MACROLICHENS OF MONTANE RAIN FORESTS IN PANAMA, PROVINCE CHIRIQUÍ

MACROLICHENS OF MONTANE RAIN FORESTS IN PANAMA, PROVINCE CHIRIQUÍ

Lichenologist 32(6): 539-551 (2000) doi: 10.1006/lich.2000.0295 Available online at http://www.idealibrary.com on IDEKL MACROLICHENS OF MONTANE RAIN ...

564KB Sizes 0 Downloads 22 Views

Lichenologist 32(6): 539-551 (2000) doi: 10.1006/lich.2000.0295 Available online at http://www.idealibrary.com on IDEKL

MACROLICHENS OF MONTANE RAIN FORESTS IN PANAMA, PROVINCE CHIRIQUI Burkhard BUDEL*, Angelika MEYERJ, Noris SALAZAR§, Hans ZELLNERJ, Gerhard ZOTZ$§ and Otto L. LANGEJ Abstract: Sixty-eight lichen species were found in the lower montane rain forest of Fortuna (1000-1200 m) in the province of Chiriqui, Panama; 33 of these were cyanobacterial lichens. The montane rain forest of the Volcan Barti (2200-3100 m) yielded 30 species of lichens, 13 of which have cyanobacterial photobionts. Forty of the species reported are new records for Panama. Information concerning their habitats within the forest and their phytogeographical affinities is provided where possible. (; 2000 The British Lichen Society

Introduction Neotropical evergreen forests are famous for their species diversity, although the occurrence and distribution of lichens in these forests have only been investigated in a few cases or regions (e.g. Sipman & Harris 1989; Lucking 1992; Aptroot 1997a; Biidel 1997). Aptroot & Sipman (1997) estimated that some 50% of the lichen flora is still unknown in the Neotropics. Reliable estimates of species numbers exist only for three geographical regions in the tropics, namely Papua New Guinea, The Guianas, and Columbia (Aptroot \991a,b; Aptroot & Sipman 1997; Sipman 1989; Sipman & Harris, 1989). Our knowledge of rain forest lichens from tropical Central America is poor, making it impossible to provide even a rough estimate of diversity. Frequently, it has been stated in the literature that the abundance of lichens in the tropics increases dramatically with altitude. In the cooler and wetter lower montane and montane forests, higher biomass and species numbers of lichens are attained (e.g. Seifriz 1924). However, in a study of Papua New Guinea by Aptroot (19976), the number of species of lichens found at all elevations (1-230 m asl; 2300-2750 m asl, and 3500-4500 m asl) was always between 400 and 500. While in the lowlands crustose growth forms dominated, foliose and fruticose species were much more abundant in the upper mountain regions (Sipman & Harris 1989). A similar picture emerged in a study in the Central Colombian Cordillera by Sipman (1989). There the author found 295 lichen taxa along a transect from 600 to 4600 m asl. Between 600 and 2000 m asl crustose growth forms dominated, whereas *Universitat Kaiserslautem, FB Biologie, P.O. Box 3049, D-67653 Kaiserslautern, Germany. flulius-von-Sachs-Institut fur Biowissenschaften, Julius-Maximilians-Universitat, Julius-vonSachs-Platz 3, D-97082 Wiirzburg, Germany. ^Smithsonian Tropical Research Institute, Box 2072, Balboa, Panama City, Panama. 0024-2829/00/060539+13 S35.00/0

'(:, 2000 The British Lichen Society

540

THE LICHENOLOGIST

Vol. 32

foliose lichens became dominant above 2000 m. Montfoort & Ek (1990) discovered 209 different lichen species in the canopy and the understorey of a lowland rain forest in French Guiana. Of these, 160 were crustose, eight leprose, 15 squamulose, 33 foliose (19 completely attached and 14 attached to the substratum with a specific part only), and two had a filamentous growth form. The observations of Sipman (1989), Sipman & Harris (1989), and Montfoort & Ek (1990) suggest that there is a characteristic distribution pattern of lichen growth forms along an altitude gradient in rain forest ecosystems. Two reports suggest that knowledge of lichen species richness in rain forest is generally poor. First, Aptroot (19876) listed 173 species on just one fallen tree in a montane forest of Papua New Guinea and, second, Lucking (1998) found as many as 80 foliicolous lichen species on a single branch of a palm tree in Costa Rica. Given such high numbers of species found in just two opportunistic studies, the real diversity can be expected to be significantly greater. However, further explorations are necessary in order to derive reliable estimates of lichen diversity in tropical evergreen forests. Ecophysiological field studies in September and October 1993 gave us the opportunity to collect lichens in the rain forest of Fortuna in the province of Chiriqui, Panama (the results of our ecophysiological work are published elsewhere: Lange et al. 1994; Zotz et al. 1996; Zotz et al. 1998; Lange et al. 2000). Collecting activities were confined to an area of about 1-5 km2 surrounding the main camp, where a surprisingly high number of macrolichens (in the sense of Swinscow & Krog 1988) was recorded. These results are reported here. Site description and lichen collection The collections were made at the 'Reserva Forestal Fortuna', an extensive area of dense, almost virgin, tropical 'lower montane rain forest' in the basin of the Chiriqui River, south of Lago Fortuna (Panama, Province of Chiriqui, north east of David, 8°45'N, 82°15'W; 10001400 m asl). The annual mean air temperature at Fortuna is between 21 and 22°C (Anonymous 1988). At the Centro de Investigaciones Tropicales Jorge L. Arauz of the Instituto de Recursos Hidraulicos y Electrificacion (IRHE), where the work was undertaken, annual rainfall was 3509 mm and 3113 mm for 1991 and 1992, respectively. Monthly precipitation was never less than 100 mm in the 3 years preceding our measurements. The forest at Fortuna has an open canopy. Trees are generally 20-30 m tall, but emergents reach up to 40 m. The physiognomic characteristics suggests that the forest is a lower montane (premontane) tropical rain forest in the sense of Grubb (1977). A brief description of the forest vegetation can be found in Cavelier (1992), and a preliminary species list of phanerogamic taxa is given in Adames (1977). No published data are available on the number of vascular epiphytes but a preliminary investigation by G. McPherson (pers. comm.) indicated that it exceeds 200. Additional collections were made in a montane rain forest at the Volcan Baru (22003100 m asl), situated about 25 km west of Fortuna. Here the trees are up to 20 m tall, and the understorey is less shady compared to the lower montane forest at Fortuna. Lichen samples were taken from the ground, from stems and branches of shrubs and trees in the lower part of the canopy, inside the forest, at clearings and along the forest margin. Canopy samples were taken from freshly fallen branches and by climbing into the canopy of a few trees. Sampling was confined to lichens of foliose or fruticose growth form. Herbarium specimens are deposited at the Smithsonian Tropical Research Institute and in the private herbarium of B. Biidel. Phytogeographical regions are named according to Galloway (1996) and Sipman (1989). The phytogeographical affinities of the lichens were determined using recentflorasor monographs

2000

Macrolichens of Panamanian rain forests—Budel et al.

541

(e.g. Swinscow & Krog 1988; Galloway 1991, 1994). New records for Panama were estimated on the basis of the catalogue of Imshaug (1956) and several more recent monographs.

Results Lower montane rain forest, Fortuna (1000-1400 m) The evergreen forest of Fortuna is rich in vascular and non-vascular epiphytes, and lichens are quite abundant at open sites such as clearings and along forest edges, where higher light exposure is usually experienced. In total 68 macrolichen species were found in this area (Table 1). Almost half of the species (33) were lichens with a cyanobacterium as primary photobiont (Table 1). The cyanobacterial lichens ( = cyanolichens) belong to the following families: Collemataceae (gelatinous lichens, Leptogium, Collema), Coccocarpiaceae (Coccocarpia), Lobariaceae {Lobaria, Pseudocyphellaria, Sticta), Pannariaceae {Erioderma, Pannana, Parmeliella), Peltigeraceae {Peltigera) and Thelephoraceae (Basidiolichens, Dictyonema). The green algal lichens ( = phycolichens) belong to the families Cladoniaceae {Cladonia, Cladia), Gyalectaceae {Coenogonium), Parmeliaceae {Hypotrachyna, Oropogon, Parmotrema, Usnea), Physciaceae {Heterodermia). Peltigeraceae {Peltigera), Lobariaceae {Lobaria, Pseudocyphellaria, Sticta), Sphaerophoraceae (Sphaerophorus), Stereocaulaceae {Stereocaulon) and Teloschistaceae {Teloschistes).

Seven species were found only inside the forest, whereas nine occurred inside the forest, in clearings and at the forest margins. Forest margins or clearings are the richest habitats for lichens; here, most of the species are epiphytes (Table 1). Five species, Cladonia peltastica, C. ceratophylla, Dictyonema sericeum, Sphaerophorus melanocarpus and Usnea sp. 3, occurred in the lower and upper canopy region (Table 1). Sphaerophorus melanocarpus was found only on larger branches from the upper canopy, while Cladonia ceratophylla also occurred on the ground in clearings. However, these observations have to be considered with reservation since access to the upper strata of the forest was rather limited. The biogeographical affinities of many of the lichen species (39%) found at Fortuna are to a large extent unknown, 39% have a pantropical to wide pantropical distribution, 7% are known only from the Neotropics and 6% have a neotropical and African distribution. About 4% of the species have a cosmopolitan distribution pattern (Fig. 1; Table 1). Montane rain forest, Volcan Baru (2200-3100 m) Collections were made between 2200 and 3100 m asl. Thirty species were found, of which 14 were cyanolichens belonging to the two genera Leptogium and Sticta (Table 1). The genus Leptogium and the green algal lichen genus Lobaria have the highest species richness with eight species per genus (Table 1). Contrary to the situation at Fortuna, most species were found inside the forest, where they grew either on twigs, stems of trees or shrubs, or were found on the ground growing on larger rocks and boulders. This forest, however, was less densely shaded than the forest at Fortuna. As at Fortuna,

On twigs

On stems

Collema sp. 2 (fertile)

Cryptotheeia sp.

On stems

Collema sp. (thick, thallus bluish when wet)

C. linkii Fhrenb. ex Nees

On twigs

1000 1200

1000 1200

1000-1200

Pantropical? 1000-1200

On stems On stems

C. pellita (Ach.) Mull. Arg. Coenogonimn interplexmn Nyl.*

1000-1200 1000 1200

1000-1200

Cyanobacterium

On stems and tree logs

C. palmicola (Spreng.) L. Arvidss. & D. J. Galloway

Cyanobacterium Green alga, Trentepohlia Green alga, Trentepohlia Cyanobacterium, Nostoe Cyanobacterium, Nostoe Green alga

1000-1200

Cyanobactenum

On stems

C. erythroxyli (Spreng.) Swinscow & Krog

Neotropical

Widespread in tropical and temperate regions Neotropical

Phytogeography

Neotropical, extensions to warm temperate regions and Fiji Worldwide, tropical to temperate regions Worldwide, tropical to warm temperate regions Pantropical Pan tropical? 1000-1200

Cyanobacterium

Coccocarpia domingensis Vain.*

Cladonia sp. 2

C. ramulosa (With.) J. R. Laund.* Cladonia sp. 1 1000-1200

2200-3100 1000-1200 1000-1200

1000 1200

Altitude (m asl)

Green alga

Green alga Green alga Green alga

Green alga

Primary photobiont

1000-1200 1000-1200

On soil

Forest margin or clearing

Green alga Green alga

On ground Twig in canopy On liana

Inside forest

On soil and decomposing log On rocks On ground and on rock On soil and rocks On leaves

Cladonia andesita Vain.* C. peltastica (Nyl.) Mull. Arg.* C. ceratophylla (Sw.) Spreng.*

Cladia aggregala (Sw.) Nyl.

Species

TABLE 1. Macrolichens recorded in Panamanian montane rain forests (lower montane rain forest,Fortuna, 1000-1200 m asl; montane rainforest, Vokan Barii, 2200-3100 m asl)

o

H

o

S

o

Z

M

X

I-H

r n

ffl

H X

stems stems stems rocks

H. rhabdiformis (Kurok.) Hale

H. osseoalba (Vain.) Y. S. Park & Hale

H. imbricatula (Zahlbr.) Hale

H. exsplcndens (Hale) Hale*

On trees

log

On decomposing

log

On decomposing

log

On decomposing

On stems

On decomposing

Hypotrachyna costaricensis (Nyl.) Hale log

On stems

On tree

log

On decomposing

On On On On

On stems and twigs, on ground On stems and twigs

Forest margin or clearing

H. leucomelos (I..) Poelt

On rocks

On lianas and twigs

D. scriceum (Sw.) Berk.*

Enodernia zvrightii Tuck.* Hcteroderniia barbifera (Nyl.) W. A. Weber H. cf. casarettiana (A. Massal.) Trevis. H. flabellata (Fee) D. D. Awasthi* H. isidiophora (Vain.) D. D. Awasthi H. japonica (Sato) Swinscow & Krog*

On twigs

Inside forest

Dichosporidium nigrocinctum (Ehrenb. Fr.) Thor* Dictyonema glabratum (Spreng.) D. Hawksw.

Species

TABLE 1. Continued

Green alga

Green alga

Green alga

2200-3100

1000-1200

1000 1200

1000-1200

1000-1200

Green alga Green alga

1000-1200 2200-3100

1000-1200 1000-1200 1000-1200 1000 1200 2200-3100 1000-1200

1000-1200

1000-1200 1000-1200

Altitude (m asl)

Green alga

Cyanobacterium Green alga Green alga Green alga Green alga Green alga

Scytonema

Green alga Cyanobacterium, Scytonema Cyanobacterium,

Primary photobiont

Pantropical

Pantropical

Pantropical

Neotropical and Africa

Wide pantropical, extending into northern and southern temperate zones Neotropical Pantropical Neotropical and Africa Pantropical Pantropical Africa, Asia, New Zealand Wide pantropical, extending into northern and southern temperate zones Pantropical

Neotropical and Africa

Phytogeography

I

to

5' s

3.

SO

3

~

n

03

o o o

bo

On stems and branches On trees and rocks

L. cyanescens (Rabenh.) Korber*

On rocks

Leptogium sp. 3

Leptogium vesiculosum (Sw.) Malme*

On trees

Leptogium sp. 2

Leptogium sp. 1

On stems twigs

L. phyllocarpum (Pers.) Nyl.

On twigs

decomposing log On stems

and

On rocks

On stems

On stems and branches

On stems

Forest margin or clearing

1. Continued

L. marginellum (Sw.) Gray*

L. cf. mandonii P. M. Jorg. (pale form)*

L. laceroides de Lesd.*

L. foveolatum Nyl.*

On trees

On trees

L. coralloideum (Meyen & Flot.) Vain.*

L. burgessii (L.) Mont.*

On stems and twigs On rocks

Inside forest

Leptogium azureum (Sw. ex Ach.) Mont.*

Species

TABLE

Cyanobacterium, Nostoc Cyanobacterium, Nostoc Cyanobacterium, Nostoc Cyanobacterium, Nostoc

Cyanobacterium, Nostoc

Cyanobacterium, Nostoc Cyanobacterium, Nostoc Cyanobacterium, Nostoc Cyanobacterium, Nostoc Cyanobacterium, Nostoc Cyanobacterium, Nostoc Cyanobacterium, Nostoc Cyanobacterium, Nostoc

Primary photobiont

Cosmopolitan

1000-1200

1000-1200

2200-3100

2200-3100

1000 1200

Tropical America, West Indies

Africa, tropical America, West Indies, Philippines, Australia Widespread in the tropics and subtropics

1000-1200

1000 1200 2200-3100

Africa, the Americas, New Zealand Neotropical

1000-1200 2200-3100 2200-3100

2200-3100

2200-3100

2200-3100

Widespread in tropical and temperate regions Wide pantropical and temperate Pantropical

Phytogeography

1000-1200

Altitude (m asl)

<<.

H

or o0

r n X w

X a

H

544 r

u> to

ol.

On trees On stems On On On On

L. deplanata (Nyl.) Yoshim.* L. dissecta (Sw.) Raeuschel*

L. pallida (Hook, in Kunth) Trevis L. peltigera (Delise) Vain, s.lat. L. pulmonaria (L.) Hoffm.* L. ravenelii (Tuck.) Yoshim.* Lobaria sp. 1 Lobaria sp. 2 Megalospora sp. Oropogon loxensis (Fee) Th. Fr. Oropogon sp. Pannaria rubigmosa (Ach.) Bory

Green alga Cyanobacterium

On rocks On rocks and soil On stems

Peltigera laciniata (Merr. ex Riddle) Gyeln.*

Penusaria sp.

Green alga

Green alga Green alga

P. tinctorum (Nyl.) Hale

Parmotrema cf. abnuens (Nyl.) Hale P. mellissii (Dodge) Hale

Cyanobacterium

Green alga Green alga Green alga Green alga Green alga Green alga Green alga Green alga Green alga Cyanobacterium

Green alga Green alga

Green alga

Primary photobiont

On stems and decomposing log On twigs On decomposing log

On stems

On stems

On stems

On stems

Forest margin or clearing

Parmeliella pannosa (Sw.) Mull. Arg.*

On trees On trees

On trees

trees trees trees shrub

On stems

Inside forest

Lobaria crenulata (Hook, in Kunth) Trevis

Species

TABLE 1. Continued

1000-1200

1000-1200

1000-1200

1000-1200 1000-1200

1000-1200

1000-1200 2200 3100 2200-3100 1000-1200 2200-3100 2200-3100 2200-3100 2200-3100 2200-3100 1000-1200 2200-3100 1000 1200 2200-3100 2200-3100 1000-1200

Altitude (m asl)

Widespread in tropical and warm temperate regions Widespread in tropical and temperate regions

Worldwide from cool temperate regions to cool regions of the tropics Widespread in the tropics and subtropics

Neotropical

Cosmopolitan Probably neotropical

Neotropical

Tropical America, New Zealand, Hawaii Neotropical and Pacific Tropical America

Phytogeography

to S:

3

o

05

3

05

tr

n

os n

to o o o

2200 3100 1000-1200 2200-3100 Green alga Cyanobacterium,

On stems On stems On stems On stems

On trees

5. ferax Mull. Arg.* A', fuliginosa (Hoffm.) Ach.*

.S\ kutithu Hook.*

S. macrophylla Delise

Sticta sp. 1

Nostoc

Cyanobacterium,

Nostoc

Cyanobactcrium,

Nosloc

Cyanobacterium,

Nostoc

1000 1200

1000-1200

1000-1200

2200-3100

Green alga

On trees

2200-3100

5. dicholoma Delise*

Nostoc

Cyanobactcrium,

1000 1200

1000-1200 1000-1200

1000-1200

1000-1200

1000-1200

Altitude (m asl)

On trees

Green alga

Green alga Green alga

Nostoc

Cyanobacterium,

Nostoc

Cyanobactcrium

Green alga

Primary photobiont

Sticla ambavillaria (Bory) Ach.*

Stereocaulon virgatum Ach.*

On ground

On stems

Relicina abstrusa (Vain.) Hale Sphaerophorus melanocarpus (Sw.) DC. 1

On big branch in canopy

On stems

P. intricata (Delise) Vain.*

Pseudocyphellaria sp.

Forest margin or clearing On stems, branches and decomposing log On stems

Inside forest

Pseudocyphellaria aurata (Ach.) Vain.*

Species

TABLE 1. Continued

Probably pantropical (Mauritius, Comoros, South Africa, Panama)

Wide pantropical, extending to temperate regions

Probably pantropical (Mascarenes, E-Africa, Panama)

Pantropical Wide pantropical and temperate Neotropical? (West Indies, Central America) Pantropical

Pantropical, extending to temperate regions

Pantropical, extending to temperate regions

Phytogcography

< o

H

00

o r o0

n X w

I HH /•*»

M

w

H X

*New record for Panama.

Teloschistesflavicans(Sw.) Norman* Usnea rubicunda Stirt. Usnea sp. 1 Usnea sp. 2 Usnea sp. 3 (yellowish-green) (7s«ea sp. 4 (pale green) Usnea sp. 5 (fertile) On big branch

On trees

On stems On stems

On stem

On stems On stems

On stems, branches and decomposing log On stems

S. weigelii (Ach.) Vain.

S. weigelii var. xanthotropa (Krempelh.) Hue

On stems

*S\ tomentosa aggr.

On stems

On decomposing log

S. tomentosa (Sw.) Ach. On stems and decomposing log

On stems

Sticta sublimbata (J. Steiner) Swinscow & Krog*

On trees

Sticta sp. 6

On stems

Sticta sp. 4 On trees

On stems

Siicta sp. 3

Sticta sp. 5

On leaves

Inside forest

Forest margin or clearing

li 1. Continued

Sticta sp. 2

Species

TAHI

Cyanobacterium, Nostoc Green alga Green alga Green alga Green alga Green alga Green alga Green alga

Wide pantropical

1000-1200 2200-3100 1000-1200 1000-1200 2200-3100 1000-1200 1000-1200 1000-1200 1000-1200

Wide pantropical Cosmopolitan

Wide pantropical

1000-1200

Pantropical

Pantropical

1000-1200 1000-1200

Palacotropical

EL

r-t

S:

ba

r-t

ft

o1

3

5

3 B

3

2200-3100 1000-1200

o

2200-3100

1000-1200

o

1000-1200

n

JO

1000-1200

Cyanobacterium, Nostoc Cyanobacterium, Nostoc Cyanobacterium, Nostoc Cyanobacterium, Nostoc Cyanobacterium, Nostoc Cyanobacterium, Nostoc Cyanobacterium, Nostoc Cyanobacterium, Nosloc Cyanobacterium, Nostoc

Phytogeography

Altitude (m asl)

Primary photobiont

<_>

200( 547

THE LICHENOLOGIST

548

Vol. 32

D Lower montane • Montane

35% 30% 25% -

1l.fl.n 'a.

Z

n



C

CO

1

"o

ate

'o.

J,...

opica] frican opical acific

S.

eotro cal

mopo tan

n%

antro cal

5%

antro cal

10% —

CO

c

'a.

CO

P.

Jncer stribi ion



11 Ij

15% -

eotro cal

i

20% -

3

PH

FIG. 1. Phytogeography of the lichens of the lower montane and montane rain forest.

many of the species (35%) have uncertain distributions, 34% of the species showed a pantropical to wide pantropical distribution and 27% of the species belong to the neotropical distribution group (Fig. 1). Only seven lichen species, Heterodermia leucomelos, Leptogium laceroides, L. phyllocarpum, Lobaria crenulata, L. dissecta, Sticta fuliginosa, and S. weigelii var. xanthotropa occurred

in both forest types considered in this study. A total of 40 lichens species are new records for Panama.

Discussion Diversity Although the number of vascular epiphytes exceeds 200 species around Fortuna (G. McPherson, pers. comm.), the total number of macrolichens found in the two forest types (68 species in the lower montane and 30 in the montane rain forest, with seven species found in both forests) is surprisingly high, despite the fact that the canopy layer could not be sampled intensively. The relatively high number of about 40 species recorded here for the first time for Panama, reflects the insufficient state of lichenological investigation, rather than an unusual species richness of lichens in Panama when compared with the neighbouring countries. The large number of foliose and fruticose lichens in the lower montane forest of Fortuna corresponds well with the findings of Aptroot & Sipman (1997), who observed an increase in macrolichens in montane rain forests at an altitude as low as 1000 m. In both studies, most of the macrolichens in

2000

Macrolichens of Panamanian rain forests—Budel et al.

549

the montane and lower montane forest types belong to the lichen order Lecanorales. Phytogeographical affinities The high percentage (39%) of pantropical species found in the lower montane rain forest of Fortuna, agrees well with the values recorded, for example, by Sipman & Harris (1989), who found that pantropical species (e.g. Cladia aggregata, Heterodermia leucomelos, Sphaerophorus melanocarpus) may

constitute about one third of all lichen species in neotropical rain forests. Only 14% in our study were neotropical species (e.g. Cladonia ceratophylla, Lobaria dissecta), some showing a neotropical and African distribution (e.g. Dictyonema glabratum, Heterodermia casarettiana). In a Colombian montane rain forest, species with a wide tropical distribution were prevalent below 2000 m and species with a restricted neotropical distribution were most frequent between 2000 and 3500 m (Sipman 1989). To some extent, our results reveal a similar pattern, with a higher proportion of neotropical species in the montane forest between 2200 and 3100 m. A comparison of the lichen flora of the Fortuna region (1000-2000 m) with that of the Colombian montane forests investigated by Sipman (1989) revealed close affinities of the former with that of the Colombian montane forest between 2000 and 4000 m (29 species in common), but less so with the lower montane zone at 1300 m (six species in common). Sixteen species of the lower montane forest at Fortuna and 17 species of the montane forest at Volcan Baru were also found in a montane forest in Costa Rica, by Kappelle and Sipman (1992), who found 66 foliose and 28 fruticose lichens species there. Cyanolichen ratio In both forest types that we investigated, almost half of the species found were cyanolichens, often with the genus Nostoc as the primary photobiont. Most of them grew in shaded habitats (e.g. Leptogium), whereas others (Sticta and Dictyonema) also occurred in sunny, exposed habitats. Only three genera with cephalodia, Lobaria, Pseudocyphellaria and Stereocaulon, were found

during this survey. Our results confirm the observations of Forman (1975) who reported an increase in cyanolichen species richness and biomass in the more light-sheltered and humid parts of a Colombian rain forest at 2700 m. A study of rain forests at different elevations in Papua New Guinea (Aptroot 19976) revealed five cyanolichen species in comparison to 495 phycolichen species in the lowland forest zone (i.e. ~ 1%), 40 cyanolichens in comparison to 360 phycolichens in the lower montane forest zone (i.e. ~ 10%), and 70 cyanolichens in comparison to 380 phycolichens in the montane forest zone (i.e. ~16%). When only foliose and fruticose lichens are considered, this trend is even more pronounced, with cyanolichens comprising 12-5% of species richness in the lowland rainforest, 31 % in the lower montane zone, and 33% in the montane zone (Aptroot 19976). In our study, these values were even higher, with 48-5% cyanolichens in the lower montane rain forest and 46-6% in the montane forest. Our data, however, are preliminary and more intensive collecting may well alter these provisional ratios.

550

THE LICHENOLOGIST

Vol. 32

We acknowledge the support of P. M. Jorgensen (Bergen), H. Sipman (Berlin) and M. Schultz (Germany). The work was supported by the Deutsche Forschungsgemeinschaft within the 'Sonderforschungsbereich 251 der Universitat Wiirzburg'. The Panamanian 'Institute Nacional de Recursos Naturales Renovables' granted permission for the study in the Reserva Forestal Fortuna. We are grateful to the 'Instituto de Recursos Hidraulicos y Electrificacion (IRHE)', especially to I. J. Ortega and I. J. F. Victoria, for the opportunity to make use of the 'Centra de Investigaciones Tropicales Jorge L. Arauz'. REFERENCES

Adames, A. J. (1977) Evaluacion ambiental y efectos del Proyecto Hidroelectrico Fortuna. Revista Loteria 254-255-256: 1-538. Anonymous (1988) Atlas Nacional de la Republica de Panama. Panama: Instituto Geografico Nacionale, 'Tommy Guardia'. Aptroot, A. (1997a) Species diversity in tropical rainforest ascomycetes: licheniscd versus non-lichenised; foliicolous versus corticolous. Abstracta Botanka 21: 37-44. Aptroot, A. (19976) Lichen biodiversity in Papua New Guinea, with report of 173 species on one tree. Bibliotheca Lichenologica 68: 203-213. Aptroot, A. & Sipman, H. J. M. (1997) Diversity of lichenised fungi in the tropics. In Biodiversity of Tropical Microfungi (K. D. Hyde, ed.): 93-106. Hong Kong: University Press. Biidel, B. (1997) Flechten der Tropen. In Flechten—Geschichte, Biologic, Systematik, Okologie, Naturschutz und kulturelle Bedeutung (H. Scholler, ed.): 139-148. Kleine Senckenbergreihe Nr. 27. Cavelier, J. (1992) Fine root biomass and soil properties in a semi-deciduous and a lower montane rain forest in Panama. Plant and Soil 142: 187-201. Forman, R. T. T. (1975) Canopy lichens with blue-green algae: a nitrogen source in a Colombian rain forest. Ecology 56: 1176-1184. Galloway, D. J. (1991) Phytogeography of southern hemisphere lichens. In Quantitative Approaches to Phytogeography (P. L. Nimis & T. J. Crovello, eds): 233-262. The Netherlands: Kluwer. Galloway, D. J. (1994) Studies on the lichen genus Sticta (Schreber) Ach.: I. Southern South American species. Lichenologist 26: 223-282. Galloway, D. J. (1996) Lichen biogeography. In Lichen Biology (T. H. Nash III, ed.): 199-216. Cambridge: Cambridge University Press. Grubb, P. J. (1977).Control of forest growth and distribution on wet tropical mountains: with special reference to mineral nutrition. Annual Reviezvs of Ecology and Systematics 8: 83-107. Imshaug, H. A. (1956) Catalogue of Central American lichens. Bryologist 59: 69-114. Kappelle, M. & Sipman, H. J. M. (1992) Foliose and fruticose lichens of Talamanca montane Quercus forests, Costa Rica. Brenesia 37: 51-58. Lange, O. L., Biidel, B., Zellner, H., Zotz, G. & Meyer, A. (1994) Field measurements of water relations and C O , exchange of the tropical, cyanobacterial basidiolichen Dictyonema glabratum in a Panamanian rainforest. Botamca Acta 107: 279-290. Lange, O. L., Biidel, B., Meyer, A., Zellner, H. & Zotz, G. (2000) Lichen carbon gain under tropical conditions: water relations and CO 2 exchange of three Leptogium species in a Panamanian rainforest. Flora 195: 172-190. Lucking, R. (1992) Zur Verbreitungsokologie foliikoler Flechten in Costa Rica, Zentralamerika. Nova Hedidgia 54: 309-353. Lucking, R. (1998) Ecology of foliicolous lichens at the 'Botarrama' trail (Costa Rica), a neotropical forest side. Part II. Ecotropica 4: 1-24. Montfoort, D. & Ek, R. C. (1990) Vertical distribution and ecology of epiphytic bryophytes and lichens in a loivland rainforest in French Guiana. M.Sc. Thesis, University of Utrecht. Seifriz, W. (1924) The altitudinal distribution of lichens and mosses on Mt Gedeh, Java. Journal of Ecology 12: 307-313. Sipman, H. J. M. (1989) Lichen zonation in the Parque Los Nevados transect. Studies on tropical Andean ecosystems 3: 461-483. Sipman, H. J. M. & Harris, R. C. (1989) Lichens. In Tropical Rain Forest Ecosystems (H. Leith & M. J. A. Werger, eds): 303-309. Amsterdam: Elsevier. Swinscow, T. D. V. & Krog, H. (1988) Macrolichens of East Africa. London: British Museum (Natural History).

2000

Macrolichens of Panamanian rain forests—Budel et al.

551

Zotz, G., Budel, B., Meyer, A., Zellner, H. & Lange, O. L. (1996) Water relations and CO 2 exchange of tropical bryophytes in lower montane rain forest in Panama. Botanica Ada 110: 9-17. Zotz, G., Budel, B., Meyer, A., Zellner, H. & Lange, O. L. (1998) In situ studies of water relations and CO 2 exchange of the tropical macrolichen, Sticta tomentosa. New Phytologist 139: 525-535~ Accepted for publication 9 June 2000