

....

U1

'-D

M. Kappelle et al.

460

Lower montane secondary plant communities 1. Pennisetum clandestinum-Holcus lanatus grassland 1.1. Orthrosanthus chimboracensis-Cheilanthes notholaenoides grassland 1.2. Geranium guatemalense-Oenothera epilobifolia grassland 1.3. Thelypteris rudis-Carex jamesonii grassland 1.4. Galium mexicanum- Vaccinium consanguineum shrubland 2. Chusquea tomentosa-Oreopanax xalapensis forest 2.1. Freziera candicans-Quercus seemannii forest 2.2. Wercklea lutea-Cornus disciflora forest Upper montane secondary plant communitiess 1. Plantago australis-Rumex acetosella grassland 1.1. Bromus sp.-Gnaphalium americanum grassland 1.2. H alenia rhyacophylla- Lolium perenne grassland 1.3. Ageratina subcordata-Muehlenbeckia tamnifolia grassland 1.4. Pteridium aquilinum-Polypodium macrolepis shrubland 2. Chusquea tomentosa- Weinmannia pinnata forest 2.1. Fuchsia arborescens-Abatia parviflora forest 2.2. Quercus costaricensis-Quercus copeyensis forest Description of lower montane secondary plant communities (Table 2) 1.

The Pennisetum clandestinum-Holcus lanatus grassland

The different grassland communities belonging to this type are all dominated by Pennisetum clandestinum, an introduced high-quality grass, locally known as 'kikuyo'. In general, this grass species is present with cover values over 30 %, and accompanied by the grass Holcus lanatus, the forb Conyza bonariensis, the fern Blechnum occidentale, and the shrubs Ageratina subcordata, Monochaetum neglectum and Rubus eriocarpus, among others.

These grasslands are about 0.3 m tall, sometimes up to 1 m at bushy sites.

In a few cases isolated Buddleja nitida trees are observed, reaching heights of 5 m. These individuals are secondary forest remnants left over after weeding activities ('chapeo') practiced by the local farmers in the rainy season.

In the present study four different Pennisetum clandestinum - Holcus lanatus grasslands are distinguished, which are described below.

1.1.

The Orthrosanthus chimboracensis-Cheilanthes notholaenoides grassland

Structure and composition This grassland community is about 0.5 m tall, sometimes up to 1.35 m. It is made up of large patches of grasses and herbs covering a 75 % of each releve, intermingled with small patches of low shrubs (15 % cover). On the ground a terrestrial moss layer is present, covering about 15 % of the surface.

Montane secondary vegetation in the Cordillera de Talamanca

461

The iridaceous Orthrosanthus chimboracensis and the less abundant fern Cheilanthes notholaenoides are diagnostic for this grassland community.

They are accompanied by the codominating grass Pennisetum clandestinum, and species such as the shrubs Ageratina subcordata and M onochaetum neglectum, the forbs Conyza bonariensis and Gnaphalium attenuatum, the grass species Holcus lanatus, and the ferns Adiantum andicola, Blechnum occidentale and Pityrogramma chrysoconia.

Ecology and distribution The Orthrosanthus chimboracensis-Cheilanthes notholaenoides community is restricted to intensively-grazed pasturelands frequently visited by dairy cattle. The diagnostic species are mainly found near heavily-trampled denudated sites, e. g. cow paths, where drainage is bad and erosion is taking place.

Here the original topsoil has largely been removed, and pH values at 15 em below ground level fluctuate between 5.1 and 5.8. The reddish-brown subsoil clay is many times exposed. This grassland community is found on both sides of the main road leading to San Gerardo de Dota, and covers very steep SandE slopes (50 to 70 degrees) behind the farmers houses.

1.2.

The Geranium guatemalense-Oenothera epilobifolia grassland

Structure and composition This plant community has an average height of 1 m, sometimes reaching 1.6 m at bushy places. A shrub layer, when present, may cover 30 %, a well-developed herb layer 80 %, and a moss layer 10 %. Scattered trees belonging to e. g. Buddleja nitida may occur as forest remnants offering shelter and shade to the cattle. The main floristic difference with the Orthrosanthus chimboracensis-Cheilanthes notholaenoides grassland community is the higher plant diversity, and the exclusive presence of dicots such as Geranium guatemalense, Oenothera epilobifolia ssp. cuprea, Phytolacca rugosa and Solanum rudepannum. The grass species Pennisetum clandestinum is even more abundant than in the former grassland community.

Other conspicuous species are the shrubs Ageratina subcordata, Archibaccharis aff. panamensis, Leandra aff. melanodesma, Monochaetum neglectum and Rubus eriocarpus, the herbs Conyza bonariensis, Hydrocotyle sp., Lachemilla spp. and Plantago australis, the grass Holcus lanatus, and the fern Blechnum occidentale.

Ecology and distribution The Geranium guatemalense-Oenothera epilobifolia grassland community is found in moderately-grazed pasturelands near settlements on N and NE slopes of 30 to 45 degrees, where drainage is good and erosion has not yet affected the topsoil seriously. At a depth of 15 em pH values range from 5.3 to 5.8.

30 Phytocoenologia 22

M. Kappelle et a!.

462

Due to a low intensity of cattle-grazing patches of densely-packed dicot species could develop inside e. g. Rubus dominated bushes, thus attracting numerous birds and insects (pollinators and seed dispersal agents).

1.3.

The Thelypteris rudis-Carex jamesonii grassland

Structure and composition This grassland community is 0.5 to 2 m tall, with a well-developed shrub layer covering over 50 %, a still large herb layer covering 70 %, and a tiny moss layer (10 % ). Characteristic species are the ferns Thelypteris rudis and Eriosorus flexuosus, the graminoid Carex jamesonii, and the forbs Hypochaeris radicata and Viola scandens.

Dominating but not exclusive species are the tree Buddleja nitida, the shrubs Ageratina subcordata, M onochaetum neglectum and Archibaccharis aff. panamensis, the thorny Rubus eriocarpus, the forb Conyza bonariensis, the grasses Pennisetum clandestinum and Holcus lanatus, and the ferns Pteridium aquilinum and Blechnum occidentale.

Ecology and distribution The Thelypteris rudis- Carex jamesonii plant community occurs on relatively steep slopes (45 to 55 degrees) with differing aspects (N, NE, SW, W). The soil sustaining this community is greyish-brown, clayey and at most places well-drained. In the topsoil at 15 em depth a pH value of 5.0 to 5.8 was measured.

Pasturelands housing this community are very rarely visited by cattle and therefore extensively grazed. Numerous palatable species can flourish without being browsed on, thus locally increasing plant species richness.

1.4.

The Galium mexican urn-Vaccinium consanguineum shrubland

Structure and composition Because of its differing structure (up to 4 m high) this community is classified as a shrubland although floristically it belongs to the Pennisetum clandestinum-Holcus lanatus grasslands. The high cover percentages of the widely-spread shrubs Ageratina subcordata, M onochaetum neglectum, Rubus eriocarpus and Vaccinium consanguineum, as well as the high abundances of the fern Pteridium aquilinum, the climber Smilax kunthii, and the treelet M on nina crepinii gives the Galium mexicanum- Vaccinium consanguineum community the appearance of a shrubland.

It has a shrub and a herb layer each covering 65 % and a moss layer covering about 10 %. The winding rubiaceous Galium mexicanum is an exclusive and almost constant feature found in this shrub land (cover percentages ranging from 1 to 7). Occasionally a forest remnant, e. g. Quercus seemannii or Comarostaphylis arbutoides var. arbutoides, is observed, which

Montane secondary vegetation in the Cordillera de Talamanca

463

locally may cover 30 to 75 %. Other species, such as the forbs Hieracium irazuense, Gnaphalium attenuatum and G. americanum, as well as the hemiparasitic Castilleja talamancensis, are less common, but characteristic for this community.

¡

Ecology and distribution The Galium mexicanum-Vaccinium consanguineum shrubland community has established itself on moderately steep E and NE slopes (20 to 50 degrees) with well-drained clayey soils. pH values in the topsoil at 15 em depth oscillate between 4.7 and 5.8.

This plant community is observed in recently abandoned pasturelands, called 'charrales'. These areas are now covered by numerous species, that could not flourish during the former period of cattle-grazing. Seeds-coming in by wind and dispersal agents from nearby forests germinate massively, giving rise to a rapid increase in plant diversity, especially woody species (KAPPELLE 1993 ).

2.

The Chusquea tomentosa-Oreopanax xalapensis forest

The lower montane Chusquea tomentosa-Oreopanax xalapensis forest includes at least two forest communities, that could establish themselves during secondary succession following abandonment of the Pennisetum clandestinum-Holcus lanatus grasslands. Both secondary forest communities are characterized by the exclusive presence of the aggressive bamboo species Chusquea tomentosa, the rapid-growing successional tree species Oreopanax xalapensis, the tall Cornus disciflora, the true pioneer tree Bocconia frutescens, the shade-bearing Clusia palmana, the shrubby Fuchsia microphylla, and the herbaceous climbers Bomarea acutifolia and Passiflora sexflora.

2.1.

The Freziera candicans-Quercus seemannii forest

Structure and composition This closed, secondary successional forest community is characterized by the dominant presence of 6 to 8 m tall Quercus seemannii and Freziera candicans trees in the canopy layer (about 30 % cover). Dense clumps of 4 to 5 m tall Chusquea tomentosa bamboos determine the appearance of the shrublayer (80 % cover). A second oak species, Quercus copeyensis, is characteristic for this community but not as abundant as Q. seemannii. Other prominent tree species are Ardisia costaricensis, Cleyera theaeoides, Comarostaphylis arbutoides var. arbutoides, flex valerii, Myrsine pellucidopunctata, Oreopanax xalapensis, and Viburnum costaricanum.

In the understory and on the ground one encounters species like the shrubby Miconia glaberrima and Leandra subseriata, the herb Relbunium hypocarpium, the terrestrial bromeliad Vriesea williamsii, the ferns Blechnum occidentale and Elaphoglossum aff. engelii. At slightly more open places

M. Kappelle et a!.

464

prominent species are Monochaetum neglectum, Vaccinium consanguineum, Pteridium aquilinum and Lycopodium thuyoides.

Ecology and distribution This plant community occupies the drier and relatively warmer, sun-exposed SE and NE slopes of the valley (slope angles of 35 to 45 degrees). The prevailing dry environment favours small coriaceous leaves as found in ericads.

The largely eroded clay-rich topsoil is extremely acid: pH values at 15 em soil depth fluctuate between 2.5 and 3.5.

Species known from the Galium mexicanum- Vaccinium consanguineum shrubland such as Monochaetum neglectum, Vaccinium consanguineum and Pteridium aquilinum are present with individuals displaying a reduced vitality. Several times M. neglectum was observed defoliating rapidly, while P. aquilinum was noticed decaying entirely, apparently giving way to young Freziera, Myrsine, Oreopanax, Quercus and Viburnum treelets.

2.2.

The Wercklea lutea-Cornus disciflora forest

Structure and composition The 8 to 12 m high Wercklea lutea-Cornus disciflora forest is the most species rich of all secondary plant communities found in the lower montane belt (Photo 2). It has a tree layer covering 60 to 70 %, which is dominated by the yellow-flowering, large-leaved tree Werklea lutea in association with tree species such as Cornus disciflora, Oreopanax xalapensis and Saurauia veraguasensis. The 5 m tall shrub layer with a cover of 50 to 70 % is again dominated by the bamboo Chusquea tomentosa. Characteristic and frequently found species are the trees Billia hippocastanum, Ocotea laetevirens, Persea rigens, Phyllanthus niruri and Tournefortia johnstonii, the palm Geonoma hoffmanniana, the shrubs Cestrum racemosa, Leandra costaricensis var. angustifolia and Palicourea brenesii, the herbs Anthurium concinnatum, Begonia udisilvestris, Hansteinia ventricosa and Piper lacunosum, the epiphytic herb Peperomia palmana, the climbers Jungia ferruginea, Liabum tonduzii and Smilax subpubescens, and the ferns Campyloneurum angustifolium, Cyrtomium macrosorum and Polypodium plesiosorum. Species repeatedly observed but not restricted to this community are the treelets Bocconia frutescens and Clusia palmana, and the climbers Bomarea acutifolia and Passiflora sexflora.

Ecology and distribution Secondary forests (co)dominated by Wercklea lutea occupy humid and relatively cooler, more shaded sites in the upper part of the lower montane belt.

So they are generally found on the somewhat wetter W slopes (slope angles: 35 to 45 degrees) or in protected valleys and along drainages, such as the Rio Savegre and its tributaries. Here, on very acid soils with pH values ranging

Montane secondary vegetation in the Cordillera de Talamanca

465

Photo 2. Secondary Wercklea lutea-Cornus disciflora forest occurs from left to centre, bordering primary Quercus seemannii forest in the background. Species-rich Geranium guatemalense-Oenothera epilobifolia grassland covers the slopes in the foreground . Two isolated Buddleja nitida trees appear on the right.

from 2.5 to 3.5, rapid-growing, large-leaved pioneer trees form a closed and dense vegetation, heavily visited by pollination and seed dispersal agents, that probably favour further forest succession towards a more mature stage.

Description of upper montane secondary plant communities (Table 3) 1.

Plantago australis-Rume x acetosella grassland

As has been stated above, the lower montane grasslands are dominated by grasses such as Pennisetum clandestinum and Holcus lanatus, whereas the upper montane grasslands are characterized by a series of exclusive forbs like Plantago australis, Rumex acetosella, Conyza bonariensis and Gnaphalium americanum. The only grass characteristic for these upper montane grassland communities is Bromus sp. Other species very abundant in the Plantago australis-Rumex acetosella grasslands, but not restricted to it, are the shrub Ageratina subcordata, the subshrub Acaena elongata, the herbs Lachemilla spp.- including L. pascuorum, L. pectinata and L. standleyi - as well as the spiny Cirsium subcoriaceum. These grasslands are 0.25 m high at sites sustaining merely herbs and up to 1 m at places where shrubs as Ageratina sub-

Table 3. Upper montane secondary plant communities, Costa Rica.

1.2.

1.3.

1. 4.

2 .1.

2. 2.

plant community

1.1.

plot numbe-r (01 to 30)

00000 00001 11111 11112 22222 22223 12345 67890 12345 67890 12345 67890

altitude (rn alt .)

22222 77777 28267 00000

22222 77777 68796 00000

22222 77777 96678 00000

22222 77777 65657 00000

22222 77777 97887 00000

22222 77877 89097 00000

! 1.1. ! ! !

Bromus sp. - Gnaphalium americanum grassland

-!><3' <3'

1.2. Halenia rhyacophylla Lolium perenne grassland

! 1.3. ! â&#x20AC;˘

Ageratina subcordata Muehlenbeckia tamnifolia grassland

aspect (exposition)

SNSEE WNNNN NNSSW ENENS SSSSS WSEEN w E E EE w EEEWE

slope angle (degrees)

12122 22333 34444 32343 33232 12222 55555 00000 50500 00500 55505 50005

topsoil pH (15 ern below level)

45555 55555 44654 54556 55555 45544

total number of vascular species

01101 11332 22222 22333 44555 43323 71391 88478 88515 49434 25211 69789

TWINSPAN division

00000 00000 00000 00111

00000 00000 11111 00001

00000 11111 00000 11000

00000 11111 11111 11111 00000 11111 11111 011 11 00001 00001

-----------------

-------2---------

plant species (001 to 114) arranged in sociological groups (I to XIII)

II

growth form

plant family Gentianaceae

TWINSPAN

1-1-3 2-112 1-------1--11 ---------------------1--1 -----------------------------1--- ----- - - ------- ----2 ----- --- --

122-----1 ------------------------------------ 1 ----1 ----1 -- ---- ---------

0001 0001 0001 0001 0001 0001 0001 0001 0001 0001 0001 0001 0001 0001 0001 0001 0001 0001 0001 0001

shrub herb herb herb herb herb herb herb herb herb herb herb fern-ally shrub herb herb herb herb herb herb

005 006 007 008 009 010 011 012 013 014 015 016 017 018 019 020 021 022 023 024

Acaena elongata Lachemilla spp .

Plantago australis Rumex acetosella Cirsium subcoriaceum Gnaphalium americanum Bromus sp.

Veronica arvensis

33225 87465 57434 45774 636-3 - 2251 24265 -421--2- ----4 -----3------------4-----------------3 3----

85888 14447 35555 24545 25442 12433 23442--32 34434 35434443--665 1-221 --232 -22-4 -323--42--23-

Juncus bufonius

division

herb herb tree herb

5255-43---24-3---

Erigeron subspicatus

Quercus costaricensis Quercus copeyensis forest

0000 0000 0000 0000

------------- - ---

Cyperaceae spp.

Eragrostis sp.

Fuchsia arborescens Abatia parviflora forest

1-------2 --------

Halenia rhyacophylla Relbunium hypocarpium Drymis granadensis Senecio oerstedianus

Oenothera epilobifolia Viola nannei Hydrocotyle sp.

Lycopodium clavatum(contiguum Hesperomeles heterophylla Holcus lanatus Lolium perenne

Pteridium aquilineum Polypodium macrolepis shrub land

--311 1-----------1

001 002 003 004

Geranium guatemalense

! 1. 4. ! ! ! ! 2 .1. ! ! ! 2.2.

66213 76555 352-1 45555 53432 23222 251-2 2-11342 --3--3---34--------------4 ---1-----

55555 53433 34332 44442 --3--1--2 42--2-----2- --3-2 ---42 -----2-----3-- --------22-

--- - -

-- -- -----

Rubiaceae Winteraceae

Asteraceae Rosaceae Rosaceae

Plantaginaceae Polygonaceae Asteraceae

Asteraceae Poaceae Scrophulariaceae Geraniaceae

Onagraceae Violaceae Apiaceae Lycopodiaceae

Rosaceae Poaceae Poaceae

Cyperaceae

Poaceae Asteraceae Juncaceae

s;:: :;>::

"'"'"'

~

" ~

~

III

IV

v

VI

VII

025 026 027 028 029 0 30 03 1 03 2 03 3 034

03 5 0 36 037 0 38

Conyza b o nariensi s

Gnaphalium attenua t um Gnaphalium elegan s Pteridium aqu i li n um Polypodium plebej um Car ex j am esoni i

Lycopodi um t huyoides S tellaria ova t a Polystichum s p .

Comarostaphyl i s arbut oi des Ageratina s ubcordat a Rubus spp .

Pleopelti s macrocarpa Chusquea talamancensis

039 Pernettya c oriacea 0 40 Styrax glabrescens 041 Eri o sorus tlexuosus

herb herb herb fern fern he r b f ern - ally herb f ern tree

Asteraceae

- ---- --- ------ --- ------ --- --

0010 0010 0010 0010 0010 0010 0010 00 1 0 0010 00 1 0

--1- 4 11 1 13 -- 1 --1-- -

0011 0011 00 11 00 11

sh ru b shru b f ern bamb oo

Asteraceae Rosaceae

----- 2 --- - 2 1-- 2 34 45 4 1- 3 - - 111--- --- - - --- - - -- - -- 22- --- - - --1------ ----- - - -1- -- - -- - --- 2 ---- -

0 1 00 0 100 0 100

s hrub t ree fe r n

Ericaceae

2- 432 11 3-2 -12-2 76756 3442 ----3 - 331- -1 -2 - 2-- 5 ---

1-----1- - --- ----1- -- --- --

-1 413 3-333 5 477 4 4444 5 --- - - -4553 44444 455 44 -- --- -- --3 - ---- 2 - 22 -- - -- -5--- ----4

42233 3 1111 -- - - - - ---

22222 1 22 3 3 - 1112 - 4- 32 -2 - - 5-122 - - - -- - 111 --1--------3 - -- --

- - 223 ---------------- --- - -- -- --2- -- -- -

---2 4 -- 33 -- 12 2 --44 - - 2-4 2---- 2 4- -- -2-----

Me l a stomatac e ae

shrub climb er fern climber herb herb tree

Al s troemeri acea e

2 4--44---2-------

2---33415 2--3-----

- 33342246 111-1 -----

3 -1-2 11-2 --11--1--

011 011 011 011

fern shrub fern fern

Dr yopteridacea e On a graceae Po lypo diace a e Polypodiacea e

- ---- --55- 3-1-- -43-- -64-- 33222 23-31 31123 11--3 --224 ----- ----- ---24 ---33 ---2 2 22 231

100 100 100

tree tree tree

Ericaceae As te raceae

-- --- ---2 - --3-- ----2 1-3-- 112-1 ----- ----- ----- ---3- ---1- 12112

101 101

tree shrub

So lanaceae Ericaceae

1100 1100 1100 1100 1100 1100 1100 1100 1100

tree

Caprifoliaceae

tree herb herb herb tree tree herb herb

Dryopteris wallichiana Fuchsia microphylla Polypodium skinneri Campyloneurum angustifolium

- - ---------------

35442 --444 ---12 ----2

Viburnum costaricanum Buddleja nitida Arenaria lanuginosa Oxalis spiralis Vriesea williamsii Escallonia myrtillioides Monnina crepinii Peperomia saligna Nertera granadensis

Styr acace a e Adian taceae

01 0 1 01 0 1 0 10 1 0 10 1 0 101 0101 0101

049 050 051 052

058 059 060 061 062 063 064 065 066

Poaceae

- 1 --- - 3 14 -1221 111-1 ----1111-21--

-- ------- - - --- ---- --- - -------

X

Polypodiaceae

4-434 411-1 --111 2 1111 41321 - - 1-1 -2- --

Monochaetum neglectum Muehlenbeck i a tamnifolia Polypodium macrolepis Bomarea acutifolia Castilleja talamancensis Peperomia galioides Myrsine pellucidopunctata

056 Solanum storkii 057 Macleania rupestris

Eri cacea e

75557 43-5444 3 2 -2 3 2 3 42343 -2 3 22 --422

042 043 044 045 046 047 048

IX

Asteraceae Asterac eae Dennstaedtiaceae Po lypodiaceae Cyperaceae Lycopodiaceae Caryophyll aceae Dryopt eri daceae

4-452 43355 12 --2--1--2----------

-- - ------- -- 3 ---2- - ------- ----

VIII 053 Vaccinium consanguineum 054 Senecio multi veni us 055 Cleyera theaeoides

---- ---- ---- 1 -- - - 3 ------ - - -

-------------------------------------

--2-2 --------3 --2---- - 2 ---5-------------

1----2422 -3--1-1-------------1--------

-2222 -3--5 --2--------2----3---2 --------1

25233 46365 41112 11111 11111 -3--5 6--1---------

12231 ----3 1 - --1----1-11 --------1 2---1----

Polygonaceae Polypodi acea e

~

0

.,g::> (1)

"'n (1)

0

::>

"'"' ~ <

(1)

""~

~

o¡ ::> s¡ ....

::r (1) () 0

'"' g;

Sc r o phulariaceae

;:;-

Piperaceae Myrsinaceae

"'(1)

"''"'

~ ;:;;-

3

"'::>n "'

Th eaceae

Loganiaceae Caryophyllaceae

Oxalidaceae Bromeliaceae Escalloniaceae Po lygalaceae

Piperaceae Rubiaceae

-l>o 0"'I

Table 3. (cont.) plant community

1.1.

plot munber (01 to 30)

00000 00001 11111 11112 22222 22223 12345 67890 12345 6789 0 12345 67890

XI

-- -----------

XII

067 068 069 070 071 072 073 074 075 076 077 078 079 080 081 082 083 084 085 086 087 088 089 090 091 092 093 094 095 096 097 098 099 100 101 102 103

XIII 104 105 106 107 108 109 110 111 112 113 114

Fuchsia arborescens

Abatia parviflora Bocconia frutescens

Centropogon costaricae Begonia udisilvestris Verbesina oerstediana

Malaxis hastilabia Carex sp. 2 Miconia sp.1 Asplenium viridissimum Cystopteris fragilis Sigesbeckia jorullensis Valeriana scandens Polystichum muricatum Calceolaria trilobata Dendropanax querceti Centropogon valerii Polypodium loriceum Asplenium harpeodes Myrsine coriacea

Oreopanax xalapensis Cornus disciflora Smilax kunthii Chusquea tomentosa Weinmannia pinnata Zanthoxylum scheryi Styrax argenteus Ardisia sp.

Jungia ferruginea Hydrocotyle ribifolia Symplocos serrulata Campyloneurum amphostenon Elaphoglossum aff. engelii Peperomia hylophila Ilex pallida Mikania cordifolia Passitlora membranacea

Quercus copeyensis Quercus costaricensis

Oreopanax capitatus Hedyosmum mexicanum Rhynchospora aristata Anthurium concinnatum

Palicourea salicifolia Elaphoglossum conspersum Blechnum costaricense Ilex discolor Ardisia compressa

1.2.

1. 3.

1. 4.

2 .1.

2. 2.

----4 7223~ ----- ---- - 34165 24131 --- - - --- -- 31123 ----11121 -3243 ----- ---- - -1111 -----33-1 ----- ----- -113----- ----- . 11-111-1-111----- --- - -- - -- --111 ----- --- -- 4---3 ----- ----- 2---1 ----- ----- --2-1 -----1-2----- ----- -- 12----1---2

-------------1--1 1---- 1--1---1----

1----1--------1----------

-I>-

cr-.

-------------

1101 1101 1101 1101 1101 1101 1101 1101 1101 1101 1101 1101 1101 1101 1101 1101 1101 1101 1101

tree tree tree herb herb tree herb herb tree fern fern herb herb fern herb tree herb fern fern

-5313 35332 25465 15433 56651 -7655 -1221 -3321-231 -1111 --112 --11-11--1-1---2----1 -1 - ----1-

51237 212 -1 34131 15531 66577 34222 41221 21 - 31 --2 3--111 2---1 1-2---11-11-21---11-1---1 ----

1110 1110 1110 1110 1110 1110 1110 1110 1110 1 110 1110 1110 1110 1110 1110 1110 1110 1110

tree tree tree climber bamboo tree tree tree tree climber herb tree fern fern herb tree climber climber

----- ---1- ----4 ----- 43- ------ -------2- --1--- --- -- ---1------- ----- ----- ----- - - -2----- ----- -------1----- ------------- ----- ----- ----- - ---------------- ----- ----- ----- --------- --------- --------- ----- ----- ----- -----

88876 37563 -1-21 4---1 -1111 -- 11 --2-1 11--1---1 -1-1-1-1-

1111 1111 1111 1111 1111 1111 1111 1111 1111 1111 1111

tree tree tree tree herb herb shrub fern

------------- --------- -----

----- --------- ------------- -----

--------- --------- --------- ----- ---- --------- -----

--------- --------- --------- ------ -----------------------------------

2--2---1----- - ---------3-

-4--2 --2---332 --5-1 ------ ---

----- - -- -- --------- ----- ----------------- -- --- --------- ----- "-- -- - -------- --------- ----- ----- --------- ----1 ----- -----

----- -----

- - --- ---- - ---- - --------- ----- ----- -----

-----

-------------

-----

---------

-----

fern

tree tree

Onagraceae Flacourtiaceae Papaveraceae Carnpanulaceae Begoniaceae Asteraceae Orchidaceae Cyperaceae Melastornataceae Aspleniaceae

00

Woodsiaceae

Asteraceae Valerianaceae Dryopteridaceae Scrophulariaceae Araliaceae Campanulaceae Polypodi aceae Aspleniaceae Myrsinaceae

Araliaceae Cornaceae Smilacaceae Poaceae Cunoniaceae

Rutaceae Styracaceae Myrsinaceae

Asteraceae Apiaceae Symplocaceae Polypodiaceae Lomariopsidaceae Piperaceae Aquifoliaceae Asteraceae Passifloraceae Fagaceae Fagaceae

Araliaceae Chloranthaceae Cyperaceae Araceae

Rubiaceae Lomariopsidaceae Blechnaceae Aquifoliaceae Myrsinaceae

~

.,~ '0

'0

a.

" ~

~

Montane secondary vegetation in the Cordillera de Talamanca

469

Table 3. (cont.) plant species found in only one plot

Andiantum andicola Arachniodes denticulata

plot cover growth plant nr. class form family

21 26 Asplenium serra 26 Cestrum racemosum 25 23 Didymaea alsinoides Digitalis purpurea 02 Elaphoglossum biolleyi 29 Elaphoglossum squamipes 17 Garrya lauri folia 27 Hydrangea asterolasia 26 25 Miconia sp. 2 Palicourea adusta 30 15 Phytolacca rugosa Roldana heterogama 09 Scheffler a rodriguesiana 30 21 Sipthorpia repens Viola scandens 26 Veronica polita 08

1 1 1 2 1 4

1 3 2 1 4

1 2 1 1 4

1 3

fern

Adiantaceae

fern fern shrub herb herb

Dryopteridac.

Aspleniaceae Solanaceae

fern

fern tree

Rubiaceae Scrophulariac.

Lornariopsidac.

Lomariopsidac.

Garryaceae

climber Hydrangeaceae Melastomatac. tree shrub Rubiaceae

herb shrub tree herb herb herb

Phytolaccac.

Asteraceae Araliaceae Scrophulariac.

Violaceae Scrophulariac.

cordata, tall herbs as Cirsium subcoriaceum, or ferns as Pteridium aquilinum dominate.

In the present study four different Plantago australis-Rumex acetosella grasslands are distinguished, which are described below.

1.1.

Bromus sp.-Gnaphalium americanum grassland

Structure and composition This 0.25 to 0.5 m tall grassland community is characterized rather by the absence than the presence of many species typical for the Plantago australis Rumex acetosella grasslands. A very poor herb layer covering over 80 % is dominated by Rumex acetosella and Plantago australis, together with different Lachemilla species (L. pectinata and L. standleyi among others), Cirsium subcoriaceum and Bromus sp. Woody Acaena elongata and Ageratina subcordata occur in a patchy pattern. Less abundant are Gnaphalium americanum, Conyza bonariensis and Veronica arvensis. The grass Holcus lanatus has been found in only one plot.

Ecology and distribution This grassland type is observed in the vicinity of farm houses and intensively grazed by dairy cattle. It appears to be less productive in terms of edible grasses then the Pennisetum clandestinum-Holcus lanatus pasturelands of the lower montane belt. Initial erosion features are common, but the original topsoil is still greatly present, especially on gentle S and E slopes (15 to 25 degrees). Topsoil acidity is not as strong as in other grassland communities displaying pH values oscillating around 5. At different places near ground level charred wood fragments are found originating from nearby charcoal

M. Kappelle et al.

470

pits. This evidence indicates a considerable nutrient influx into the topsoil, which probably affects the overall species composition. The occasional presence of horses which carry sacks of charcoal to the main road may locally increase soil nutrient levels even more.

1.2.

Halenia rhyacophylla-Lolium perenne grassland

Structure and composition This secondary plant community with heights sometimes over 1.5 m is much richer in species than the Bromus sp.-Gnaphalium americanum grassland. It has a shrub layer (average cover 60 %) dominated by the sub shrub Acaena elongata and the shrubs Ageratina subcordata and Rubus spp. (among others R. eriocarpus). A well-developed herb layer (70 %) is occupied by prominent species like Cirsium subcoriaceum, Geranium guatemalense, Halenia rhyacophylla, Hydrocotyle sp., Lachemilla spp., Oenothera epilobifolia ssp. cuprea, Plantago australis, Rumex acetosella, Viola nannei, and the fern Dryopteris wallichiana.

Ecology and distribution The Halenia rhyacophylla-Lolium perenne grasslands are mainly found on gently and moderately steep N and NE slopes of 15 to 30 degrees. Soils are clayey and relatively well-drained with topsoil pH values around 5. Cattle grazing is moderate, favouring the establishment of low shrubs inside vegetation nuclei of spiny, bushy or less palatable species ( Cirsium subcoriaceum, Acaena elongata, Rubus eriocarpus and Pteridium aquilinum among others).

1.3.

Ageratina subcordata-Muehlenbeckia tamnifolia grassland

Structure and composition This 1 to 2 m high grassland community is dominated by several species of low shrubs, forbs and ferns. The shrub layer is less dense (average 30 % cover) than the herb layer (65 % cover). The omnipresent Ageratina subcordata and Acaena elongata together with Monochaetum neglectum and Rubus spp. determine the shrub layer, while species such as Cirsium subcoriaceum, Lachemilla spp. and Rumex acetosella continue to dominate the herb layer.

The woody climber Muehlenbeckia tamnifolia characterizes the overall aspect of the vegetation, occupying sites in both the herb and shrub layer.

Other prominent herbs are Gnaphalium americanum, G. attenuatum and G. elegans, as well as Conyza bonariensis and the cyperaceous Carex jamesonii.

Ecology and distribution The Ageratina subcordata-Muehlenbeckia tamnifolia grassland community occurs on steep S, Wand NW slopes (slope angle: 35 to 45 degrees) sustaining extensively grazed pasturelands. Soils are well-drained and rich in clay, with topsoil pH values between 3.7 and 5.8.

Montane seco ndary vegetation in the Cordillera de Talamanca

471

Muehlenbeckia tamnifolia forms thick blankets of leaves covering the crowns of mainly Ageratina, Monochaetum and Rubus, thus reducing the light available for these taxa. In this way M. tamnifolia prepares the road for shade-tolerant woody follow-up species, such as Cleyera theaeoides and Solanum storkii.

1.4.

Pteridium aquilinum-Polypod ium macrolepis shrubland

Structure and composition The 3 m tall Pteridium aquilinum-Polypodium macrolepis shrubland community belongs floristically to the Plantago australis-Rumex acetosella grasslands, although it is structurally quite different from these. The aspect of the shrub layer covering 70 % is determined by bushy elements such as Monochaetum neglectum and Pteridium aquilinum (Photo 3). The herb layer covers less than 30 %, while a bryophyte layer becomes important reaching over 40 %. Taxa like the hemiparasitic Castilleja talamancensis, the shrubby Pernettya coriacea and the fern Polypodium macrolepis occur for the first time in considerable quantities. Other species known from the above-described upper montane grasslands are still present in high numbers. Among these occur Acaena elongata, Ageratina subcordata, Lachemilla spp., Rubus spp. and to a lesser extent Plantago australis, Rumex acetosella and Conyza bonariensis. Certain other species such as the woody Comus disciflora and Fuchsia microphylla usually dominate locally.

Ecology and distribution The upper montane fern-shrublands inhabite well-drained clay-rich soils (pH 4.3 to 5.6) on mainly moderate to steep E slopes (20 to 40 degrees). This bushy community may develop after complete abandonment of the 'true' Plantago australis-Rumex acetosella grasslands and probably marks the start of a recovery process (secondary succession) that may lead to more complex vegetation types, such as Chusquea tomentosa- Weinmannia pinnata forest communities described below. Bird- and insect-attracting shrubs favour a continuous seed input that enhances the process of forest recovery in a substantial way.

2.

Chusquea tomentosa-Weinmannia pinnata forest

Within the upp er montane Chusquea tomentosa-Weinmannia pinnata secondary forest vegetation two different forest communities have been distinguished. They have a large number of species in common, such as the dominating bamboo Chusquea tomentosa, which is also known from secondary forests in the lower montane belt (see above). The woody climber Smilax kunthii, the trees Weinmannia pinnata, Myrsine coriacea, Zanthoxylum scheryi and Styrax argenteus are characteristic species in these forest commu-

M. Kappelle et a!.

472

Photo 3. Pteridium aquilinum-Polypodium macrolepis shrubland front right. Note the large clump of decaying Chusquea tomentosa bamboos to the left and the recovering patch of secondary Quercus costaricensis-Q. copeyensis forest to th e right, behind the Pteridium aquilinum ferns. At the back primary Quercus costaricensis forest blanketing the higher ridges.

nities. Other species occurring in both types, but not restricted to them, are Cleyera theaeoides, Comus disciflora, Vaccinium consanguineum, and Viburnum costaricanum.

2.1.

Fuchsia arborescens-Abatia parviflora forest

Structure and composition This 8 to 12 m tall secondary forest has a tree layer covering about 7S % and a shrub-herb layer covering 40 to SO %. A well-developed bryophyte layer comprising terrestrial mosses reaches a cover of about SO %. Characteristic codominating tree species are Fuchsia arborescens, Abatia parviflora and Bocconia frutescens. Other important tree species are Weinmannia pinnata, Myrsine coriacea, Zanthoxylum scheryi and Styrax argenteus. The bamboo Chusquea tomentosa dominates the shrub and herb layers, while the climber Smilax kunthii ascends its culms. Among the characteristic herbaceous species one finds Begonia udisilvestris, Oxalis spiralis ssp. vulcanicula and the orchid Malaxis hastilabia. Numerous successional species present in the

Montan e second ary vegetation in the Cordillera de Talamanca

473

Plantago australis-Rumex acetosella grasslands have not yet disappeared and occur frequently in this forest type. We mention Ageratina subcordata, Buddleja nitida, Fuchsia microphylla and Monochaetum neglectum. Others, such as Cornus disciflora and Viburnum costaricanum known from the Pteridium aquilinum-Polypodium macrolepis shrubland, flourish even better here.

Ecology and distribution The Fuchsia arborescens-Abatia parviflora forest community is situated on well-drained, humid and moderately steep S slopes (20 to 35 degrees). Topsoil pH values at 15 em depth in studied releves range from 4.1 to 5.3.

This secondary forest community could establish itself after a recovery period of a dozen years since farmers and their cattle abandoned the area (KAPPELLE 1993 ). A number of early secondary trees such as Fuchsia and Abatia, but also Cornus, Viburnum and Buddleja in an earlier stage, could form a matrix of dense patches of regrowing forest, that offer a new habitat for displaced avian and mammal communities coming in from nearby tracts of mature (primary) oak forest.

2.2.

Quercus costaricensis-Quercus copeyensis forest

Structure and composition This secondary oak forest community is represented by closed, 10 to 12 m tall stands dominated by Quercus costaricensis and Q. copeyensis in the tree layer (average cover 70% ), and 5 m tall Chusquea tomentosa bamboos in the shrub layer (average cover 50 % ). Accompanying species with high cover percentages are the tree species Cornus disciflora, Myrsine coriacea, Weinmannia pinnata and Zanthoxylum scheryi, and the climber species Smilax kunthii. Less abundant, although frequent are the woody Vaccinium consanguineum and Viburnum costaricanum. The understory tree species !lex discolor var. lamprophylla and I. pallida, as well as the facultative epiphyte Anthurium concinnatum known from primary oak forests KAPPELLE et al.

1989) are several times found.

Ecology and distribution Upper montane secondary oak forests are spread over gentle, relatively drier E and W slopes (15 to 25 degrees), further away from farmers' houses, and many times adjacent to primary (mature) oak forest stands. They root in clayey soils with topsoil pH values ranging from about 4.1 to 4.7.

These successional Quercus forests have developed a more complex forest structure expressed in a canopy of densely-packed oak tree-crowns creating dark conditions in the understory. Thus at ground level shade-tolerant species get a chance to germinate and thrive, while taking effective advantage of dispersed light pouring through the canopy. The earlier mentioned aroid Anthurium concinnatum is one of the many specialists displaying such a strategy.

L 0

M 0

W E R

I""'' ' ' ' ' ' ' ' '

m

LI 10

cover(%) 1 .1

tree layer shrub layer herb layer moss layer

M M U N I

100

0

100

0

C 0

N T A N E

I

I

I

1. 2

}~

E

S

-!>'-1 -!>-

100

0

10

'E

~r-

E

Ol

·a;

..c:

rn 0

I

t"

I

cover(%)

T

-0

100

0

100 I

0

I

~

I

I

I

cover(%)

I

110 2 .1

I E

Ol

·a;

..c:

100

0

l:~:;:~~~:::n:t~tf:~r:,::):;:::::r~~:~:~:t:m:,:::

cover(%)

;~

. 2 I 10

.s :.cOl

·a;

..c:

~ 7::

~

'U ~

" ~

0

~

U P P E R

0

100

I

I

I

I

cover(%)

L1o

I

1.1

C 0 M M U N I

MONTANE

100

0

I

I

I

I

cover(%)

I

1.2

T I

E S

100

0

11 o

I

I

I

I

cover(%)

I

1 .3

2:::

11o

0

" ""'"' M

"'"'0 ()

",_,

Q..

E" ~

.:E 0> ·o:;

"'

-

II

II

.s::

'-<

<

"""'g

~

o· s·

::>

;;..

Ism

\ : :r::\:rr\ 1 I-o lr'\r 111111111111111111111111111

rn

0

100 cover(%)

1. 4

I

I

lill~rrr~~:!:n III1111111111

o

100

0 2.1

I

0 0

I

"'() ~

o

100

s,_,

""' Q..

10

"'

~

'"8"' ::> ()

"'

-1>'..J

Fig. 4. Comparative structure diagrams showing height (m) and cover (% ) of the tree, shrub, herb, and moss layer of six lower and six upp er montane secondary plant communities in the Costa Rican Cordillera de Talamanca.

V1

M. Kappelle et a!.

476

Differences in structure, diversity and growth form spectrum Comparing structure diagrams that belong to the different plant communities described above, one immediately notices the great similarity between physiognomical aspects occurring at lower and upper montane elevations (Fig. 4).

In general, montane grasslands (1.1., 1.2. and 1.3.) are characterized by a dominant herb layer, next to a less prominent shrub layer, which may develop successfully when grazing intensity is reduced. Shrublands (1.4.) are dominated by a shrub layer, harbouring a variety of herbaceous and woody species.

These are accompanied by a number of species not reaching above the herb layer. Secondary forests (2.1. and 2.2.), however, are not always dominated by a prominent tree layer. In the lower montane successional forests instead, the shrub layer defines the overall aspect. This is expressed in the open character of these forests, probably due to the short period of recovery following abandonment. In the upper montane secondary forests, on the other hand, a more complete canopy closure has taken place. Here an important tree layer, sometimes over 10 m high, covers over 70 % of the surface.

To finally illustrate differences in structure and composition the lateral aspect of the studied communities is visualized in Fig. 5. Species typical of the distinguished structural layers are shown. Sometimes important species are found in both vegetation belts. As a rule structural and compositional changes occur gradually along the two supposed seres. Yet the change from shrubland to forest is abrupt. A probable explanation is the difference in landuse history and the length of the recovery period already mentioned above. According to VAN VELZEN & WIJTZES (1990) these forests may have passed through a period of 10 to 20 years of regrowth, whereas the shrubLegend

to

the characteristic species shown in Fig. 5.

®

Abatia parviflora

® CD

Acaena elongata

0

@

•

Oreopanax xalapensis Quercus copeyensis

®

Quercus costaricensis

Buddleja nitida

@

Quercus seemanii

®

Cirsium subcoriaceum

@)

Rubus spp.

0

Comus discif/ora

~

Viburnum costaricanum

<ID CD 0

Freziera candicans

0

Weinmannia pinnata

Q

Mannina crepinii

CID

Monochaetum neg/ectum

@

Myrsine coriacea

Ageratina subcordata

Fuchsia arborescens Fuchsia microphylla

e

Wercklea Jutea

~ Nt;jJ ~

Chusquea tomentosa

herbs and ferns

'-'

1.2

1 .1

10

1. 3

1.4

2.2

2.1

0

'<

8

8

8

0 0

"'0 .,0()

6

6

4

4

0 ;:;·

aq N N

"'

;:l

"' 'J>

()

"';:l

s::: 0

g

>-;:;

::r-

;:l

~ <

"'

aq

~

;:; 2

2

c;·

;:l

5" 0

5-

"'

()

.., §:;

0

UPPER

MONTANE

"..,"'

COMMUNITIES

height (m)

0<1>

:;;'

;:;--

10

10

8

"';:l ()

8

8

6

6

4

4

2

2

0

I I

\1 Yl

I [

I V

I Vf I \,I I J

1 N

V

I I l,l1 I 1'=, et:J:I I I I 1/

y

XI

....-r-,.,) V

I J J'\

I I (I/ I I I

n

I I

J

I 1/ I I I I 1 !)

Ct::=.b

N

I I I I I

Q

Fig. 5. Schematic vegetation profile diagrams of six lower and six upper montane secondary plant communities in the Costa Rican Cordillera de Talamanca.

"'

4>-

'-J '-I

478

M. Kappelle et a!.

land types have been abandoned only recently. Studies concerning forest recovery at 2900-3000 m alt. actually being carried out may reveal more explicit details on the topic (KAPPELLE 1993).

Considering patterns in diversity and growth form spectra along the two supposed successional seres, the following main trend is seen: an increase in species number with increasing structural complexity and decreasing human influence (Table 4 and 5). While the number of tree species is rising quickly, the number of herbs remains more or less the same. The numbers of shrub and climber species rise much slower, whereas fern species may double or triple along the successional gradient. All in all, trees and herbs are the most important growth forms in montane secondary vegetation types. Palm species are very few and only found at recovering lower montane sites. Bamboos (Chusquea spp.) are represented only poorly, but dominate entire structure layers .

Table 4. Number of species per growth form for six lo wer montane secondary plant communities in Costa Rica.

1.1.

1.2.

1.3.

1.4.

2.1.

2.2.

total

trees shrubs herbs climbers palms bamboos ferns fern -allies

3 8 14 1 0 0 7 0

5 6 17 1 0 0 7 0

6 6 13 3 0 1 8 0

7 5 13 4 0 1 5 1

33 9 8 5 0 1 13 2

39 16 15 10 3 1 22 1

53 23 40 12 3 1 31 2

total

33

36

37

36

71

107

165

Table 5. N umber of species per growth form for six upper montane secondary plant communities in Costa Rica.

1.1.

1.2.

1.3.

1.4.

2.1.

2.2.

total

trees shrubs herbs climbers palms bamboos ferns fern-allies

0 2 16 0 0 0 0 0

10 7 28 1 0 0 7 2

9 6 25 3 0 8 0

15 8 22 3 0 1 7 2

29 8 27 6 0 1 13 0

28 10 20 7 0 2 13

36 12 50 7 0 2 23 2

total

18

55

52

58

84

81

132

Montane secondary vegetation in the Cordillera de Talamanca

479

Discussion and conclusions

"

The phytosociological analysis of Costa Rican montane secondary vegetation here-presented shows a clear gradient of decreasing human impact from short, heavily-grazed pastureland communities to relatively tall secondary forest communities. This drop in intensity of anthropogenic activities leads to more diverse and structurally more complex plant communities. Although the same structural types (grassland, shrubland, forest) were found at both altitudes, i.e. near 2300 (lower montane) and 2800 m a.s.l. (upper montane), floristic compositions differed largely, resulting in a main division between both vegetation zones during the TWINSPAN classification process. This led to two different vegetation tables (a lower and an upper montane one), each representing a possible successional sere of plant communties inhabiting either actively-used, or recently-abandoned, or partly-recovered sites. These communities could be classified and described by a series of characteristic species with different growth forms and occupying one or more structural layers.

Firstly, all these communities belong to a Monochaetum neglectum-Rubus eriocarpus vegetation complex, of which the syntaxonomical status sensu BARKMAN et al. (1986) is still uncertain. Secondly, if one takes a look at both TWINSPAN dendrograms (Fig. 3) a clear division between grasslands (including shrub lands) and (successional) forests is observed. This evidence made us suggest provisional names for the complete series of lower and upper montane grasslands s.l. and secondary forests, respectively. The syntaxonomical status of these broad grassland and fores t types, too, is not yet completely clear.

On the other hand, specific plant communities within these structural formations have been recognized, treated extensively, and described on basis of five releves each. They have been named after their most characteristic species. Since further research in the study area is needed, especially at elevations around 2500 and 2600 m, no official syntaxonomical names have been proposed for the present communities. Moreover, the primary plant communities in the area have not yet been classified with multivariate techniques (KAPPELLE et al. 1989). These undisturbed, primary (oak-bamboo forest) communities form the basis for a thorough understanding of the secondary plant communities which in some way are derived from the former. Fortunately, primary, mature Chusquea- Quercus communities inhabiting the Chirrip6 National Park in the central-western part of the Cordillera de Talamanca are actually being studied and will subsequently be described, putting emphasis on their altitudinal distribution (KAPPELLE 1991, KAPPELLE, VAN UFFELEN & CLEEF in prep.). This will make possible a complete phytosociological analysis of the whole complex of primary and secondary (terrestrial) plant communities. In such a phytocoenological analysis the communities present in the nearby tropic-alpine paramo vegetation belt above 3200 m alt. will be included as well (CHAVERRI & CLEEF in prep.), especially since many species characteristic for upper montane grasslands have turned out to be true paramo species, that could migrate downslope as a result of human activities like indiscriminate deforestation (KAPPELLE et al. in prep.).

480

M. Kappelle et a!.

Considering the here-described plant commumues as apparent stages along two possible successional seres, one must take into account the unfortunate lack of reliable data on the former land use and cover in the study area on the one hand, and on the period of recovery since abandonment with respect to the shrublands and forests on the other hand. Oral information from local peasants farming on land, where releves were laid out, was rather incomplete and did not lead to a continuous record of the land use history.

Usually local people do not remember exactly what they did, when and where, and what has happened to the different sites since clearing of primary forest took place in the past. Moreover, many of the farmers actually inhabiting the study area did arrive after clearing and initial grazing started about fourty years ago. Thus, in order to obtain more reliable information on the vegetational past of the study area, aerial photographs were consulted (GEUZE 1989, SPREUWENBERG 1989, VAN VELZEN & WIJTZES 1990). However, only a set of photos on a scale of 1 : 60 000 which dated from 1969 was available at the Costa Rican National Geographic Institute at the moment of field research. On these photographs, secondary grasslands, shrublands and forests could be depicted, but no specific plant communities were identified.

Thus only general information on landuse in the area two decades ago was generated. This restriction made us decide to describe the floristic composition of the secondary vegetation types and not the hypothetical chronosequence. A monitoring study of permanent plots can focus on these aspects in the near future. Moreover, plant communities now occurring in one vegetation table may represent stages belonging to different successional pathways. The case of the two lower montane secondary forest communities is illustrative. Community 2.1. dominated by Freziera candicans and Quercus seemannii inhabits much drier locations than forest type 2.2., which is characterized by the pioneering, large-leaved malvaceous species Wercklea lutea.

Numerous ericaceous and theaceous species in the Freziera-Quercus forest are xeric in distribution, while palms, rubiaceous trees, herbs like Begonia udisilvestris and Hansteinia ventricosa as well as many ferns flourish in the hygric habitat of the Wercklea-Cornus forests . The latter is also the most species-rich community described in the present study, precisely because of its warmer and wetter environment on W slopes, which are better protected from strong tradewinds and intense sunlight than e. g. NE slopes sustaining Freziera-Quercus forests. Thus it is not expected that the humid Wercklea forest will make way for a drier Freziera forest when time advances.

A brief comparison between the here-presented secondary Quercus forests and the primary oak-bamboo forest communities, earlier described

by KAPPELLE et al. (1989), reveals an interesting similarity. It results that the lower montane Freziera candicans-Quercus seemannii forest has numerous species in common with the primary Quercus seemannii- Q. copeyensis forest, known from 2300 m alt. in the study area. The dominant presence of Quercus seemannii and Chusquea tomentosa is diagnostic for both the primary and the secondary forest type. Other woody species frequently observed and occurring at undisturbed as well as at partially regrown forest sites are Ardisia costaricensis, Fuchsia microphylla, Miconia glaberrima, Oreopanax

Montan e secondary vegetation in the Cordillera de Talamanca

481

xalapense and Styrax argenteus. A similar pattern is seen in the upper montane belt at an altitude of about 2800 m. There, secondary Quercus costaricensis - Quercus copeyensis forest stands show a floristic composition similar to that of primary forests dominated by the same combination of oak species. In the latter KAPPELLE et al. (1989) found characteristic species such as !lex discolor, I. pallida, Macleania rupestris, Symplocos sp. and Schefflera rodriguesiana, which also occur in the secondary forests here presented. A profound study by the first author and fellow-workers on differences between these primary (mature) and secondary (recovering) Quercus forests near 3000 m a.s.l. in the same area will help us understand the prevailing trends in diversity along the successional gradient (KAPPELLE et al. in prep.).

One interesting mechanism stimulating the further development of recovering communities has already been detected. It concerns the role of small bushy islands in, for instance, moderately-grazed pasturelands. These nuclei are protected from cattle herbivory by unpalatable species and spiny taxa, such as the dwarf-shrub Acaena elongata, the bracken fern Pteridium aquilinum, the blackberries Rubus spp., the composite Cirsium subcoriaceum and the large-leaved Solanum rudepannum. According to A.M. CLEEF (pers. obs.) these centres of recovery are many times situated near rock outcrops or large stones that provide an initially suitable habitat (microclimate). In the core of such 'refugia' a variety of flowering plant species is protected from cattle. The flowers and fruits of these taxa are supposed to attract numerous pollinators and seed dispersal agents, mainly insects and birds, which in their turn bring in new propagules originating from better-developed nearby forest patches. In this way, the unpalatable species mentioned above help establish new regeneration patches, which become richer in species with time. Similarly primary forest trees like Quercus spp., left over after clearing, may act as centres of recovery. The authors and collaborators observed, for instance, the occasional presence of squirrels, which distribute Quercus acorns from isolated oak trees over vegetation patches in different phases of recovery, thus stimulating the process of recovery. These forest remnants like the earlier-mentioned bushy islands - are also supposed to attract a variety of birds, which locally improve the seedbank by short-range seed dispersal (GEUZE 1989). This has been proved at least for secondary vegetation in tropical lowlands (GUEVARA et al. 1986), but is still to be studied systematically in disturbed tropical montane habitats.

The main objective of our study was to organize the montane secondary vegetation types in such a way that they can be recognized easily in the study area and other parts of the Costa Rican Cordillera de Talamanca. Due to limited time, however, not all types observed in the field could be studied.

Therefore we underline at this point the great need of future additional sampling of man-induced vegetation occurring in the area. Especially data collection at intermediate altitudes (2400-2700 m), as well as in communities structurally differing from the ones here presented (e. g. in tall shrub lands, bamboo and fern brakes, and roadside vegetation) will enable us to grasp the complete phytosociological hierarchy of the plant communities of montane secondary vegetation in the Costa Rican Cordillera de Talamanca.

482

M. Kappelle et al.

Acknowledgements. The authors are very much indebted to Antoine M. CLEEF (University of Amsterdam), who took the initiative to the present research and supported the authors during all stages of the study. Adelaida CHAVERRI (Universidad Nacional, Heredia) and Pablo SANCHEZ (Museo Nacional, San Jose) provided logistic support in Costa Rica (laboratory facilities, transport, herbarium). Margot SPREUWENBERG and Thorwald GEUZE helped with fieldwork and prepared initial reports. Collected plant specimens were identified by different specialists, among many others Luis D. G6MEZ, Luis J. PoVEDA and Nelson ZAMORA (CR). Antoine M . CLEEF, Otti WILMANNS and an anonymous reviewer con;tmented on earlier versions of the manuscript. The English text was corrected by Mary E. KAPPELLE-MOHR and the Spanish summary by Marta E. KAPPELLE-JUAREZ. This study was financially supported by the Hugo de Vries Foundation, the University of Amsterdam, the Universidad Nacional at Heredia (ECOMA project 762090) and grant W 84-331 of the Netherlands Foundation for the Advancement of Tropical Research (WOTRO-NWO).

References Barkman, J.J. (1990): Controversies and perspectives 111 plant ecology and vegetation science. - Phytocoenologia 18 (4): 563-589.

Barkman, J.J., Moravec, J. & Rauschert, S. (1986): Code of Phytosociological Nomenclature, 2nd ed. - Vegetatio 67: 145-195.

Beard, J.S. (1944 ): Climax vegetation in tropical America.- Ecology 25: 127-158.

Beard, J.S. (1955): The classification of tropical American vegetation types.- Ecology 36: 89-100.

Braun-Blanquet, J. (1965): Plant sociology. The study of plant communities. - Hafner, London, 439 pp.

Burger, W. (ed.) (1971-1990): Flora Costaricensis.- Fieldiana Bot. 35 and 40, and Bot. New Ser. 13 and 23. Chicago, U.S.A. Castillo, R. (1984): Geologia de Costa Rica. Una sinopsis.- Universidad de Costa Rica, San Jose de Costa Rica, 182 pp.

Chaverri, A. & Cleef, A.M. (in prep.): Zonal plant communities of the paramos of Costa Rica.

Cleef, A.M., Rangel, 0., Vander Hammen, T. & Jaramillo, R. (1984): The forest vegetation of the Buritaca transect.- In: Vander Hammen, T. & Ruiz, P.M. (eds.), La Sierra Nevada de Santa Marta (Colombia), the Buritaca-La C umbre transect. Stud. on Trop.

Andean Ecosyst. 2: 267--406.

Cuatrecasas, J. (1934): Observaciones geobotanicas en Colombia. - Trab. Mus. Nac. Cienc. Nat. Ser. Bot. 27: 1-144. Madrid.

Eiten, G. (1968): Vegetation forms. A classification of vegetation based on structure, growth form of the components, and vegetative periodicity. - Bol. Inst. Bot. 4, Sao Paulo. 33 pp.

Gauch, H. G. (1982): Multivariate analysis in community ecology.- Cambridge University Press, Cambridge. 298 pp.

Geuze, T. (1989): A silvicu ltural analysis of secondary succession in the montane oak forest belt, Cordillera de Talamanca, Costa Rica. - Thesis. Larenstein International School for Agriculture. Yelp (NL). 41 pp.

Guevara, S., Purata, S.E. & Vander Maarel, E. (1986): The role of remnant forest trees in tropical secondary succession. - Vegetatio 66: 77-84.

Hastenrath, S. (1973): On the Pleistocene glaciation of the Cordillera de Talamanca, Costa Rica.- Z. f. G letsc herk. u. G lazialgeol. 9 (1-2): 105-121.

Heaney, A. & Proctor, J. (1990): Preliminary studies on forest structure and floristics on Volcan Barva, Costa Rica.- J. Trop. Ecol. 6: 307-320.

Montane secondary vegetation in the Cordillera de Talamanca

483

Hill, M.O. (1979): TWINSPAN- A Fortran program for arranging multivariate data in an ordered two-way table by classification of individuals and attributes. - Cornell University Press, Ithaca, New York.

Instituto Meteorol6gico Nacional (1988): Catastro de las series de precipitaciones medidas en Costa Rica. - MIRENEM, San Jose de Costa Rica. 363 pp.

Jongman, R.H.G., Ter Braak, C.F.J. & Van Tongeren, O .F.R. (eds.) (1987): Data analysis in community and landscape ecology.- Pudoc, Wageningen. 299 pp.

Kappelle, M. (1991): Distribuci6n altitudinal de Ia vegetaci6n del Parque Nacional Chirrip6, Costa Rica.- Brenesia 36: 1-14.

(1992): Structural and floristic differences between wet Atlantic and moist Pacific montane Myrsine-Quercus forests in Costa Rica. - In: Balslev, H. & Luteyn, J.L. (eds.), Paramo: an Andean ecosystem under human influence: 61-70. Academic Press, London.

(1993 ): Recovery following clearing of an upper montane Quercus forest in Costa Rica.- Rev. Bioi. Tropic. (Costa Rica) 41 (1): 47-56.

Kappelle, M., Cleef, A.M. & Chaverri, A. (1989): Phytosociology of montane ChusqueaQuercus forests, Cordillera de Talamanca, Costa Rica.- Brenesia 32: 73-105.

Kappelle, M., Kennis, P.A.F. & De Vries, R.A.J. (in prep.): Changes in diversity along a successional gradient in a neotropical montane Quercus forest, Costa Rica.

Kappelle, M., Van Uffelen, J.-G. & Cleef, A.M. (in prep.): Altitudinal zonation of montane Quercus forests along two transects in the Chirrip6 National Park, Costa Rica.

Kappelle, M., Zamora, N. & Flores, T. (1991): Flora leiiosa de Ia zona alta (2000-3819 m) de Ia Cordillera de Talamanca, Costa Rica.- Brenesia 34: 121-144.

Lellinger, D.B. (1989): The ferns and fern-allies of Costa Rica, Panama and the Choc6 (Part 1: Psilotaceae through Dicksoniaceae).- A mer. Fern Soc. Nr. 2 A. 364 pp.

Monasterio, M., Sarmiento, G. & Solbrig, O.T. (eds.) (1987): Comparative studies on tropical mountain ecosystems. Planning for research. - Bioi. Int. Spec. issue 12. IUBS, Paris. 48 pp.

Mueller-Dombois , D. & Ellenberg, H. (1974): Aims and methods of vegetation ecology.Wiley, New York. 547 pp.

Reserva de Ia Bi6sfera La Amistad (1990): Estrategia para el desarrollo institucional de Ia Reserva de Ia Bi6sfera La Amistad. - MIRENEM, San Jose de Costa Rica. 174 pp.

Salamanca, S. (1991): Vegetation of the paramo and its dynamics in the volcanic massif Ruiz - Tolima (Cordillera Central, Colombia).- Ph.D. thesis, University of Amsterdam. 122 pp.

Spreuwenberg, M. (1989): Preliminar description of pastureland communities in the upper Rio Savegre watershed area, Cordillera de Talamanca, Costa Rica. - Int. report, Universidad Nacional, Heredia, Costa Rica. 43 pp.

Standley, P. (1937-1938): Flora of Costa Rica. - Field Mus. Nat. Hist. 18 (1-4). Pub!. 391.

Chicago. 1571 pp.

Stutz de Ortega, L. (1990): Etudes floristiques de divers stades secondaires des formations forestieres du Haut Parana (Paraguay oriental). Structure, composition floristique et recril forestier: analyse de cinq stades de succession secondaire.- Candollea 45: 81-123.

Sugden, A.M. (1983 ): Determinants of species composition in some isolated neotropical cloud forests. - In: Sutton, S.L., Whitmore, T.C. & Chadwick, A.C. (eds.), Tropical rain forests, ecology and management: 43-56. Blackwell, Oxford.

Tryon, R.M. & Tryon, A.F. (1982): Ferns and allied plants. With special reference to tropical America. - Springer, New York. 857 pp.

UNESCO (1973): International classification and mapping of vegetation. -Ecology and conservation 6. UNESCO, Paris. 92 pp.

Van der Hammen, T., Mueller-Dombois , D. & Little, M.A. (eds.) (1989): Manual of methods for mountain transect studies: comparative studies of tropical mountain ecosystems.- TME-IUBS and MAB-UNESCO, Amsterdam. 67 pp.

484

M. Kappelle et a!.

Van Gils, H.A.M.J. & Van Wijngaarden, W. (1984): Vegetation structure in reconnaissance and semi-detailed vegetation surveys.- ITC J. (3): 213-218.

Van Uffelen, J.G. (1991): A geological, geomorphological and soil transect study of the C hirrip6 massif and adjacent areas, Cordillera de Talamanca, Costa Rica. - Thesis, CATIE/MAG-Costa Rica and Wageningen Agric. Univ. 72 pp.

Van Velzen, H.P. (1992): Priorities for conservation of the biodiversity in the Colombian Andes.- Noved. Color!'lb. 4. Spec. issue. Mus. Hist. Nat. Univ. Cauca, Popayan, Colombia. 33 pp.

Van Velzen, H.P. & Wijtzes, W.H. (1990): Secondary vegetation in the upper Rio Savegre watershed area, Costa Rica. - Thesis.,University of Amsterdam. Int. rep. 277. 80 pp.

Van Velzen, H.P., Wijtzes, W.H. & Kappelle, M. (in press): Lista provisional de las plantas de Ia vegetacion secundaria del piso montano Pacifico de Ia Cordillera de Talamanca, Costa Rica. - Brenesia.

Vasquez, A. (1983): Soils.- In: Janzen, D.H. (ed.), Costa Rican Natural History: 63-65.

Univ. Chicago Press, Chicago.

Verweij, P.A. & Beukema, H. (1992). Aspects of human influence on upper Andean forest line vegetation.- In: Balslev, H. & Luteyn, J.L. (eds.), Paramo: an Andean ecosystem under human influence: 171-175. Academic Press, London.

Walter, H. (1985): Vegetation of the earth and ecological systems of the geo-biosphere.Springer, New York. 318 pp.

Westhoff, V. & Vander Maarel, E. (1973): The Braun-Blanquet approach of phyto-sociology.- In: Whittaker, R.H. (ed.), Handbook of vegetation science V: 619-726. Junk, The Hague.

Weyl, R. (1980): Geology of Central America.- Gebr. Borntraeger, Stuttgart. 371 pp.

Wolf, J.H.D. (1993a): Epiphyte communities of tropical montane rain forests in the northern Andes. I. Lower montane communities.- Phytocoenologia 22: 1-52.

- (1993b): Epiphyte communities of tropical montane rain forests in the northern Andes. II. Upper montane communities.- Phytocoenologia 22: 53-103.

Woodson Jr., R.E. & Schery, R.W. (eds.) (1943-1980): Flora of Panama.- Missouri Bot. Gard. St. Louis, U.S.A. Zonneveld, I.S. (1979): Land evaluation and land(scape) science.- ITC, Enschede. 134 pp.

Address of the authors: (Please address all correspondence to the first author.)

M. KAPPELLE, H.P. VAN VELZEN and W.H. WIJTZES, Hugo de Vries Laboratorium, Universiteit van Amsterdam, Kruislaan 318, NL-1 098 SM Amsterdam, The Netherlands.