Epiphyte communities of Tropical montane rain forests in N Andes II - Upper communities - Wolf 1993

.

53- 103

Phyto coeno logia

Berli n-Stu ttgart , June 30, 1993

Epi phy te com mu niti es of trop ical mo nta ne rain forests in the nor the rn Andes II. Up per mo nta ne com mu niti es

by Jan H. D. WoLF, Ams terda m with 10 figures and 37 tables

Intr odu ctio n communities from tropiIn a series of two papers epiphytic shade and canopy ibed for the first time, cal mon tane rain forests in the north ern Andes are descr (WEsTHOFF & VAN y iolog using the Braun-Blanquet approach in phytosoc altitudinal transect ranDER MAAREL (1973). The stud y was carried out in an Cent ral Cordillera in the ging from 1000 m to 4130 m on the west slope of N 75° 30 W). A descripColo mbia , near Pereira, capital of Risaralda (ca. 4°50 of sampling used and an tion of the stud y area, the forest climate, the methods forest may be found in account of the communities of Lower Mon tane Rain the first pape r (WoLF 1993). ted in 20 communiA TWI NS PAN analysis (HILL 1979) of 528 releves resul ps. Two additional grou main ties, clust ered into four, altitudinally separated, e in any of the denc confi species poor communities could not be placed with four groups. will be described folIn this pape r the communities from higher altitudes yte communities in the lowed by a summary and a general discussion of epiph altitu dina l trans ect.

Res ults t, 2460 -367 0 m 3. Com mun ities of the Upp er Mon tane Rain fores (Table 1) Upp er Montane Rai? Exclusive taxa for the epiphyte communities of the B. hookeri, Lepicolea prutforest are Adel anth us pittieri, Bazz ania breuteliana, ochila bursata, Riccardia Plagi a, nosa, Lepi dozia spp., Lophocolea trapezoide ntosa. spp., Sema toph yllum insul arum and Trichocolea tome t are clustered into The 264 releves made in the Upp er Mon tane Rain fores es. five tree base communities and five canopy communiti 0340-269X/93/0022-0053 $ 12.75 n· D-70176 Stutt gart © 1993 Gebri ider Borntraeger, D-14129 Berli

subito e.V. licensed customer copy supplied for Universidad Ceu San Pablo (SLI11X00058E)

l ]. H. D. Wolf

54

Table 1. Synoptic table of the epiphyte communities in the Upper Montane Rain forest. In addition to the presence degrees of species, their minimal cover value, characteristic cover COMMUNITY 3

NUMBER OF RELEVES IN THE COMMUNITY MEAN RELEVE AREA (dm2) TOTAL NUMBER OF TAXA IN THE COMMUNITY

3 1 1 2 32 6.2 133

1 1 1 25 7.2 107

3 1 2 61 7.3 226

pres cover(%) pre& cover(%) pres cover ~•)

a. !;;QmmuDiti~ gf l~ 1.!12121:[ Ms!Dlill!il BiliD wwl Trichocolea tomentosa (Sw.) Gott.

Lepicolea pruinosa (Tayl.) Spruce Lepidozia spp.

Lophocolea trapezoidea Montagne Bazzania hookeri (Lindenb.) Trevis.

Adelanthus pittieri (Step h.) Grolle Campylopus asperifolius Mitt.

Riccardia spp.

Plagiochila bursata (Desv.) Lindenb.

Sematophyllum insularum (Sull.) Mitt. !:iUlQ~)l ~2mmuoili~ 2f tb!: u~~!:[ MsmliiD!: RiliD f2wt Jamesoniella rubricaulis (Nees) Grolle s.lat.

Prionodon fusco-lutescens Hampe Trachylejeunea dominicencis Step h.

Frullania convoluta Lindenb. & Hampe Leptoscyphus jackii (Steph.) Grolle Plagiochila echinella Gott. group Dictyonema glabratum (Spreng.) D. Hawksw. (=Cora pavonia Fr.) Hypotrachyna prolongata (Kurok.) Hale Hypotrachyna physcioides (Nyl.) Hale Leptoscyphus porphyrius (Nees) Grolle Hypotrachyna bogotensis (Vain.) Hale Bazzania longistipula (Lindenb.) Trevis.

Drepanolejeunea spp.

Hypotrachyna laevigata ((Smith) Hale)- group Anoplolejeunea conferta (Meissn.) Step h.

Daltonia longifolia Tayl.

Cheilolejeunea spp.

Diogenesia cf. tetranda (A.C. Smith) Sleumer Hypotrachyna imbricatula (Zahlbr.) Hale Hypotrachyna caraccensis (Tayl.) Hale Frullania sphaerocephala Spruce Herbertus pensilis (Tayl.) Spruce Hypotrachyna reducens (Nyl.) Hale Grammitis apiculata (Kl.) Seymour Herbertus juniperoideus (Sw.) Grolle Hymenophyllum sp. A (# 750) Squamidium leucotrichum (Tayl.) Broth.

Herbertus subdentatus (Steph.) Fulf.

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44 <1 11 40 3 19 <I 5 20 44 34 <1 <1 2 16 44 <1 3 10 13 47 <1 13 70 7 91 I 19 55 77 16 <I 5 20 13 6 I 6 10 3 69 <I 13 70 7 69 <1 2 15 26 38 <1 6 30 2 13 <I 9 30 8 56 <1 <I I 49 59 <I 4 32 18 56 <I <I 5 49 3 I I I 15 19 <I 1 I 25 3 2 2 2 3 28 <I 4 15 11 34 <1 8 30 3 <1 <1 <1 7 34 <I 2 5 13 9 <1 2 5 3 6 <1 <1 <1 7 9 <1 <1 <1 16 16 <I 4 10 3 13 <1 <1 2 20 41 <1 7 62

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55

Epiphyte communities of tropical montane rain forests. II

value(= average cover value when present) and maximum cover value are given. The'#' refer to collection numbers.

3 1

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P""' cover (%} P""' cover (%)pres cover (%)pres cover(%) pres cover(%) pres cover (%)pres cover(%) pres cover (%) pres cover(%) o/o .. av + o/o .. av + o/o - av + o/o - av + 'Yo ¡ av + 'Yo .. av + o/0 .. av + o/0 .. av + o/0 .. av + 17 <1 < 74 1 <I 66 <I 6 66 <1

<1 2 30 <1 7 30 91 <I 3 25 43 <I <I <I 11 <I 13 60 66 <1

1 13 9 <I 12

6 98 80 I

27 18 73 18 90 9

<1 <1 65 <I <I <I 60 I 40 85 75 <I <I <I 30 10 10 10 60

<1 10 45 14 <1 <1 <1 78

22 65 80 20 60 43 <I I 2 20 5 14 <I <I <I 44 <I 5 15 20 45 11 5 5 5 43 <1 15 99 34 <1 7 85 64 <1 11 35 25 <1 <1 2 57 2 5 2 2 9 3 3 3 S <1 <1 <I 14 2 2 <1 1 5 13 <I <1 <I 9 1 I I 10 <1 3 5 44 <1 6 10 ~ <1 12 40 9 5 5 5 5 1 1 1 14 90 90 90 11 <I <1 <I 20 9 <1 <1 <1 20 <1 <1 <1 11 30 30 30 11 <1 <1 1 9 <1 <1 <I <I <I <1

2 16 I 18

20

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6 17 30 30 5 5 <1 <I

2 2 2

60 <1 16 65 13 <I <I 1 4

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17 <I 5 15 2 I I I 15 <1 5 20 23 <1 10 35 <1 12 55 9 89 <I 18 75

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17 <1 <1 1 6 <1 2 5

10 <1 <1 <1

5 1 1 1

<1 <I <1 1 1 1

23 <1 1 2

6 5 13 20 2 <1 <1 <1

15 <1 9 40 6 <1 <1 2 96 <1 4 30 n <1 1 1 15 <1 3 20 17 <1 <1 2

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7

2 2 2

10 <1 <1 1

4 <1 <I <I 2 <I <I <I

14 <1 <1 <1

43 <1 <1 2 9 <1 <1 <I 34 <1 4 40 4 <1 <1 <1 45 <1 1 15 11 <1 <1 <1 9 <1 <1 <1

47 <1 <I

6 <1 <I 1 2 <1 <1 <1 15 <1 <1 <1 6 <1 <1 <I 6 1 2 17 3 5

60 <1 <1

19 <1 2 10 4 2 4 5

4

7

<1 2 2

2 38 <1 22 60 26 <1 10 85 9 <I <I <1 2

5 5 5

11 <1 <I 2

6 1 I 2 19 <1 11 55 9 <1 10 35 47 <I 4 20 64 <1 3 15

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J.H.D. Wolf

56 Table 1. (cont.)

COMMUNITY 3 1 1 1 Macromitrium trachypodium Mitt.

Amphilejeunea viridissima Schust.

Harpalejeunea ancistrodes (Spruce) Schiffn.

Anzia leucobates (Nyl.) Milll. Arg. Cladonia cf. ochrochlora Flk. Heterodermia vulgaris (Vain.) Follm. & Redan Frullania apiculata (Reinw. et al.) Nees Zygodon reinwardtii (Homsch.) Braun Epidendrum sp. B (#609) Sticta cf. damaecornis (Sw.) Ach. Elaphoglossum aff. lindigii (I<arst.) Moore Grammitis meridensis (1<1.) Seymour Everniastrum fragile Sipm.

Epidendrum spp. elleantoides group Elaphoglossum sp. A (# 826) Herbertus acanthelius Spruce Hypotrachyna microblasta (Vain.) Hale Lepyrodon tomentosus (Hook.) Mitt.

Hypotrachyna gigas (Kurok.) Hale Everniastrum cirrhatum (Fr.) Hale Pachyphyllum sp. A (# 751) Zygodon squarrosus (Tayl.) C. Mull.

Menegazzia sp. A(# 993) Leptotheca boliviana Herz.

Syzygiella integerrima Steph.

Blepharolejeunea incongrua (Lindenb. & Gott.) v. Slag. & Kruijt Anastrophyllum auritum (Lehm.) Step h.

Dictyonema spp.

Anastrophyllum leucocephalum (Tayl.) Step h.

Hastifera sp.

Brachiolejeunea laxifolia (Tayl.) Schiffn.

Lepidopilum haplociliatum (C.M.) Par (incl. L. muelleri) Aureolejeunea fulva Schust.

Adelanthus crossii Spruce Ceratolejeunea grandiloba Jack. & Steph.

Plagiochila sp. B (# 1354) Elaphoglossum sp. A (# 771) Sematophyllum cuspidiferum Mitt.

Leptodontium viticulosoides var. panamense (Lor.) Zander Cyrtolejeunea holostipa (Spruce) Evans ~.l.l, Ql.at!:E s;ano~, triigl!:l,

3 1 1 2 3 <1<1<1 1 1 1 3 <1 <1 <1

4 12 <1 <1 1 28 16 <1 1 3 13 13 12 <I <1 <1 3 4 <I <1 <I 3 4 <I <1 <I 6 3 13 3

3 1 2 2 2 2 2

28 <1 <1 <I <1 <1 2 1 1 <I <1 <I 3 <1 <I <1 <I <1 2 <I <I <I <I <1 <1 16 <1 <1 2 <I <I <1 33 <1 1 5 <I <I <I 18 <1 <I 2

2 2 2 2 2 2 2 5 10 I 1

8 <I I 2 8 1 1 I

6 <I 5 10 6 <I I 2 11 <I 3 10 2 1 I I

12 <I 4 10 16 <I 2 3 3 1 I 1

I 7 15 2 <I <I <I

7

22 <1 3 10

12 1 2 2 9 <1 <1 <1 13 <I 4 10 9 <1 <I <1

20 1 2 5 16 <I 2 5 3 <I <I 5 5 5 2 <I <I <I 10 30 6 <I <1 <1 15 <I <I 25 <I <1 2 <1 <1 <1 6 <I <I I 2 <I <I 9 <1 <I 2 <1 <1 <1 9 <I <I I <1 <I <1 11 <I <I 3

12 4 4 8

<I <I I <I

2 8 <1 <I <I

12 <1 <I

<1

7 <I <I <I

3 <I <I <I

8 3 4 5

15 <I 3 15 6 <I <I 1 33 <II 5 3 <I <I <1 2 I I 3 <1 <I <I 20 <I <I

S:smlml.lllilillll

Oropogon bicolor Essl.

Oropogon loxensis (Fee) Th. Fr. Aureolejeunea paramicola (Herz.) Schust.

Everniastrum catawbiense (Degel.) Hale

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57

Epiphyte communities of tropical montane rain forests. II

3 1 3 2 2 4 4 23 6 2 6 4 4 6 4 4 9 11

3 2 1 1

3 1 4 <1 <1 <1 <1

<1 <1 <1 <1

<1 2 <1 <1 <1

19 <1 6 10 21 <1 3 10 13 <1<1 2

3 2 2 1

3 2 2 2

4 1

4 2

15 <1 <1 <1

5 <1 <1 <1

15 <1 12 30

13 <1 4 5

26 1 5 10 11 <1 5 18 6 <1 <1 2 6 <1 <1 2 2 1 1 6 <1 2 5 15 <1<11

2 19 2 2 6

3 2 1 3

5 5 5

<1 <1 1 6 <1 <1 1 <1 <1 <1 <1 <1 <1 <1 <1 <1 1 1 1 10 15 20 1 2 3 5 2 4 5 2 3 <1 5 10

3 2 1 2

10 2 4 5

23 8 46 80

5 5 5 <1 3 12 13 <1 4 10 1 13 <1 29 82 <1 <1 <1 <1 47 95 2

2 2 2

4 <1 <1 15 <1 5 10

13 <1 <1 1 6 <1 <1 <1 2 <1 <1 <1 2 <1 <1 <1 2 <1 <1 <1 6 <1 <1 <1 2 2 6 2

<1 <1 <1 <1 <1 <1 2 <1 <1 <1 <1 <1 <1 <1 <1 <1 2 <1 <1 <1

11 1 1 1

11 1 20 35 2 <1 <1 <1 6 1 4 10 4 <1 <1 <1 4 <1 <1 1

51 19 21 4

<1 <1 <1 <1

<1 <1 2 <1

5 <1 10 <1

17 6 13 2

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subito e.V. licensed customer copy supplied for Universidad Ceu San Pablo (SLI11X00058E)

93 <1 5 20 31 <1 <1 1 60 <1 2 5

J.H.D. Wolf

58 Table 1. (cont.)

COMMUNITY 3

Drepanolejeunea andina Herzog Frullania peruviana Gott.

Anastrophyllwn tubulosum (Nees) Grolle Plagiochila sp. C (II 1367) Pertusaria s pp.

3.1.11. Diplasiolejeupea pauckertii- Usnea spp. community Diplasiolejeunea pauckertii (Nees) Steph.

Usneaspp.

Graphidaceae Ramalina sp. A (# 793) Frullania cf. paradoxa Lehm & Lindenb.

Colura naumannii (Schiffn.) Steph.

Colura tenuicomis Steph.

Ramalina cochlearis Zahlbr.

3 1 2. Omphalanthus filiformis community Omphalanthus filiformis (Sw.) Nees (incl. 0. platycoleus Herz.)

Frullania brasiliensis Raddi Macromitrium guatemalense C.Miiii.M.Iongifoliwn Adelanthus decipiens (Hook.) Mitt.

Plagiochila comiculata Dum. Lejeunea sp. A(# 572) Taxilejeunea pterigonia (L. & L.) Schiffn.

Leptoscyphus amphibolius (Nees) Grolle Omphalanthus ovalis (Lindenb. & Gott.) Gradst.

Lejeunea reflexistipula (Lehm. & Lindenb.) Step h.

Plagiochila dominicensis Tayl.

Pireella cavifolia (Card. & Herz.) Card.

3.1.3. Upper Montane Rain forest canswy" t,ypicum" community Anastrophyllwn piligerum (Nees) Steph.

3.1.4. Pla~iochila fuscolutea community Plagiochila fuscolutea Tayl.

3 2 Tree base communities of the Upper Montane Rain forest Calypogeia peruviana Nees & Mont.

Telaranea nematodes (Gott. ex Arnst.) Howe Lophocolea aff. connata (Sw.) Nees Riccardia andina (Spruce) Herz.

Plagiochila esmeraldana Step h.

Prionolejeunea spp.

Racopilum tomentosum (Hedw.) Brid.

Porothamnium cf. comosum Hymenophyllum sp. B (# 588) Leskeodon cubensis (Mitt.) Ther.

Thuidium delicatulum (Hedw.) B.S.G. var. peruvianum Cyclolejeunea accedens (Gott.) Evans Syrrhopodon lycopodioides (Brid.) C. Miill.

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1 1 1 40 <I I 2 60 <I 4 25 12 <I <I <I 4 10 10 10 16 <I 9 20 100 <I I 3 100 2 20 40 64 <I 9 30 52 <I <I 5 40<1520

28 <I <I <I 20 <I <I <I 4 <I <I <I 4

3 1 2

1 1

3 1 3

2 28 <I 2 10 2

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I 1 56 <I 4 20 7 <17 2513 9 <I <I <I 6 <I 5 10 ' 6 I I I

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25 <I <I <I S <1 <1 <1'15 91 <I 3 10 25 <1 1 3i ss 19 I 8 30 8 1 2 9 6

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5 5 5

3 <I <I <I

3 3 3

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4 <I <1 <1 8 <I <I <1

77 <I 8 70 13 76 <I 12 65 3~

64 <I 10 56 <1 3 56 <I <1 52 <I 2 46 <1 2 41 <1 3 39 <1 3 31 <1 2 21 <I 9 18 <1 1

85 2

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4

6 8 10

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Epiphyte communities of tropical montane rain forests. II

3 1 4

3 1 3

-

3 2 1 1

3 2 1 3

3 2 1 2

3 2 2 1

59

3 2 2 2

4 1

1 6 <1 <1 <1 9 <1 <1 <1 9 <1 <1 <1 5 <1 <1 <1 2513 <1 1 2 23 <1 1 5 '

7

7 <1 <1 <1 67 <1 3 10 7 1 1 1

4 <1 <1 <1 5 23 <1 <1 1 1 <1 2 <1 <1 <1

<1

5 2 <1 <1 <1

11 <1 <1 <1

70t3<11 2 65 32 <1 2 10

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11 <1 <1 <1

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11 <1 <1 <1

3 15 <1 10 55 4

50 <1

2 2 2

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<lt5 <1 <1 3i 55 <1 <1 2 1 1 9 <1 <1

151 10

4 2

35 <1 6 40

<1 3 5 81 <1 32 95

55 <1 1 1 2 <1 <1 <1 36 <1 <1 <1 18 <1 <1 <1 2 <1 <1 <1 18 <1 1 2 27 1 33 98 2 15 15 15 9 <1 <1 <1 9 <1 <1 <1 9 <1 <1 <1 2 <1 <1 <1 9 5 5 5

65 25 15 10 5

10 35 25 18 <1 <1 <1 5 15

<1 2 10 71 <1 <1 1 43 <1 <1 <1 14 <1 <1 <1 14 <1 <1 <1 <1 1 <1 <1 10

6 9 2 <1 12

11

15 5 <1 15

3 <1 2 5 33 <1 <1 <1 <1 33 <1 2 5 1 1 1 22 <1 3 5 2 <1 <1 <1 22 <1 22 1 8 15

40 60 20 20

1 3 5 <1 <1 2 5 5 5 1 1 1

14 <1 <1 <1 33 <1 <1 2 20 <1 <1 20 <1 <1 11 1 1 40 <1 1 11 5 5 5 40 1 1 14 <1 <1 <1 20 5 5

9 3 3 3 5 <1 <1 <1 14 <1 <1 <1 18 <1 5 10

subito e.V. licensed customer copy supplied for Universidad Ceu San Pablo (SLI11X00058E)

<1 <1 2 1 5

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J.H.D. Wolf

60 Table 1. (cont.)

COMMUNITY 3 1 1 1

3 1 1 2

3 1 2

13 <1 2 5

8 <1 4 15

3 2 1 l:b::m~:mph)%.am liml~:oii - ~z:Zlloiii!lt~:!.lt!lli;am !:2mm!.loiti!::l

Hymenophyllurn lindenii Hook.

Bazzania breuteliana (Lindenb. & Gott.) Trevis.

J,21.2, ~PiiDiii~ZQWD'!il~i:i- l!li!l;iQtb~illm DQXQ-Im\Dil!ilD:i!il 'Qmml!Dit): Scapania portoricensis Hampe & Gott.

Plagiotheciurn novo-granatense (Hampe) Mitt.

Porotrichodendron superbum (fayl.) Broth.

Blepharostoma trichophylla (L.) Dum. Riccardia ciliolata (Spruce) Gradst.

21:2 Bbi2:12!;llDillm liodigii 'l2mmuoit)l Rhizogonium lindigii (Hampe) Mitt.

Cephalozia crassifolia (Lindenberg & Gottsche) Fulford Calypogeia cyclostipa (Spruce) Step h.

Vittaria remota Fee 3,2 2. S)::m5!~~Dil hmo~iilttii - H:4!D!:IIii 5!ili~m 'l2mm!.loiti~:~ Symphyogyna brongniartii Mont.

Hypnella pilifera (Hook. & Wils.) Jaeg.

Heteroscyp hus polyblepharis (Spruce) Schiffner Plagiomnium rhynchopho rum (Hook.) T.Kop. Lophocolea bidentata (L.) Dum. Cyclodictyon rubrisetum (Mitt.) Kuntze Monoclea gottschei Lindb.

Stenodictyon wrightii (Sull. & Lesq.) Crosby 3 2 21. Milli:mtbomoi!.lm mptll~ 'Qmm!.loit)l Mittenothamnium reptans (Hedw.) Card. I Ctenidium malacodes Mitt.

Taxilejeunea lancifolia Steph.

Lophocolea muricata (Lehman) Nees Plagiochila macrifolia Tayl.

Aneuraspp .

Taxilejeunea sulphurea (Lehm. & Lindenb.) Schiffn.

3 2 2.2. Ri";amiooberro~ '2mm!.loit)l Riccardia aberrans (Steph.) Gradst.

Pyrrhobryum mnioides (Hook.) Manuel Plagiochila rutilans Lindenb. i. !:12mmuoiti!::l 2f th~: ~l21lll~i~ dw:od f2~t Zygodon fragilis Robins Blepharolejeunea securifolia (Step h.) Schust.

4 <1 <1 Frullania tetraptera Nees & Mont.

28 <1 <1 Metzgeria metaensis Kuw. Parmelina sp. A (# 1185) Leptogium mandonii Jorg.

Bryaceae sp. A(# 1180) Plagiochila sp. A (# 1164) Tortula andicola Mont. ~.1, !:!ltm&:i~twm !11.1hitll~ - !:, !lSI!.IiW2&:iense '2mmuDi~

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Epiphyte communities of tropical montane rain forests. II

3 2 1

3 1 4

3 1 3

1

3 2 1 2

3 2 1 3

3 2 2 1

3 2 2 2

4 1

4 2

6 <I <I I 30 <I 2 10 64 <I 8 30 65 <I 2 65 43 I 2 5 11 2 2 2 15 <I I 2 4 <I <I <I 100 <I 26 70 40 <I 6 20 71 <I 19 80 ~I

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11 <I <I <I 4 <I 2 4 18 <I 8 15 50 <I 14 60 65 <I 10 45 14 <I <I <I 18 <I <I <I <I 2 I I I 45<1865 43 <I 20 90 20 <I <I <I 20 I 3955 2 2 2 2

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<I <I <I 100 <I 37 27 <I <I <I 36 <I <I <I <I <I <I 71 <I <I <I <I <I 9 5 5 5 15 <I <I <I 57 I 26 43 5 7

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<I I 45 10

5 5 5

10 <I <I <I 2

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18 <1 3 5

<1 <I <I

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2 4

2 4

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I I

10

9 <I <I <I 15 <I <I 2 14 5 5 5 78 <I 10 <I <I I 78 2 20 <I I <I 44 <I 22 <I 33 <I <I <I <I 33 <I

:I

27 <I 13 45 77 <I 28 80

<I <I <I 14 I

89 <I 5 67 <I 4 56 <I <I 44 <I 7 1 I 44 <I 9 33 <I 2

8 <I <I <I

IS 40 <I 3 5 2 20 <I <I <I 2 20 <I <I <I 25 35 4

<I <I <I 18 <I 2 4 SS <I 18 60 14 I I I 11 20 20 20 100 I 43 80 60 <I <I I 10 <I <I <I I 3 5 40 5 8 10 14 <I <I <I

<I <I <I

<1 <1 <1

II 25 100 <I 7 20 67 93 <I 4 20 54 <I <I <I 100 <I 14 65 8S <1 2 20 20 <I <I I 31 <I <I <I 80 <I 13 50 46 <I 33 <I 1 5 31 <I 7 1 I 1 15 <I 7 <I <I <I 23 <I 7 I I 1 8 <I

subito e.V. licensed customer copy supplied for Universidad Ceu San Pablo (SLI11X00058E)

2 10 2 5 45 90 <I I <I <I

J.H.D. Wolf

62 Table 1. (cont.)

COMMUNITY 3 1 1 1

3 1

1 2

3 1 2

Cetrariastrum dubitans Sipm.

4 2 2 2 Cetrariastrum equadoriense (Sant.) Sipm.

Leptodontium cf. viticulosoides (P.-Beauv.) Wijk & Marg.

Hypotrachyna sinuosa (Smith) Hale 28 <I <I 2 Parmeliella nigrocincta (Mont.) Miill. Arg. Hypotrachyna boquetensis (Hale) Hale 4 2 Campylopys pittieri community Campylopus pittieri Williams I C. fragilis (Brid.) B.S.G. 13 <1 <1 <1 2 <1 <1 <1 4.: Peltigera pulverulenta (fayl.) Nyl. Leptodontium wallisii (C. Miill.) Kindb.

Leptodontium capituligerum C. Milll.

Nephromopsis? sp. A (# 1172) Zygodon pichinchensis (Tayl.) Mitt.

Accompanying taxa with a preference for the Upper Montane Rain forest lor dwarf forest>.

Chorisodontium mittenii (C. Miill.) Broth.

4 <1 <1 <1 63 <1 4 30 2 <1 <1 <1 5~ Catagonium brevicaudatum C. Miill. ex Broth.

3 <1 <1 <1 Hypotrachyna densirhizinata (Kurok.) Hale 4 <I <1 <1 9 <1 <1 1 2 I I 1 Sphaerophorus formosanus (Zahlbr.) Asah 4 1 1 1 25 <1 3 10 1~ Hypotrachyna andensis Hale 4 1 1 1 2 1 1 I Fernandezia sanguinea (Lindt.) Garay & Dunst.

4 <1 <1 <1 2 2 2 2 Riccardia capillacea 6 32222555 Stelis sp. (# 972) 3 5 5 5 2 Stelis sp. A(# 971) 3 1 1 1 2 Bromeliaceae sp. A (# %1) 3 5 5 5 4 Femandezia sp. A(# 941) 3 <1 <1 <1 2 Leprocaulon albicans (Th. Fr.) Nyl. ex Hue 3 <1 <1 <1 2 Lepanthus sp. A (#638) 2 <1 <1 <1 ' 2 Plagiochila sylvicultrix Spruce 3 1 2 2 2 Plagiochila bogotensis Gott.

2 <1 <1 <1 Semiramisia cf. speciosa 2 1 Leciophysma? sp. A(# 621) 3 1 2 2 Prionolejeunea cf. aemula (Gott.) Evans 3 1 2 2 Grammitis senilis (Fee) Morton 3 <1 <1 <1 Radula plumosa Mitt. ex Steph.

3 <1 <1 <1 Leptoscyphus physocalyx (Hampe & Gott.) Gott.

2 3 1 :2 :2 Cyrtolejeunea sp. A 5 <1 <1 <1 2 Metzgeria leptoneura Spruce 7 <1 <1 <1 Cavendishia cf. pubescens (HBK.) Hemsl.

6 10 <1 2 5 Disterigma acuminata (HBK.) Nied.

10 <1 4 10 Adelanthus lindenbergianus (Lehm.) Mitt.

2 Rhynchostegiopsis cf. flexuosa (Sull.) C.Miill.

2 Scaphosepalum sp. A ( # 824) 4 Elaphoglossum squamipes (Hook.) Moore vel valde aff.

4 Lophocolea quadridentata Spruce 6 Elleanthus kermesinus (Lindl.) R.f.

11

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63

Epiphyte communities of tropical montane rain forests. II

3 1 3

3 1 4

3 2 1 1

3 2 1 3

3 2 1 2

3 2 2 1

3 2 2 2

4 1

4 2

67 <I 9 50 60<1720 13 <1 <I <I

33 <I 5 15 20 <I <I I 13 I 18 35 :I

43 <1 4 60 45 <I 6 65

25 <I 4 20 14 <1 <1 <1

13 <1 <I I 92 <I 6 30 46 2 7 15

31 15 15 8 :I

55 <I 8 60 87 <I 12 90 9 <I <I <I 25 <I 2 IO 11 1 3 10 10 <I <I <1 4 1 <1 1 19 <I <I 2 28 <I <1 2 18 <I <I <I 5 I I I

6

:1

z

<I <1 <1

18 <1 1 2

2 2 2 2 2 4 2 4 5 2 2 2 2 2 <1 <1 <1 2 <I <1 <I 2 <1 <I <I 9

:1 1

I

5

3 3 3

I

20 1 1 1

1

2 2 2

22 <I <1

10 <1 <1 <1

14 5 5 5

:1

11 I I I

'1

2 20 20 20 2 <1 <1 <1 6

1 2 5

0 2 <1 <1 <1 2 <1 <1 <1 4 I I 1 2 I 1 4 2 4 5 6 <I 27 72 30 <1 <I 11 2 3 5 6 <I 3 5

5 <I <I <1 15 <1 2 5 <1 <1 <1 9 5

<1 <I <I <1 <1 <1

27 <I <I 2 85 <I 30 70 8 <I <I <I

5 <1 <I <1

5

<I 18 73 <I I 2

4 4 4

20 <I 3 5 9 <1 <I <I 5 60 60 60

subito e.V. licensed customer copy supplied for Universidad Ceu San Pablo (SLI11X00058E)

20 1 I I

]. H. D. Wolf

64 Table 1. (cont.)

COMMUNITY 3 1 1 1

3 1 1 2

Hymenophyllum (Sphaerocionium) elegantulum v.d.Bosch # 725 Symphyogyna brasiliensis Nees Bartramia angustifolia Mitt.

Campylium ? sp. A Relbunium hypocarpium subsp. nitidum (HBK.) Ehrendf.

Dicranodontium meridionale Bartr.

Plagiochila superba Lindenb.

Symphyogyna marginata Steph.

Accidental taxa with a preference for the Lower Montane Rajn forest Heterodermia casarettiana (Massal.) Trevis 4 <I <I <I 3 <I <I <I Lopezaria versicolor (Fee) Kalb. & Haf. 4 <I <I <I Hypotrachyna endochlora (Leight.) Hale 4 1 1 1 Parmotrema peralbidum (Hale) Hale 4 <I <I <I Phyllopsora sp. A (# 427) 4 2 2 2 Chiodecton spp.

8 <I <I I 9 <I <I 2 Frullania caulisequa (Nees) Nees 3 <I <I <1 Sticta fuliginosa (Dicks.) Ach. 3 <1 <I <I Hypotrachyna costaricensis (Nyl.) Hale 3 2 2 2 Aptychella proligera (Broth.) Herz.

Frullania arecae (Spreng.) Gott.

Lejeunea flava (Sw.) Nees • Sticta weigelii (Isert) Ach. • Acanthocoleus aberrans (Lindenb. & Gott.) Kruijt Meteoridium remotifolia (Hornsch.) Broth. • Metzgeria spp. (sect. Metzgeria) Marchesinia brachiata (Sw.) Schiffn. • Lejeunea sp. B (# 561) Ceratolejeunea cf. maritima (Spruce) Steph.

Heterodermia obscurata (Wulf.) Trevis.

Frullania standaertii Steph. • Squamidium nigricans (Hook.) Broth.

Lejeunea megalantha Spruce Lejeunea laetevirens Nees & Mont.

Isopterygium tenerum (Sw.) Mitt.

Plagiochila subplana Undenb.

Cheilolejeunea choachina (Gott.) Grolle Coccocarpia pellita (Ach.) Milll. Arg. em. Sant. • Lobaria spp.

Heterodermia Ieucomela (L.) Poelt • Phyllopsora spp.

Fissidens spp.

Hypopterigium tamariscinum (Hedw.) Brid.

Accompanyjn~ taxa withoyt a preference for ejther the Lower or Upper Montane Rajn forest. crustose lichen indet (sterile) 80 <I s IS 28 <I s 10 Sticta spp.

8 <I <1 <I 25 <I 18 60

subito e.V. licensed customer copy supplied for Universidad Ceu San Pablo (SLI11X00058E)

3 1

2

7 <I 2 5

I

1 1

3 <I

2 7 <1 <1 1 '

2 2 5 <1

2

2

<1 1

I

13 <I 2 15, 10 <I <1 <1, 7 I 11 25'

16 d<l 2 15 <I 1 9 <1 1 3 5 <1 <1 1 2 <I <1 <1 <1 1 2 <1 2 5 <1 <1 <1 <1 <1<1.:1 <1 <1 1 <1 3 5 <1 <1 .d <I <1 <1

11 <I 1 <I <1

10 8 8 7

7 7 7

5 5 5 3 3 2

2151515 7 <1 <1 <1 7 <I <1 <1 11<1850 8<1<11 3

<1 5 tO

3 1 3 5 64 <1 5 (,5

Epiphyte communities of tropical montane rain forests. II

-I

2 2 2 2 9

3 2 1 2

3 2 1 1

3 1 4

3 1 3

10 10 10 <1 <1 <1 10 10 10 1 1 1 <1<11

5 5 5 5 5 27 <1 5 15 15 5

3 2 1 3 5 <1 <1 <1 <1 <1 1

5 <1 <1 <1 <1 <1 1

3 2 2 1

5 <1 <1 <1 <1

65

3 2 2 2

4 1

4 2

20 1 1 1

1

20 <1 <1 <1

14 3 3 3 11 1 1 1 5

I

1

1 1 ' 2

1

I

15, <1,

z 4

<1 <1 <1 <}<11

25' 2

9 1

5 d

2 d

2

5

.d 1

5 .:1 ~1

20 <1 <1 <1

15 ~1 ~1

so tO

22 2 6 10 6 4 2 2 2

<1 <1 <1 <1 3 5 303030 <1 <1 <1 5 5 5 9

14 <1 <I <I

<1 <1 <1

22 3 4 5

5

65

6 <I 8 15 2 30 <I 4 15 6

I I 14 35

5 <I <I <I 20 <I <I 2

5 Phytocoenologia 22

subito e.V. licensed customer copy supplied for Universidad Ceu San Pablo (SLI11X00058E)

20 <I <I I 8 <I <I <I 60 <I 23 55 38 <I 11 25

66

J.H.D. Wolf

Table 1. (cont.)

COMMUNITY 3

Metgeria decipiens (Mass.) Schiffn.

Sticta cf. laciniata (Sw.) Ach. Leptogium burgessii (L.) Mont.

Porotrichum spp.

Plagiochila diversispina Step h.

Dimerella spp.

Lejeuneaceae Microlejeunea spp.

Orchidaceae Odontoglossum ramosissimum Lind!.

Leptoscyphus cuneifolius (Hook.) Mitt.

Normandina pulchella (Borr.) Nyl. Tillandsia sp. (sect. inflata??)

Megalospora tuberculosa (Fee) Sipm.

Drepanolejeunea inchoata (Meissner) Step h.

Taxilejeunea pallescens (Mitt.) Steph.

Leptogium spp.

Pannaria rubiginosa (Ach.) Bory Radula tenera Mitt. ex Step h.

Leptogium resupinans Nyl. Erioderma verruculosum Vain.

Radula sonsonensis Step h.

Lepidolejeunea spongia (Spruce) Thiers Epidendrum sp. A (# 645) Stelis spp.

Phyllogonium fulgens (Hedw.) Brid.

Zygodon cf. peruvianus Sull.

Lindigianthus cipaconeus (Gott.) Kruijt & Gradst.

Lobaria crenulata (Hook.) Trevis.

Plagiochila bidens Gott.

Bromeliaceae Plagiochila choachina Gott.

Polypodium angustifolium Sw. complex Plagiochila alpina Gott.

Prionodon densus (Hedw.) C. Mull. var.luteo-virens Heterodermia microphylla (Kurok) Swinsc. & Krog Radula voluta Tayl.

Leptolejeunea elliptica (Lehm. et Lindenberg) Schiffn.

Polypodium glaucophyllum Kunze Plagiochila adiantoides (Sw.) Dum. Prionolejeunea sp. A Ceratolejeunea patentissima (Hpe. et Gott.) Evans Rare taxa Megalospora admixta (Nyl.) Sipm.

Splachnaceae sp. A (# 603)

subito e.V. licensed customer copy supplied for Universidad Ceu San Pablo (SLI11X00058E)

3 1 1 1 1 1 2 12 <1 <1 1 13 <1 <1 2 6 5 10 15 4 <1 <1 <1 3 3 3 3

3

10 10 10

88 <1 <1 2 56 <1 <1 5 9 1 2 3 64 28 28 24 16 8 4 4 4 4 4 4

<1 1 5 50 <1 <1 <1 19 <1 <1 2 6 10 3 10 <1 1 3 <1 <1 1 1 1 1 <1 <1 <1 <1 <1 <1 2 2 2 3 5 5 5 <1 <1 <1 9 6 3 3 3 3 3

<1 <1 2

<1<1<1 1 1 1

3 1 2 <1 <! ~ 1 4 to 1 4 to <1 <1 ( 3 <1 5 to 5 <1 1 ( 39 <1 1 ~ 44<1<1( 28 1 5 ts 3 5 5 ~ 43<1<1\ 10 <1 <1 l 3 <1 <I ""1 7 2 s to 25 <1 <I ~

52 5 5 10

29 <1 <1 ~ 5 <1 <I l 11 <1 1

s

<1 <1 <1

<1 1 20 3 1 <1 <1

<1 3 20 3 1 <1 <1

<1 5 20 3 1 <1 <1

2 13 26 5 15 13 11

<1 <1 ""' <1 1 IO <1 3 10 1 2 s <1 3 15 <1 3 10 <1 3 20 7 <1 <1 1 13 <1 6 18 25<1520 29 <1 3 15 2 <1 <1 "' 2 1 1 1 3 <1 1 1 20 <1 4 25 2 2 2 2 10 <1 <1 1 8 <1 2 s 18 <1 1 s 10 <1 1 s 10 <I <1 2 7 <I 2 s

8

1 1 I 5

15 32 60

-

67

Epiphyte communities of tropical montane rain forests. II

3 1 4

3 1 3 .;

I

0 0

11 <1 <1 <1 6 <11 2 2 3 3 3 6 <1 <1 2

2 2 28 13 2 13 6 4

3 3 3 2 2 2 1 1 1 3 2 1 9 <1<1<1 45 <1 <1 5 5 1 1 1

3 2 2 1

3 4 4 2 1 2 2 2 56 <1 <1 1 80 <1 3 10 33 <1 <1 1 23 <1 <1 <1 13 5 8 10 8

1 1 1

8

1 1 1

78 <1 15 70 20 2 2 2 45 <1 26 60 10 2 6 10 43 5 15 25 11 1 1 1 27 5 25 40

<1 <1 <1

4 <1 <1 <1 <1 <1 2 <1 <1 <1 <1 2 11 <1 <1 1 2 5 5 5 5 2 5 5 <1 <1 <1 6 <1 <1 <}<1<1 2 <1 <1

1 <1 9 <1 <1 <1 <1

33 <1 2 5

7

1 1 87 <1 <1

5 <1 <1

15 <1 <1 <1

13 <1 <1 <1 15 <1 <1 <1 13 <1 <1 <1 8 <1 <1 <1

<}<1<1

4

<1 <1 <1

z

<1 <1 <1

5 <1 <1 <1 11 5 5 5 27 1 2 3 10 <1 <1 I

I

0 0

5 <I <I <I

11 <I <1 <1 2 <1 <I <I

5 10 10 10

5 0 0

4

<1 5 10

5 <I <1 <I 2

8

5

20 5 5 5

20 5 39 70

0

15 <1 19 60 2 10 10 10 5 2

I

I

I

I

22 <I 13 25

I

5 505050

I

22 <I I 2

5

7 <I <I <I 2

8 <1 <I <1

<1 <1 <I

9 <I <I <I

7

subito e.V. licensed customer copy supplied for Universidad Ceu San Pablo (SLI11X00058E)

I

I

I

68

J.H.D. Wolf

Table 1. (cont.)

COMMUNITY 3 1 1 1

Lejeunea aff. ambigua Lindenb. &: Gott.

Grammitis semihirsuta (Kl.) Morton Araceae Elaphoglossum aff. cuspidatum Moore Hymenophyllum nigrescens Liebm.

Plagiochila densispina Steph.

Themistoclesia recondita A.C. Smith Heterodermia comosa (Eschw.) Follm. &: Redon Hypotrachyna cf. pulvinata (Fee) Hale Parmotrema subsumptum (Nyl.) Hale â&#x20AC;˘ Pseudocyphellaria aurata (Ach.) Vain.

Epidendrum xylostachyum Lindley Dictyonema sp. A (# 1008) Plutarchia monantha A. C. Smith Hydrocotyle Bonplandii A. Rich Bazzania sp. (aff. chilensis)

3 1 1 2

3 1 2

5 5 5 5 5

1 ~ 5 .::1 1 2 .::1 .<:1 1 1 3 5 .::1 4 tO .::1 .<:1 1 , , 5

3 3 3 .::1 .<:1 <1 3 1 ~ 3 3 .::1 .<:1 <1 3 .::1.::11

In addition, the following taxa were only recorded once: In commu":ity_4.2.: Cladoniq subsquamosa Kremp (1 %), Pannariaceae (< 1 %), Rhodobryum grandifolzum (Tayl.) C. Miill. (<1 %). In community 3.1.1.1.: Bacidia sp. (<1 %), f!uell~r: ~odesta (Krempelh.)

Miill. Arg. (2 %), Catillaria sp. A #984 (10 %), Caloplaca brebzssomz (Fee) Haffeln. & Poelt (< 1 % ), Hypotrachyna rhabdiformis (Kurok.) Hale (2% ), Lecidea sp. B # 805 (2% ), Pacby, phyllum hispidulum (Reichb. f.) Garay & Dunst. (1 %), Phyllopsora sp. A #427 (2 %), Telos, chistes jlavicans (Sw.) Norm. (<1 %). In community 3.1.1.2.: Coccocarpia erythro:x;yli (Spreng.) Swinsc. & Krog (< 1 % ), Epidendrum cerneum group# 890 (5% ), M acromitrium cirrosum (Hedw.) Brid. (1 % ), Myriotrema concretum (Fee) Hale (2% ), Trichosalpin:x: sp. A # 859 (30 %). In community 3.1.2.: Cheilolejeunea rigidula (Mont.) Schust. (< 1 %), Chiodec, ton sp. A #687 (1 %), Clusia sp. (3 o/o ), Dioscorea sp. A# 1431 (2 o/o ), Erioderma sorediatum Galloway & Jorgensen (< 1 % ), Erioderma sp. A # 1281 (< 1 % ), Frulliana cf. moritziana Lindenb. & Gott. (5% ), Grammitis jamesonii (Hook.) Morton(< 1 % ), Groutiella apicu[a, ta (Hook.) Crum & Steere(< 1 %), Harpalejeunea cf. o:x:yphylla (Nees & Mont.) Steph. (< 1 % ), Heterodermia circinalis (Zahlbr.) W.A. Weber(< 1 % ), H.lutescens (Kurok.) Follm. & Redon (< 1 %), H. speciosa (Wulf.) Trevis. (< 1 o/o ), Hymenophyllum sp. C # 1432 (5 %), Lepanthes dasyphylla (2 o/o ), Lepidolejeunea involuta (Gott.) Grolle (1 o/o ), Leucobryum po[a, kowskyi (C. Mi.ill.) Card. (1 %), Metzgeriafruticola Spruce(< 1 %), Oreopana:x: cf. capitatum Qacq.) Dene & Pl. (2% ), Pachyphyllum sp. (< 1 o/o),Peltigera dolichorhiza (Nyl.) Nyl. (5 %), Peperomia angularis C.DC. (2 %), Prionodon densus (Hedw.) C. Mi.ill. var. densus (< 1 %), Pseudocyphellaria intricata (Del.) Vain. (1 %), Pyrrhobryum spiniforme (Hedw.) Mitt. (< 1 % ), Syzygiella perfoliata (Swartz) Spruce (1 o/o ). In community 3.1.3.: Dicranoweisia sp.

A# 969 (1 %), Dicranum cf. paramicola C. Mi.ill. (5 %), Dictyonema sericeum (Sw.) Berk.

(30% ), Neckera chilensis Schimp. ex Mont.(< 1 % ), Parmotrema sancti-angeli (Lynge) Hale (1 % ), Peperomia saligna HBK. (5% ), Pleurothallis antenniferae Lind!. (40% ), Pleurothallis sp. (1 %), Polypodium remotum Desv. (1 %), Scaphyglottis bicornis (Lind!.) Garay (5 %), Si, phocampylus venosus (Gleason (5 %), Sphyrospermum bu:x:ifolium Poepp. & End!. 2 %). In community 3.1.4.: Cladonia squamosa (Scop.) Hoffm. var. squamosa(< 1 %), Fernandezia

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69

Epiphyte communities of tropical montane rain forests. II 3 1

3 1 3

4

3 2 1 1

3 2 1 2

3 2 1 3

3 2 2 1

3 2 2 2

4 1

4 2

5 I

5 10

5 d 3

d

6

4

1 3 5 <1 3 5

13 1 4 10 4 1 2 2 10 2 9 15

aff. ionanthera ~~chbf. f. & Warsc.) Schltr. (5 %), Grammitis aff.jamesonii (Hook.) Morton (1 %), Grammztzs heteromorfa (Hook. & Grev.) Morton (1 %), Herbertus colombianus Van Reenen (< 1 %), Leiomela ecuadorensis Robbins (2 %), Lycopodium sp. A # 1092 (5 %), Myrrhidendron glaucescens (Benth.) Coult. & Rose (5 %), Pleurothallis sp. sect. Corocodilanthe (2 %). In community 3.2.1.1.: Cephalozia bischlerae Fulford(< 1 %), Plagiochila divaricata Lindenb. (5 %), Polypodium aff. mindense Sodiro (1 %), Radula nudicaulis Steph.

(1 %). In community 3.2.1.3.: Kurziaflagellifera (Steph.) Grolle (< 1 %), Pileafallax Wedd.

(5 %). In community 3.2.2.1.: Plagiochila laetevirens Lindenb. (1 %), Thuidium minutulum (Hedw.) B.S.G. (<1 %). In community 3.2.2.2.: Radula sinuata Gott. ex Steph. (<1 %).

The most common taxa exclusive for the canopy communities are Anoplolejeunea conferta, ] amesoniella rubricaulis, Leptoscyphus jackii, L. porphyrius, Plagiochila echinella sensu lata (incl. P. hansmeyeri and P. paludosa), Prionodon fusco-lutescens, Trachylejeunea dominicensis and Hypotrachyna laevigata (not separated from H. producta). Within the canopy a twiglet community and an outer/middle branch community, both with a relatively high abundance of lichens, and three inner crown communities dominated by bryophytes, are distinguished.

Exclusive species for all five tree base communities are, among others, Calypogeia peruviana, Cephalozia crassifolia, Lophocolea aff. connata and Telaranea nematodes. The tree base communities may be divided in a Hymenophyllum lindenii-Bazzania breuteliana group with three communities and in a group of two communities with Symphyogyna brongniartii and Hypnella pilifera as most common exclusive taxa.

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]. H. D. Wolf

70 3.1. Canopy communities 3.1.1.1. Diplasiolejeunea pauckertii-Usnea spp. community

Composition and structure (Table 2; Fig. 1) ::·NR = 25; ::·NTt = 107; ::·NT10% = 48; ::·sR

= 35 % .

Table 2. Composition and structure of the Diplasiolejeunea pauckertii-Usnea spp. communi· ty (type 3.1.1.1.); 25 releves. per community no. of taxa, no. of taxa, total freq. >10% mosses liverworts crustose lichens foliose lichens fruticose lichens bryophytes lichens bryophytes & lichens flowering plants ferns, excl. filmy ferns bryophyte & lichen layer dry weight (g/dm2) thickness (em)

1

6

25 4 8 9 26 21 47 1

46 14 23 15 52 52 104 3

0

0

mean 1.6 1.2

s. dev.

1.3 I

per releve mean no. mean cover of taxa (%) 0.4 0.3 10.2 16.3 2.3 12 3.1 9.3 4.6 27.4 10.6 16.6 7.7 48.7 18.3 65.3 0.2 0 0

®~ Fig. 1. The Diplasiolejeunea pauckertii- Usnea spp. community at a height of 22m and 3190 m altitude. Scale bar equals 5 em.

•:· NR, NTt and NT10% stand for number of releves, total nr. of taxa and nr. of taxa with frequencies(= presence degrees) over 10 %, respectively. SR stands for the mean similarity ratio, a quantitative coefficient of similarity between relevcs. Species with frequencies 10 % were excluded from the calculations.

<

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Epiphyte communities of tropical montane rain forests. II

.·

71

Exclusive taxa are Diplasiolejeunea pauckertii, Frullania cf. paradoxa, Graphidaceae, Ramalina sp. A and Usnea spp. (high abundance). Accompanying taxa are Anoplolejeunea conferta, Aureolejeunea paramicola, Drepanolejeunea spp., Frullania peruviana, Leptoscyphus cuneifolius, Microlejeunea spp., £verniastrum catawbiense, Hypotrachyna laevigata, Oropogon bicolor and 0. loxensis.

The high abundance of fruticose lichens, mainly Usnea spp., and the pendent liverwort Frullania peruviana are most conspicuous .

Ecology and distribution':-::· (Table 3) The Diplasiolejeunea pauckertii-Usnea spp. community is an outer crown community with an altitudinal range from 2740-3670 m. This community shares many species with the Diplasiolejeunea pauckertii-Brachiolejeunea laxifolia community, an outer crown community mostly recorded between 2460 and 2550m.

Out of 2 releves in the Diplasiolejeunea pauckertii- Usnea spp. community sufficient soil could be gathered for total nutrient analysis (in mg/g): N 11.4 and 8.6, P0 4 -P0.9 and 0.7, K 1.2 and 1.9, Ca 2.4 and 2.0, 50 4 2.6 and 2.1. These values are lower than recorded in the Diplasiolejeunea pauckertii-Brachiolejeunea laxifolia community.

Table 3. Ecology and distribution of the Diplasiolejeunea pauckertii-Usnea spp. community (type 3.1.1.1.); 25 releves.

mean I preference altitude (m) 2740-3670 3370-3670 absolute height (m) 7.0-27 18.2 relative height(%) 57-96 86.6 diameter branch/trunk (em) 1-5.0 1.8 position within the tree (I-V) IV-Y V inclination bark surface (0 ) 0-70 40.5 thickness suspended soil (em) <0.1 <0.1 diffuse site factor n= 6 (s. dev.)

21.4-38.7 28 (6.6) phorophytes: Axinaea sp., Clusia aff. multiflora H.B.K., range

Escallonia myrtilloides L.f. var. myrtilloides,Weinmannia mariquitae

Szyszyl.,W. pinnata L. and W. pubescens H.B.K.

3,1,1,2, Oropagon bicolor-0. loxensis community

~ 90

th nICS

Composition and structure (Table 4; Fig. 2) NR = 32; NTt = 133; NTlO % = 53; SR = 17 % .

Exclusive taxa for the Oropogon bicolor-0. loxensis community are lacking.

However, the community may easily be separated by the characteristic taxa of

::·::· Zonification within the tree is after J oHANSSON (1974): I = tree base, II = tree trunk, III first third part of a canopy branch, IV middle part of branch, V = outer part of

=

=

branch.

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]. H. D. Wolf

72

the two canopy lichen communities , the Oropogon bicolor-0. loxensis commu~ nity and the Diplasiolejeunea pauckertii-Us nea spp. community, and by many accompanyin g taxa. Exclusive taxa for both lichen communities are Aureolejeunea paramicola, Frullania peruviana, Everniastrum catawbiense, Oropogon bicolor and 0. loxensis. Accompanyin g species with a high presence degree are: Anoplolejeunea conferta, Chorisodonti um mittenii, Drepanolejeunea spp., Frullania convoluta, Herbertus subdentatus,]amesoniella rubricaulis, Lepicolea pruinosa, Lepidozia spp., Leptoscyphus jackii, L. cuneifolius, L. porphyrius, M icrolejeunea spp., Plagiochila echinella, Riccardia spp., Hypotrachyna laevigata and Usnea spp. The foliose lichen Hypotrachyna physcioides has a preference for this community.

In addition to the two Oropogon species, the community is dominated by liverworts. The liverworts form turfs of several centimetres in height on top of a thin layer of organic soil. Foliose lichens may attain great cover values.

ÂŽ

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Fig. 2. The Oropogon bicolor-0. loxensis community at a height of 8 m and 3670 m altitude.

Scale bar equals 5 em.

Epiphyte communities of tropical montane rain forests. II

73

Table 4. Composition and structure of the Oropogon bicolor-0. lox ensis community (type 3.1.1.2.); 32 releves. per community no. of taxa, no. of taxa, freg. >10% total 2 18 23 55

mosses liverworts crustose lichens foliose lichens fruticose lichens bryophytes lichens bryophytes & lichens flowering plants fems. excl. filmy ferns filmy fems bryophyte & lichen layer dry weight (g/dm 2 ) thickness (em)

2 6

8 22 13

25 18 43 0

73 43 116 14

0

2

1 mean 5.4 2.7

1 s. dev.

3.5 1.3

10

per releve mean no. mean cover of taxa (%) 1.7 3.5 12.4 49.4 0.8 3.3 3.9 25.6 3.7 6.7 14.1 52.9 8.4 35.6 22.5 88.5 3.2

0 0.6

Table 5. Ecology' and distribution of the Oropogon bicolor-0. loxensis community (type ).1.1.2.); 32 releves. mean/ preference altitude (m) 3190-3670 3190-3670 absolute height (m) 1.0-21 13.7 relative height(%) 7.0-92.9 68.7 diameter branch/trunk (em) 3.0-45 12.6 position within the tree (I-V) II-IV III-IV inclination bark surface (0 ) 5.0-90 55.4 exposition ( 0 ) 10-340 thickness suspended soil (em) <0.1-1 0.2 diffuse site factor n= 25 (s. dev.)

15-40.1 19.8 (5.8) phorophytes: Axinaea sp., Ocotea cf guianensis Aubl., Weinmannia mariquitae Szyszyl. andW. pubescens H.B.K. range

Table 6. C hemical composition of suspended soil in the Oropogon bicolor-0 . loxensis com111 unity (type 3.1.1.2.). residue on ignition(%) pH H20 conductivity fl.S em- 1

t

N03-N N02-N Nf14-N total N P04-P S04 K Ca

n range mean 16 2.5-5.1 3.8 10 3.6-4.3 3.7 10 350-1535 624.5 IN WATER SOLUBLE IONS (mg kg- 1 dr~ weight) n mean s. dev.

10 0.6 0.5 10 2.2 1.9 44.5 8 47.3 10 10 9 10

179.8 841.6 802.9 224.6

41.2 438.7 499.8 126

s. dev.

0.9 0.2 341.5 NUTRIENT CONTENT (mg g-1 dr~ weight) n mean s. dev.

16 15 15 15 15

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11.4 0.7 3.6 1.8

3.1

3.2 0.1 2.2 1 0.9

74

]. H. D. Wolf

Ecology and distribution (Tables 5, 6) The Oropogon bicolor-0. loxensis community has an altitudinal range from 3190-3670 m and was mostly recorded at the inner and middle sections of rather steep and exposed branches. The average diffuse site factor is 19.7 %.

Compared with communities at lower altitudes, the suspended soil has a relatively high amount of organic matter, a low pH and a low total nutrient content.

3.1.2. Omphalanthus filiformis community Composition and structure (Table 7; Fig. 3) NR = 61; NTt = 223; NT10% = 71; SR = 12%.

Exclusive taxa, within the epiphyte communities of the Upper Montane Rain forest, are Adelanthus decipiens, Frullania brasiliensis, Lejeunea sp. A., Leptoscyphus amphibolius, M acromitrium guatemalense (not separated from M. lon-

Fig. 3. The Omphalanthus filiformis community at a height of 11 m and 2550 m altitude.

Scale bar equals 5 em.

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Epiphyte communities of tropical montane rain forests. II

75

Table 7. Composition and structure of the Omphalanthus filiformis community (type 3.1.2.); 61 releves.

mosses liverworts crustose lichens foliose lichens fruticose lichens bryophytes lichens bryophytes & lichens Dowering plants ferns, excl. filmy ferns filmy ferns bryophyte & lichen layer dry weight (g/dm 2) thickness (em)

per community no. of taxa, no. of taxa, freg. >10% total 12 33 45 102 0 10 7 39 2 9 57 135 9 58 66 193 3 21

2 0

9 3

mean 4.8 3.7

s. dev.

3.4 2.2

per releve mean no. mean cover of taxa (%) 3.6 12.9 17.6 65.6 0.4 1.5 2.9 9 0.7 0.4 21.2 78.5 4.0 10.9 25.2 89.4 4.1 1 0.3

gifolium), C?mp?alanthusfiliformis, 0 . ovalis, ?lagiochila corniculata, Taxilejeunea pterzgoma a~d Stzcta spp. A~companymg species are Anoplolejeunea conferta, Drep_a:zolejeur:zea SJ?p., Lepzcolea pruinosa, Leptoscyphus cuneifolius, Metzgeria deczpzens, Mzcrolejeunea spp., Plagiochila bursata, Plachiochila echinella s.l., Prionodon fus~o-lU;tescens, Riccardia spp. and Trichocolea tomentosa.

Liverworts are do.mmatmg m term.s of richness and abundance. Local dominance may be attamed by turf formmg mosses and large foliose lichens such as in the genera M acromitrium and Sticta. Thickness of the bryophyte layer is ca.

3.5 centimetres and corresponding dry weight approximately 5 g/dm 2 â&#x20AC;˘ Ecology and distribution (Tables 8, 9) The Omphalanthus filiformis community was recorded in a wide range of different habitats from 2460- 3190 m. Many of its characteristic species, such as Table 8. Ecology and distribution of the O mphalanthus filiformis community (type 3.1.2.); 61 releves. range mean/ preference altitude (m) 2460-3190 2550-2970 absolute height (m) 7.5-23 16.1 relative height(%) 27-90 67.3 diameter branch/trunk (em) 3.5-45 12.4 position within the tree (I-V) II-IV III inclination bark surface (0 ) 0- 115 51.5 exposition (0 ) 0-360 thickness suspended soil (em) <0.1- 18 1.3 diffuse site factor n=49 (s. dev.)

7.9-34.9 16.4 (6.8) phorophytes: Brunellia cf goudotii Tul., B. occidentalis Cuatr., Clusia aff. multiflora H.B.K., Gordonia cf pubescens Pl. & Lindbl. ex Tr. et Pl., Ocotea cf. guianensis Aubl., Tetrorchidium boyacanum Croiz., Weinmannia pinnata L., W. pubescens H.B.K., W. rollottii Killip var. rollottii Tul. and Weinmannia sp.

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J.H.D. Wolf

Table 9. Chemical composition of suspended soil in the Omphalanthusfiliformis community (type 3.1.2.).

residue on ignition (%) pH H20 conductivity 11S em-I

N03-N N02-N Nt4-N total N P04-P so4 K Ca

range mean n 21 3.2-8.2 5.6 4.2 11 3.4-4.8 II 5I8.2 282-703 IN WATER SOLUBLE IONS (mg kg-1 dry weight) s. dev. n mean 10 0.6 0.5 1.5 11 1.3 11 143.5 104.1 11 11 10 11

212.7 444.5 677.8 228.6

84.1 93.9 406.1 I31.1

s. dev.

1.4 0.5 166.8 NUTRIENT CONTENT (mg g-1 dry weight) n mean s. dev.

21 22 21 21 21

14.5 0.9 4.1 3.1 6.5

2.1 0.2 1.7 2 3

Omphalanthus filiformis, Frullania brasiliensis and Taxilejeunea pterigonia are known neotropical generalists, commonly invading newly created man-made habitats such as road banks etc. (GRADSTEIN 1992).

3.1.3. Upper Montane Rain forest canopy "typicum" community Composition and structure (Table 10; Fig. 4) NR = 47; NTt = 157; NT10% = 55; SR = 14%.

The "typicum" community of the canopy in the Upper Montane Rain forest lacks high fidelity characteristic taxa. The exclusive liverwort Anastrophyllurn piligerum was only observed in 15% of the releves. The community is characterized by the common presence of the following species: A delanthus pittieTable 10. Composition and structure of the Upper Montane Rain forest canopy "typicum" community (type 3.1.3.); 47 releves.

mosses liverworts crustose lichens foliose lichens fruticose lichens bryophytes lichens bryophytes & lichens flowering plants ferns, excl. filmy ferns filmy ferns bryophyte & lichen layer dry weight (g/dm2) thickness (em)

per community no. of taxa, no. of taxa, freq. >10% total 4 23 33 63

0

8

6

18 13 86 39 125 23 7

5 37 11 48

4 2 1 mean 6.7 5.5

2

s. dev.

4.9 2.8

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per releve mean no. mean cover of taxa (%) 1.9 9.9 11.1 73.0 0.2 1.4 1.8 6.6 2 1.1 13 83.3 4 9.2 17 92.5 6.1 1.0 2.1

77

Epiphyte communities of tropical montane rain forests. II

ÂŽ

Fig. 4. The Upper Montane Rain forest canopy "typicum" community at a height of 19m and 3370 m altitude. Scale bar equals 5 em.

ri A noplolejeunea conferta, Bazzania hookeri, Chorisodontium mittenii, j ames;niella rubri~aulis, ~epicolea pruinosa, Lepidozia spp., Leptoscyphus porphyrius Plagiochda echmella s.l., Hymenophyllum sp. A and Oropogon btcolor.

The bryophyte/lichen layer, on top of 1.4 em of organic soil, is well developed as indicated by its high mean cover of over 90 %, its average thickness of 5.5 em and dry weight _o_f 6.7 g/dm2: Turf forming liverworts like the showy reddish Herbertus pensdts are conspicuous.

Table 11. Ecology and distributi~n of the Upper Montane Rain forest canopy "typicum" community (type 3.1.3.); 47 releves. range

mean/ preference altitude (m) 2740-3670 2740-3670 absolute height (m) 0.5-20.5 15.9 relative height(%) 2.1-95 64.2 diameter branch/trunk (em) 4.0-80 15.9 position within the tree (1-V) (I) II-4 III inclination bark surface (0 ) 0-100 48.7 exposition (0 ) 0-340 thickness suspended soil (em) <0.1-8 1.4 diffuse site factor n= 40 (s. dev.)

6.8-34.1 18.5 (8.4) phorophytes: Laplaca cf. pubescens Pl. & Lindbl. ex Tr. et Pl., Ocotea cf. guianensis Aubl., Tetrorchidium boyacanum Croiz, Weinmannia pubescens H.B.K., W. pinnata L. and W. mariquitae Szyszyl.

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J.H.D. Wolf

Table 12. Chemical composition of suspended soil in the Upper Montane Rain forest canopy "typicum" community (type 3.1.3.). residue on ignition(%) pH H20

conductivity jlS cm-1

N03-N N02-N ~-N

total N P04-P S04 K

Ca

n range mean 36 2.1-6.0 4.3 37 2.8-4.5 3.6 37 310-1270 600 IN WATER SOLUBLE IONS (mg kg-1 d~ weight) n mean s. dev.

34 1.2 0.8 35 0.6 0.7 21 60.1 48.8 36 37 29 37

177.5 650.1 519.3 213.2

43.3 277.4 237.8 86.8

s. dev.

0.9 0.3 303.8 NUTRIENT CONTENT (mg g-1 dry weight) n mean s. dev.

35 31 30 29 30

11.5 0.7 2.7 1.5 3.4

3.2 0.1 0.9 1.1 1.3

Ecology and distribution (Tables 11, 12) The community has an altitudinal range from 2740 m up to 3670 m and is typically recorded at exposed sites in the inner crown. Near the forest line at 3670 m it may also occur on tree bases when light intensities are high. The average diffuse site factor in such sites was 14.7 %.

3.1.4. Plagiochila fuscolutea community Composition and structure (Table 13; Fig. 5) NR = 47; NTt = 115; NT10% = 39; SR = 2 %.

The great abundance of the liverwort Plagiochila fuscolutea (= P. scopulosa) is characteristic for this community. Accompanying taxa are Adelanthus pittieri, Bazzania hookeri, Campylopus pittieri, Chorisodontium mittenii, Lepicolea pruinosa, Lepidozia spp., Porotrichodendron superbum and Hymenophyllum sp. A. A thick layer of erect, living bryophytes, packed loosely together on top of up to 1 m of accumulated organic soil gives the community a characteristic physiognomy. The (hemi-)spherical bryophyte turfs, encompassing branches, branch stumps, or protruding from tree forks have been called 'moss balls' (VAN LEERDAM et al. 1990). However, the author prefers the term 'super turf' - equivalent to a tall turf (RICHARDS 1984) of extraordinary size - since 'moss ball' has been described as an unattached growth form from forest floors, barren grounds, glaciers, lakes and sea shores (DEGUCHI & INOUE 1982).

Individual 'super turfs' may attain over 40 kg in fresh weight, depending on age of the substrate, historical factors and suitability of the location to maintain a large amount of organic matter (VAN LEERDAM et al. 1990).

On top of these turfs a whole range of vascular plants is found, e. g. Elaphoglossum sp. A, Diogenesia cf. tetranda (Ericaceae) and various Orchidaceae. At the underside pendent filmy ferns (Hymenophyllum) are conspicuous.

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Epiphyte communities of tropical montane rain forests. II

79

-ÂŽ Fig. 5. The Plagiochila fuscolutea co mmunity at a height of 8 m and 3370 m altitude. Scale bar equals 5 em.

Super turfs are poor ~n spe~ies with one or a few (co)dominant bryophyte species covering the enure avatlable substrate. As a result, the similarity ratio between releves is low, on average 2 %.

No clear succession of species is evident, even though remnants of lichen thalli sometimes could be recognized in the core of smaller turfs.

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J. H. D. Wolf

80

Table 13. Composition and structure of the Plagiochilafuscolutea community (type 3.1.4.); 47 releves. per community no. of taxa, no. of taxa, freg. > 10 % total mosses liverworts crustose lichens foliose lichens fruticose lichens bryophytes lichens bryophytes & lichens flowering plants ferns, excl. filmy ferns filmy ferns bryophyte & lichen layer dry weight (g/dm2) thickness (em)

6

16

20

51

0

2

2

12

3

8

26 5 31 4

67 22 89 13

3 2 mean 8.2

9 3 s. dev.

4.9 4.1

9

per releve mean no. mean cover of taxa (%) 2.4 27.1 7.2 67 0.1 0. 1 0.8 3.1 0.9 0.3 9.6 94.3 1.8 3.5 10.4 97.8 2.9 2.5 0 2.8

Ecology and distribution (Tables 14, 15) The inner crown Plagiochila fuscolutea community has an altitudinal range from 2740 m up to 3670 m. Super turfs are most common on the slanting trees of the 'mossy' forest near the forest line, where a single 15m tall Weinmannia mariquitae tree may carry nearl{ 115 k~ dry weight of epiphytic mass, estimated to amount up to 44 tons ha- , the h1ghest value reported so far (HoFSTEDE et al. 1993). Green biomass produced by epiphytes exceeded that of the phorophyte leaves by one third and epiphytic matter contributed greatly to above-ground nutrient capital of the phorophyte (N 49 %, P 42 %, K 20 %, Ca 28 %).

The average pH value of the suspended soil is 3.6. Nutrient concentrations are low.

Table 14. Ecology and distribution of the Plagiochila fuscolutea community (type 3.1.4. ); 47 releves. range

mean/ preference altitude (m) 2740-3670 3370-3670 absolute height (m) 0.1-18 8.7 relative height(%) 0.7-88.5 46 diameter branch/trunk (em) 5.0-70 25.4 position within the tree (I-V) I-III II-III inclination bark surface (0 ) 0-90 57.1 0 exposition ( ) 0-360 thickness suspended soil (em) 2.0-60 9 diffuse site factor n= 47 (s. dev.)

7.9-29.6 14.9 (4.5) phorophytes: Axinaea sp., Escallonia myrtilloides L.f. var. myrtilloides,Weinrru:mnia mariquitae Szyszyl. and W. pubescens H.B.K.

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81

Table 15. Chemical composition of suspended soil in the Plagiochila fuscolutea community (type 3.1.4.). residue on ignition(%) pHHzO conductivity JJ.S cm- 1

N03-N N~-N

NJ14-N total N

n range mean 41 1.7-8.8 4.4 2.8-4.4 3.6 39 285-1280 510 39 IN WATER SOLUBLE IONS (mg kg-1 d!1 weight) mean n s. dev.

0.6 39 0.7 1.4 39 1.6 67.3 29 103.8 39 39 39 39

P04-P S04 K Ca

138.7 536.3 418.4 159.3

71.1 219.2 285.9 67.5

s. dev.

1.4 0.4 220 NUTRIENT CONTENT (mg g-1 dry weight) n mean s. dev.

41 38 39 39 39

11.5 0.8 2.7 1 3

2.7 0.2 0.8 0.6 1.1

3 ..

2 Tree base communities " . , 3 .2.1.1. H_ymenophyllum lindenii-Bazzania breuteliana typ1cum commumty

Composition and structure (Table 16) NR == 11; NTt = 51; NTlO % = 26; SR = 24 % .

The Hymen~phyllum lin.denii~Bazzania breuteliana "typicum" community has no exclus1ve taxa and 1s defmed by a number of combinations of taxa characteristic of tree bases in the Upper Montane Rain forest, such as Calypogeia peruviana, .C:ephalozia c~assifolia, . Telaranea nemat?des an~ by Hymenophyllum lindemz and Bazzama breutelzana. Accompanymg spec1es are Adelanthus

Table 16. Composition and structure of the Hymenophyllum lindenii-Bazzania breuteliana "typicum" community (type 3.2.1.1.); 11 releves.

mosses liverworts crustose lichens foliose lichens fruticose lichens bryophytes lichens bryophytes & lichens flowering plants ferns, excl. filmy ferns filmy ferns bryophyte & lichen layer dry weight (g/dm2) thickness (em)

per community no. of taxa, no. of taxa, freg. >10% total 6 13 18 35

0

0

per releve mean cover mean no. of taxa (%) 2.2 15.4 75.3 7.6 0 0

0

0

1 24 1 25

1 48 1 49

0.2 9.8 0.2 10.0

90.7

0 0

0 0

1 mean 2.7 6.4

2 s. dev.

2.9 3.6

0 0

0 0

6 Phytocoenologia 22

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0

0 0.02 90.7

0.02

5.8

J. H. D. Wolf

82

pittieri, and species of Lepidozia and Porotrichum. Leafy liverworts are dominant, lichens are nearly absent.

Ecology and distribution (Tables 17, 18) Possibly, the Hymenophyllum lindenii-Bazzania breuteliana "typicum" community is a depauperate assemblage of the Scapania portoricensis-Plagiothecium novo-granatense community, which has a similar distribution: tree bases at altitudes between 2740 and 3510 m. However, the great abundance and presence degrees of thallose liverworts in the S capania portoricensis-Plagiothecium novo-granatense community suggest a higher humidity (Pocs 1982).

Therefore, both communities are distinguished until more releves and ecological data are available.

Table 17. Ecology and distribution of the Hymenophyllum lindenii-Bazzania breuteliana "typicum" community (type 3.2.1.1.); 11 releves. range

mean/ preference 2740-3510 2740-2970 0.1-8 1.1 0.3-5.9 4.3 16-45 34

altitude (m) absolute height (m) relative height(%) diameter branch/trunk (em) position within the tree (I-V) I (II) I inclination bark surface (0 ) 60-90 79 exposition (0 ) 40-360 thickness suspended soil (em) 0.1-5 diffuse site factor n= 11 (s. dev.)

2.8-13.3 7.4 (3.4) phorophytes: Brunellia cf gou.dotii T~J., Gor.do'!ia cf pubescens Pl. & Lindbl. ex Tr. et Pl., Wemmanma manquuae Szyszyl., W. pinnata L., W. pubescens H.B.K.and W. rollottii Killip var. rollottii.

Table 18. C hemical composition of suspended soil in the Hymenophyllum lindenii-Bazzania breuteliana "typicum" community (type 3.2.1.1.).

residue on ignition(%) pH H20

conductivity J..LS cm- 1

N03-N N02-N NH4-N total N P04-P

so4 K

Ca

n range mean 6 2-8.4 5.1 5 2.7-4.2 3.3 5 460-1345 728.6 IN WATER SOLUBLE IONS (mg kg-1 dry weight) n mean s. dev.

3 1.8 0.8

4

1.3

4

120

5 5 5 5

223.6 660 446.6 102

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s. dev.

2.9 0.7 366 NUTRIENT CONTENT (mg g-1 dry weight) n mean s. dev.

1 78.5 86.9 230.6 379.9 84.3

6 5 5 5 5

15 0.9 2.6 1.6 2.3

3.3 0.3 1.2 1.2 1.6

83

Epiphyte communi ties of tropical montane rain forests. II

3.2.1.2. Scapania portoricensis-Plagiothecium novo-granatense commu nity Compo sition and structur e (Table 19; Fig. 6)

NR = 20; NTt = 91; NT10%

= 35; SR = 10 %.

The liverwo rt Scapania portoricensis is exclusive togethe r with the moss Plagiothecium novo-granatense. Accomp anying taxa are: Bazzani a breuteliana, B. hookeri, Calypogeia peruviana, Lepicolea pruinosa, Lepidozia spp., Metzgeria Table 19. Composi tion and structure of the Scapania portoricensis-Plagiothecium novo-gra natense commun ity (type 3.2.1.2.); 20 releves. per communi ty no. of taxa, no. of taxa, freq. > 10 % total mosses liverwort s crustose lichens foliose lichens fruticose lichens bryophytc s lichens bryophyte s & lichens flowering plants ferns, excl. filmy ferns filmy ferns 'bryophyt e & lichen layer dry weight (g/dm 2 ) thickness (em)

8 22 0 2

21 45

1 3

0

2

30 2 32

66 6 72

0 l 2

5 3 3

mean

s. dev.

2.9 6.2

3.6 2.8

Fig. 6. The Scapania portoricensis-P lagiothecium novo-.granatense commun ity at a he1ght of 1 m and 2970 m altitude. Scale bar equals 5 em .

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per releve mean no. mean cover of taxa (%) 3.5 18.2 9.7 77.5 0.1 0 0.4 1.9 0.1 0.1 13.2 95.7 0.6 2 13.8 97.7

0.7 l 2.3

84

]. H. D. Wolf

decipiens, Porotrichodendron superbum, Riccardia aberrans, Trichocolea tomentosa and Hymenophyllum lindenii.

Ecology and distribution (Tables 20, 21) The S capania portoricensis-Plagiothecium novo-granatense community is common on tree bases at altitudes over 2740 m. Scapania portoricensis is a good indicator of the upper montane forest in tropical America (FRAHM & GRADSTEIN 1991 ). The community is absent near the forest line at 3670 m due probably to the openness of the forest adjacent to the grassy paramo vegetation.

Table 20. Ecology and distribution of the Scapania portoricensis-Plagiothecium novo-granatense community (type 3.2.1.2.); 20 releves. range

mean/ preference altitude (m) 2970-3510 3370-3510 absolute height (m) 0.1-7 0.9 relative height(%) 0.4-39 4.7 diameter branch/trunk (em) 10.0-70 38 position within the tree (I-V) I-II I 0 inclination bark surface { ) 65-95 80 0 0-290 exposition { ) thickness suspended soil (em) 0.1-25 7.5 5.8-13.7 9.4 (2.8) diffuse site factor n= 19 (s. dev.) phorophytes: Axinaea sp., Brune ilia cf. goudotii Tul., Ocotea cf. guianensis Aubl., Weinmannia mariquitae Szyszyl. and W. pubescens H.B.K. Table 21. Chemical composition of suspended soil in the Scapania portoricensis-Plagiothecium novo-granatense community (type 3.2.1.2.). residue on ignition (%)

pHHzO conductivity f:!S cm-1

N03-N N02-N Nf4-N total N P04-P so4 K Ca

range mean n 4.6 2.5-5.7 13 2.8-3.9 3.2 12 325-710 515.3 12 IN WATER SOLUBLE IONS ~mg kg-1 d!}: weight) mean s. dev. n 12 1.3 1.8 12 0.6 1 81.9 lO 101.1 12 12 12 12

136.5 562.2 264.3 113.9

3.2.1.3. Rhizogonium lindigii community Composition and structure (Table 22) NR = 7; NTt = 33; SR = 24 %.

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52.7 181.3 176.5 42.8

s. dev.

1.3

0.4 104 NUTRIENT CONTENT (mg g-1 d!)l: weight) n mean s. dev.

13 12 12 12 12

14.1 0.75 2.3 0.7 2.9

2.6 0.2 1.3 0.5 2.7

Epiphyte communities of tropical montane rain forests. II

85

The moss Rhizogonium lindigii is exclusive, together with the liverworts Cephalozia crassifolia and Calypogeia cyclostipa and the fern Vittaria remota. Accompanying taxa are: Adelanthus pittieri, Bazzania breuteliana, Calypogeia peruviana, Lepidozia spp. and Hymenophyllum lindenii. The relatively sparseness of the Rhizogonium lindigii community compared to other epiphyte communities in the Upper Montane Rain forest is shown by its low average bryophytic biomass (0.8 g/dm 2).

Table 22. Composition and structure of the Rhizogonium lindigii community (type 3.2.1.3.); 7 releves.

mosses liverworts crustose lichens foliose lichens fruticose lichens

bryophytes lichens bryophytes & lichens flowering plants ferns, excl. filmy ferns filmy ferns

bryophyte & lichen layer dry weight (g/dm2) thickness (em)

no. of taxa, total 7 22 0

0

0

29 0 29 I 2

per releve mean no. of mean cover taxa (%) 1.9 36.9 6.6 54.1 0 0 0 0

0 8.5 0

0 91 0

8.5

91 0.7 3.6 1

I

mean 0.8 2.5

s. dev.

0.9 0.9

Ecology and distribution (Tables 23, 24) The Rhizogonium lindigii community has only been recorded at 2740 and 2970 m, but is expected to occur along a wider altitudinal range in a well-

Table 23. Ecology and distribution of the Rhizogonium lindigii community (type 3.2.1.3.); 7 releves. mean/ preference altitude (m) 2740-2970 2740 absolute height (m) 0.1-1.4 0.7 relative height(%) 0.4-5.6 3.1 diameter branch/trunk (em) 45-70 51.3 position within the tree (1-V) I I inclination. bark surface (0 ) 80-110 95 0 exposition ( ) 20-300 thickness suspended soil (em) <0.1-1 0.4 diffuse site factor n= 7 (s. dev.)

1.1-7.5 3.5 (2.2) phorophytes: Gordonia cf. pubescens Pl. & Lindbl. ex Tr. et Pl. and Weinmannia rollottii Killip var. rollottii. range

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Table 24. Chemical composition of suspended soil in the Rhizogonium lindigii community (type 3.2.1.3.). residue on ignition (%) pHHzO conductivity J..LS cm-1

N03-N NOz-N

NJ4-N total N P04-P so4 K

Ca

range n mean 2-38.4 3 14.6 2.5-3.6 3 3.2 270-380 3 333 IN WATER SOLUBLE IONS (mg kg-1 dry weight) n mean s. dev.

3 1.3 0.4 3 0.4 0.4 3 17.6 81.7 3 3 3 3

96.1 75.2 278 34

88.3 343.3 299.3 89.7

s. dev.

20.6 0.6 56.9 NU1RIENT CONTENT (mg g-1 dry weight) n mean s. dev.

3 3 3 3 3

14.8 0.8 2.6 0.9 2

3.5 0.4 0.4 1.1 I

defined habitat. The community habitually occurs in dark locations which usually do not receive direct precipitation, such as hollows between roots or on the under-side of slanting trees. The inorganic fraction of the suspended soil may be up to 38.4%, suggesting the occurrence of splash. Concentrations of nutrients analysed were low.

3.2.2.1. M ittenothamnium rep tans community Composition and structure (Table 25) NR = 9; NTt =52; SR = 11%.

Exclusive taxa, within the Upper Montane Rain forest communities, are Aneura sp(p?), Lophocolea muricata, M ittenothamnium rep tans (not separated Table 25. Composition and structure of the Mittenothamnium reptans community (type 3.2.2.1.); 9 relevk

mosses foliose liverworts thallose liverworts crustose lichens foliose lichens fruticose lichens bryophytes lichens bryophytes & lichens flowering plants ferns, excl. filmy ferns filmy ferns bryophyte & lichen layer dry weight (g/dm2) thickness (em)

no. of taxa, total 15 26 8 0 1

0 49 1 50 0 0

per relev~ mean no. of mean cover taxa (%) 4.3 34.5 7.6 36.9 3 17.9

0

0

0.1

0.6

0 14.9

0 89.3

0.1 15

0 0.8

2

mean 0.9 4.4

0.6

89.9 0

s. dev.

0.5 2.4

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Epiphyte communities of tropical montane rain forests. II

from Ctenidium malacodes), Plagiochila macrifolia and Taxilejeunea lancifolia.

The moss genus Porotrichum is diagnostic in separating theM ittenothamnium reptans community from ~he Riccardia aberrans community. The t~o c.~mmu­ nities are related by shanng, among others, Symphyogyna brongnzartzz, Hypnella pilifera and Heteroscyphus polyblepharis. Other accompanying taxa are Lophocolea trapezoidea, M etzgeria decipiens, Riccardia spp. and Trichocolea tomentosa of which the latter may attain great abundance. Thallose liverworts are con~picuously present with on average approximately three taxa per releve, covering nearly 18 %.

Ecology and distribution (Tables 26, 27) The tree base Mittenothamnium reptans community has an altitudinal range from 2460 m up to ?970 m. At lower elevations Mittenothamnium reptans is often a canopy species.

Table 26. Ecol~gy and distribution of the Mittenothamnium reptans community (type 3. 2.2.1.); 9 releves. range

mean/ preference altitude (m) 2460-2970 2460-2970 absolute height (m) 0.1-1.0 0.3 relative height (%) 0.6-4.6 1.5 diameter branch/trunk (em) 4.0-50 36 position within the tree (I-V) I I inclination bark surface (0 ) 75-90 84.8 exposition (0 ) 0-320 thickness suspended soil (em) 0.1-2.0 0.5 diffuse site factor n= 7 (s. dev.)

5.6-8.8 6.6 (1.7) phorophytes: Brune ilia occidentalis Cuatr. and Weinmannia rollottii Killip var. rollottii.

Table 27. Chemical composition of a single suspended soil sample (altitude 2970 m) in the Mittenothamnium reptans community (type 3.2.2.1.). residue on ignition (%) pHHzO conductivity jlS cm- 1

N03-N NOz-N

5 4.1

635 IN WATER SOLUBLE IONS

NUTRIENT CONTENT

(mg kg-1 dry weight) <0.1

(mg g-1 dry weight)

3

Nl4-N

327

total N P04-P S04 K Ca

261 463 1402 83

10.7

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0.8 2.8 3.8 2

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3.2.2.2. Riccardia aberrans community Composition and structure (Table 28) NR = 5; NTt = 38; SR = 26%.

Exclusive species are the thallose liverwort Riccardia aberrans, often in great abundance, and the moss Pyrrhobryum mnioides. Accompanying taxa are Trichocolea tomentosa, Telaranea nematodes, Hypnella pili/era and the thallose liverworts Symphyogyna brongniartii, Metzgeria decipiens and Monoclea gottschei. Diagnostic for the Riccardia aberrans community are the high cover values of the (six) thallose liverworts.

Of the thallose liverworts the robust Monoclea gottschei and Symphyogyna brongniartii have a preference for altitudes below 3000 m, Riccardia aberrans is dominant at higher altitudes. It may be necessary to recognize two variants within the Riccardia aberrans community when more releves come available.

Table 28. Composition and structure of the Riccardia aberrans community (type 3.2.2.2.); 5 releves.

mosses foliose liverworts thallose liverworts crustose lichens foliose lichens fruticose lichens bryophytes lichens bryophytes & lichens flowering plants ferns, excl. filmy ferns filmy ferns bryophyte & lichen layer dry weight (g/dm2) thickness (em)

no. of taxa, total 11 18 6

0 0 0 35 0 35 l l l mean 2.2 6.5

per releve mean no. of mean cover taxa (%) 3.6 6.4 5 19 3.4 64.7

0 0 0 12 0 12

0 0 0 90.1 0 90.1 0.2 0.2 0.4

s. dev.

1.2

3.1

Ecology and distribution (Tables 29, 30) The Riccardia aberrans community is recorded from elevations between 2460 and 3670 m on the lowest part of tree trunks, often as an extension of the terrestrial forest floor vegetation. The community is considered the most hygrophytic of all communities described here. In Cuba a similar hygrophytic community has been recorded, growing in shady wet crevices near springs and consisting of thalloid mats of M onoclea gotscheii, Riccardia spp. and Symphyogyna spp., accompanied by Hypopterygium tamariscinum, Trichocolea spp. and various Hookeriaceae (Pocs 1982).

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Epiphyte communities of tropical montane rain forests. II

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Table 29. Ecology and distribution of the Riccardia aberrans community (type 3.2.2.2.); 5 releves. range

mean/ preference 2460-3670 2970-3670 0.1 0.1 0.1-0.7 0.4

altitude (m) absolute height (m) relative height (%) diameter branch/trunk (em) position within the tree (I-V) I inclination bark surface (0 ) exposition (0 ) thickness suspended soil (em) 3.0-20 11.5 diffuse site factor n=? (s. d~v.) .

5.0-9.6 8.6 (2) phorophytes: Brunellw occzdentalzs Cuatr., Clusia aft. multiflora H.B.K., Weinmannia mariquitae Szyszyl. and W. pubescens H.B.K. Table 30. Chemical composition of suspended soil in the Riccardia aberrans community (type 3.2.2.2.). residue on ignition (%) pH HzO I ÂŁ9nductivity JJ.S em-

N03-N NOz-N NH4-N total N P04-P S04 K Ca

n range mean 4 4.3-9.1 6.4 4 3.2-4.5 3.8 4 370-625 485 IN WATER SOLUBLE IONS (mg kg- I dry weight) n mean s. dev.

4 0.3 0.4 4 1.2 1.4 4 102.3 71.6 4 4 4 4

217 422.5 307.5 225

s. dev.

2.4 0.6 110 NUlRIENT CONTENT (mg g-1 dry weight) n s. dev. mean

85.9 78.2 101.1 84.6

4 4 4 4 4

14.7 0.9 3.3 0.7 5.3

4 0.1 0.7 0.2 2.7

4. Communities ofthe Polylepis dwarfforest, 4130 m (Table 1) Confined to the high elevational Polylepis dwarf forest are the Cetrariastrum dubitans-C. equadoriense fruticose lichen community and the Campy/opus pittieri moss community. Exclusive taxa for both communities are Blepharolejeunea securifolia, Frullania tetraptera, Zygodonfragilis, and Parmelina sp. A. 4.1. Cetrariastrum dubitans-C. equaoriense community Composition and structure (Table 31) NR = 15; NTt = 39; NT10% = 28; SR = 18 %.

The endemics Cetrariastrum dubitans and C. equadoriense, are among others exclusive for this community. The community is dominated by fruticose lichens such as Oropogon bicolor, 0. loxensis, Parmelina sp. A, Usnea spp. and the two Cetrariastrum species. Daltonia longifolia, a photophytic species with a wide altitudinal range, is most frequently recorded in this community. The

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liverworts Blepharolejeunea securifolia and Frullania tetraptera are common . . accompanymg spec1es.

Table 31. Composition and structure of the Cetrariastrum dubitans-C. equadoriense community (type 4.1.); 15 releves. per community no. of taxa, no. of taxa, freg . >10% total mosses liverworts crustose lichens foliose lichens fruticose lichens bryophytes lichens bryophytes & lichens flowering plants ferns, excl. filmy ferns filmy ferns bryophyte & lichen layer dry weight (g/dm2) thickness (em)

6

8

7

11 3 11

1 8

6

6

13 15 28

19 20 39

0 0 0

0 0 0

mean 2.5 1.3

s. dev.

2 0.7

per releve mean no. mean cover of taxa (%) 2.2 11 4.2 17.8 0.3 0.2 2.3 19.9 4.3 28 6.4 28.8 6.9 48.1 13.3 76.9 0 0 0 0

0

0

Ecology and distribution (Tables 32, 33) The Cetrariastrum dubitans-C. equadoriense community is restricted to branchlets of Polylepis sericea and Diplostephium violaceum with high diffuse Table 32. Ecology and distribution of the Cetrariastrum dubitans-C. equadoriense community (type 4.1.); 15 relevk range

mean I preference altitude (m) 4130 4130 absolute height (m) 0.9-3.9 2.2 relative height(%) 22.5-97.5 55 diameter branch/trunk (em) 0.7- 10 3.6 position within the tree (1-V) III-V lll-V inclination bark surface (0 ) 10.0-40 26.8 thickness suspended soil (em) <0.1 <0.1 diffuse site factor n= 14 (s. dev) 5-39.8 30.3 (15.8) phorophytes: Diplostephium violaceum Cuatr. and Polylepis sericea Wedd.

Table 33. Nutrient content (mg g- 1 dry weight) of suspended soil in the Cetrariastrum dubitans-C. equadoriense community (type 4.1.). residue on ignition(%) total N P04-P so4 K

Ca

n 3 3 3 3 3 3

mean 9.2 13.1 1.4 4 2.8 5.4

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s. dev.

3.2 3.3 0.4 0.6 0.8 0.8

Epiphyte communities of tropical montane rain forests. II

91

site factors, averaging 30.3 %. No preference for either Polylepis sericea, which has a flaking bark, or Diplostephium violaceum was observed.

The frequently many branched rough lichen thalli, with long rhizines orvilt possibly are morphological adaptations to capture wind-driven fog, which is a regular occurrenc~ in the ?warf forest (pers. observ.). Pendent liverworts such as F rullania peruvzana wh1ch may depend more on regular wetting by rain than by fog (MAGDEFRAU 1983), are noteworthy absent.

Fig. 7. The Campy/opus pittieri community at a height of 4 m and 4130 m altitude. Note the scaling bark of the phorophyte Polylepis sericea. Scale bar equals 5 em.

4.2. Campylopus pittieri community

Composition and structure (Table 34; Fig. 7) NR == 15; NTt == 39; NT10% = 26; SR = 22%.

Exclusive taxa within the dwarf forest communities are the mosses Campylopus pittieri (incl: Campylopus fragilis), Choriso~onti"!m mitten~i, Leptodontium wallisii, Porotrtchondron superbum and t~e foliose lichen Peltzgera pulverulenta. Accompanying species are Blepharolejeunea securifolia, Frullania tetraptera, Zygodon fragilis and Parmelia sp. A. Releves are typically dominated by cushion or turf-forming moss species, covering over 50 %.

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Table 34. Composition and structure of the Campy/opus pittieri community (type 4.2.); 13 releves.

mosses liverworts crustose lichens foliose lichens fruticose lichens bryophytes lichens bryophytes & lichens flowering plants ferns bryophyte & lichen layer dry weight (gldm2) thickness (em)

per community no. of taxa, no. of taxa, freq. >10% total 9 13 8 12

0

3

6

7

3 17 9 26 0 0 mean 9.9 3.1

4 25 14 39 0 0 s. dev.

4.5

per relev~ mean no. mean cover of taxa (%) 4.9 82.2 2.9 3.1 0.2 0.1 1.9 8.7 1

1.1

7.8 3.1 10.9 0 0

85.3

9.9 95.2 0

0

2

Table 3;>. Ecology and distribution of the Campy/opus pittieri community (type 4.2.); 13 releves. range

mean/ preference 4130 1.1 27 7.2

altitude (m) 4130 absolute height (m) 0.1-3 relative height(%) 2.5-75 diameter branch/trunk (em) 2.0-13 position within the tree (1-V) I-III inclination bark surface (0 ) 0-105 29 thickness suspended soil (em) <0.1-1.5 0.5 diffuse site factor n= 9 (s. dev.)

15-44.3 27.4 (10.8) phorophytes: Diplostephium violaceum Cuatr. and Polylepis sericea Wedd.

Table 36. Chemical composition of suspended soil in the Campy/opus pittieri community (type 4.2.). residue on ignition (%) pH H20 conductivity f:l:S cm-1

N03-N NOz-N

Nl4-N total N P04-P so4

range n mean 6.9-15.2 11.7 8 4 4.3-5.1 4.8 4 340-470 406 IN WATER SOLUBLE IONS (mg kg-1 dry weight) n mean s. dev.

4 9.2 8.6 4 1 1.8 4 93.8 13.1 4 4

253.3 347.5

40.5 71

4

306

66.2

K

Ca

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s. dev.

2.9 0.4 59.4 NUTRIENT CONTENT (mg g-1 dry weight) n mean s. dev.

8 8 8 8 8

13 1.2 3.8 1.7 6.4

3.2 0.3 0.6 0.3 1.6

Epiphyte communities of tropical montane rain forests. II

93

Ecology and distribution (Tables 35, 36) The community shows a preference for lower heights in the dwarf forest than the Cetrariastrum dubitans-C. equadoriense community. Diffuse site factors are slightly lower.

The characteristic gregarious growth form of the mosses, building cushions or short turfs, appears not to be an adaptation to maintain foothold on the scaling bark surface of Polylepis trees, since Diplostephium trees carry the same community. Such growth forms are often seen as an adaptation to dry climatic conditions (GrMINGHAM & BrRsE 1957, PRoCTOR 1990).

General discussion Altitudinal zonation The altitudinal di~tribution ~f.20 epiphyte ~omn:tunities shows that a marked change in the species co.mposltl~n the el?iphyuc vegetation occurs between 2130 and 2460 m (Fig. 8). With mcreasmg altitude the (living) biomass of ca~opy bryophytes and lichens also increases rapidly in this belt together with ~he average thic~ness of the bry.op?ytic layer and the amount of'suspended soil accumulated (F1g. 9) .. The van~b1~1ty ?et~een samples per altitude is great, presumably due to d1fferences m mclmatlon of the substrate and historical factors.

The transition zone for epiphytic vegetation coincides with a climate zone that shows a s_ha~p increase in relative hum~dity both in cle~rings and at varius heights w1thm the forest (WoLF 1993, m press). Cloudmess and fog were 0 ommonly observed at altitudes above 2100 m. Thus, the data on epiphytic bryophytes and lichens are in agreement with the hypothesis that fogginess may exert a great in~uence on the boundari~s between zonal ÂŁ?rest formatioJ?-s on tropical mountams (GRUBB 1974). The importance of mmsture content m poikilohydric bryophytes and lichens for net carbon gain is substantiated by ecophysiological research (e. g. PROCTOR 1990, RICHARDSON 1973).

The variation in abundance of fruticose lichens also seems to support GRUBB's (1974) fog~in7ss hypothesis. ~.ruticose lichens in the _l.Jpper Montane Rain forest Diplaswlejeunea pauckertu-Usnea spp. commumty cover 27.4% on average. In con~rast,_ average frutico~e lichens cover in ;he Hypotrachyna imbricatula-Rimelta retzculata commumty from lower altitudes 1s less than 2%. Fruticose lichens are commonly most abundant on the outer branches of trees and the rough, many branched, thalli of Usnea species in particular suggest that they are capable to make maximum use of s.mall a~ounts of ?ew or fog precipitation (PIKE et al. 1975, STONE 1989). Fruttcose hchens are Cited by KERFOOT (1968) as being favoured in their development by California sea fog and Usnea as having a prevalence in Somalia for the mist belt in the North.

However the data presented here may only be regarded as indicative. Next to changes in relat_ive ~umidity along the a_lti~udi_nal gr~dient simultaneous changes in other chmatlc factors such as rad1at1on mtensity, temperature and

ot

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(west

Paramo

Upper Montane Rain forest

j

Lower Montane Rain forest

Tennales

Santa Rosa de Cabal ~-------

1000

~Rio Cauca

""""'"""'""'""'"" outer canopy community

..,,,,;-,,,,,,/. inner/middle canopy community â&#x20AC;˘ ...._ tree base/trunk community

Lowland forest

distance 0

10

20

30

40

50

60

(km)

Fig. 8. The altitudinal range of the epiphyte communities.

rainfall occur. These factors also influence desiccation. The net intensity and frequency of the drying-wettin g cycle will be the result from complex interactions between these factors. The species response to the environmenta l variable 'altitude' is far from understood. In addition characteristics of the substrate, i.e. bark or suspended soil, may influence the establishmen t of epi-

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Epiphyte communities of tropical montane rain forests. II

95

30 25 20 0 0

15

0

10

.

•

•

9 t:J

: 0 0

5

o 0

500

altitude (m) 1000 1500

2000

2500 3000 3500 4000

::: thickness (em) of the bryophytic layer 0 ::: thickness (em) o~ the layer. o~ accumulated suspended soil; values <0.2 omitted 0 • ::: dryweight (g/dm ) of the hvmg bryophytes and lichens Fig. 9. Structural characteristics of the epiphytic vegetation from the inner and middle canopy.

ce I

phytes. The relative importance of various ecological factors and the occurrence of host-specifici~y is the subject of a future paper.

Species turn-over, bwmass and cover values of tree base (0-3 m) and terrestrial bryophytes ha~e been used to recognize altitudinal belts on tropical mountains world-wrde (FRAHM 1990a, GRADSTEIN & FRAHM 1987, VAN REENEN 1987, VAN REENEN & GRADSTEIN 1983, GRADSTEIN et al. 1989).

FRAHM & GRADSTEIN (~ 991) develop~d a scheme of altitudinal zonation. On high equatorial mountams on the mamland they recognized five altitudinal forest belts. Three of these belts occur in the transect where this study was carried out (GRADSTEIN et al. 1989). The boundary between 'lower tropical montane forest' (=Lower Montane Rain forest, sensu GRUBB et al. 1963) and 'upper tropical montane forest' (= Upper Montane Rain forest) in our transect was established at ca. 2200 m. The 2200 m boundary falls within the transition zone for epiphytic vegetation. Thus, the altitudinal zonation based on terrestrial and tree base bryophytes is in agreement with the zonation of this study of canopy species and which included lichens (and vascular plants).

Consequently , the floristic and structural characteristic s of the canopy vegetation may provide parameters for the identification of belts in addition to the characteristic s proposed by FRAHM & GRADSTEIN (1991).

Useful identifiers of the Lower Montane Rain forest in the transect are the pantropicalli chen c;:occo_carpia pell!ta, He~ero.4ermia lu_tescens, Hypotrachyna costaricensis, Rimelta retzculata, Stzcta wezgelu and vanous Parmotrema spe-

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cies. The amount of accumulated epiphytic soil is less than 1 em and the thickness of the bryophytic layer less than 5 em.

Exclusive lichen species of the Upper Montane Rain forest are Hypotrachyna caraccensis, H. laevigata and H. physcioides. Characteristic also is the great abundance of conspicuous fruticose Usnea species in the outer canopy. Other fruticose lichens such as Oropogon bicolor, 0. loxensis and Everniastrum catawbiense commonly occur. The thickness of the bryophytic layer is often over 5 em. The layer of suspended soil in the canopy is generally over 1 em thick and may attain over 15 em on older branches and in tree forks.

Near the forest line the Upper Montane Rain forest grades into Subalpine Rain forest (GRUBB 1974). This forest formation, optimally developed at an altitude of 3670 m where adult trees attain a height of approximately 15 m and where canopy cover is less than 40 %, lacks exclusive epiphyte communities (Fig. 8). Whereas the epiphyte communities in its canopy are identical as in the more dense Upper Montane Rain forest from lower altitudes, the tree base vegetation is characteristic. At the forest floor typical canopy communities from lower altitudes may be found such as the Plagiochila fuscolutea community. Skiophytic tree base communities, e. g. the Hymenophyllum lindeniiBazzania breuteliana "typicum" community and the Scapania portoricensisPlagiothecium novo-granatense community are lacking.

The greater uniformity in the epiphytic vegetation adjacent to the paramo is also shown by the drop in richness of non-vascular epiphytes from 132. taxa at 3510 m to 108 taxa at 3670 m (WoLF in press). Uniformity seems associated with low habitat (beta-)diversity: diurnal courses of temperature and relative humidity in the understory are similar to a clearing nearby (WOLF 1993).

Moreover, diffuse site factors in the understory from 2970 m up to 3510 m and at 3670 m were 8.5% (s. dev. 2.9) and 15.8% (s. dev. 3.4), on average.

The Polylepis dwarf forest at 4310 m is structurally different from the forest at 3670 m (WoLF 1993). Its two epiphyte communities, the Cetrariastrurn dubitans-C. equadoriense community and the Campylopus pittieri community ~~~~. '

Bark acidity Bark acidity values are single measurements per tree species only and do not take into account possible variation with season, age, or within the tree (Table 37).

All bark samples are acidic in reaction. Drying of bark fragments prior to acidity detection hardly influences the outcome, giving values of about 0.3 pH units lower (BISANG 1985).

Most values fall in the range between pH 4.5 and 5.5. An Axinaea (Melastomataceae) tree showed the lowest value of pH 3.5 and a Talauma (Magnoliaceae) tree the highest value of 6.2. Similar pH values have been reported from montane Dipterocarp forest on Mt. Kinabalu, Malaysia and from Amazonian lowlands (FRAHM 1990b, LISBOA 1976).

In temperate areas a correlation between bark acidity and the floristic com-

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97

Table 3Z Bark acidity values of various phorophyt es.

Phorophyte Aegiphyla bogotensis (Spreng.) Mold.

Axinaea sp. (Melastomataceae) Brune Ilia cf goudotii Tul. Brunellia occidentalis Cuatr.

Chrysophyllum cf auratum Miq. Cinchona sp. (Rubiaceae) Clusia aff. multiflora H.B.K. Dendropanax cf macrocarpum Cuatr.

Diplostephium violaceum Cuatr..

Escallonia myrtilloides Lf var. myrtilloides Ficus cf velutina Willd.

Hyeronima scabrida (Tull.) MU!l. Arg. Gordonia cf pubescens Pl. & Lmdbl. ex Tr. et Pl. Lauraceae Lauraceae Lauraceae Lauraceae Machaerium capote Triana ex Dugand Matisia sp.

Nectandra cf globosa (Aub1et.) Mez. Ocotea cf guianensis Aubl.

Polylepis sericea Wedd.

Rosaceae (cf) Senna pistaciifolia (H.B.K.) Irwin & Barneby Solanum inopinum Ewan Talauma sp. (Magnoliaceae) Tetrorchidium boyacanum Croiz.

Weinmannia mariquitae Szyszyl.

Weinmannia rollottii Killip var. rollottii Weinmannia pubescens H.B.K.

altitude (m a.s.l.)

2130

3510

2970 2460 1500 1500 2970 1725

4130 3670 1500

1980 2740 1210 1210 1725 1980 1000 1210 1980 3190 4130 1725

2130 2130 1725 2740 3370 2970 3370

pH KC1

5.8 3.5 4.5 4.7

5.0 4.1 4.6 5.6 5.4 4.6 5.5 4.5 4.3 5.0 5.4 4.4 5.4 5.8 5.3 4.4 4.7 3.8 5.0 5.7 5.3 6.2 5.4 4.4 5.0 4.3

5.3

3.2 3.9 4.2 4.7 3.8 4.1 5.0 5.0 4.2 4.9 4.1 3.6 4.4 5.0 3.7 4.7 5.5 5.1

4.1 4.0 3.4 4.5 5.2 5.0 6.2 4.7 4.0 4.6 3.8

position of epiphytic vegetation has been documen ted (e. g. BARKMAN 1958, N rMIS 1985). In the tropics bark acidity measurements are few (FRAHM 1990b). In th~ transect th.e bark hydrogen ion c.oncentration is less likely to determin e eptphyte establishment above approximately 2500 m due to the accumulati on of humus on the bark surface and small amounts of stem flow (VENEKLAAS & VANEK 1990).

Chemical composition of organic suspended soils The nutrient content of suspended soils from tree base and canopy communities reveal patterns along the altitudinal gradient and within host trees (Fig. 10). At higher altitudes soils are more organic and nutrient concentrations are lower. Tree base soils are richer than canopy soils at altitudes below 2000 m. In contrast at altitudes above 2550 m the tree base soils were not richer than in the can;py, with the excepti.on of ni~ro~e.n. . . .

The significantly lower restdues on tgmuon at htgher elevations conftrm observations made in El Salvador (KLINGE 1963). In the study area, towns, roads

7 Phytocoen ologia 22

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I

Upper Montane rain forest 'typicum' community alt. range: 2550 m - 3670 m

Cheilolejeunea trifaria Schlotheimia angustifolia community alt. range: 1250 m - 1980 m residue after ignition (%)

9. 95 total N 4.4 17.3 _ _ _ _...=::::..:..:.______ 11.5 t

t

canopy

I I

,,In I

P-1'04

2.84

K

f

s~

5.1

n~

2.1

t s.I3

3.39 t t :

t I

I 1

•

+ 11.51

tree base

6.8 4.5

I I

• 1.75 18.3 -

I

Ca

I

1

2.86 + I 2.27.

S04

canopy

f ' I I I

:

I

I

r

1.04 t

I

1

I

0.~ ~

K

P-P04

tree base

I

0.75 • .. 14.1

_!.otal N -

• residue after ignition (%) • 13.4 ---===-==:....:E=:::.::~!......- 4.6 Bryopteris filicina community alt. range: 121 0 m - 1980 m

Scapania portoricensis Plagiothecium novogranatense community alt. range: 2740 m - 351 0 m

A) NUTRIENT CONTENT (mg/g) Cheilolejeunea trifaria Schlotheimia angustifolia community alt. range: 1250 m - 1980 m

Upper Montane rain forest 'typicum' community alt. range: 2550 m - 3670 m

pH

5.1 -----""N'"'-N""'H"'"'4,-------- 3.5 280.1 60.1 P-1'04 190.8·-------i<--------177.5 t 1264.8 ... 519.3 514.3 .. -- ~~--- .. 650.11 2 393.1 Ca

t

canopy

t

canopy

]"! :

ea

:

113.H

I

S04 562.2 ~ K 264.5

levels of significance

tree base

~

P-1'04 136.5

p$0.0005 p$0.005 p$0.05 - - - - - - - - - - - - - - - n.s.

N-NH4 81.9 t pH 3.2

Scapania portoricensis Plagiothecium novogranatense community alt. range: 2740 m- 3510 m

B) IN WATER SOLUBLE IONS (mglkg) Fig. 10. Chemical composition of suspended soil of two canopy and two tree base communities. Levels of differences in significance, using Student's t-test between the mean values on the diagonals are indicated. A Kolmogorof-Smirnov test for normality gave no significantly different distribution from normality for all variables; p > 0.1.

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and arable lands are mainly found at lower altitudes. Hence, dust input below ca. 2000 m is expected to be higher. Moreover, the recorded increase of epiphytic soil accumu.lation with al~itude, should result in a decrease of inorganic matter concentratiOns at equal mput rates.

Lower macronutr ient concentrat ions at higher elevations may reflect a general decrease in nutrient concentrat ions of main suspended soil building componen ts with increasing altitude, as reported for mature foliage and litterfall (GRUBB 1977, VENEKLAAS 1991, VITOUSEK 1984). Differences in the decomposi tion sub-system (catabolism, communiti on and leaching) at various altitudes due to differences in th~ decompose r community , the resource quality and t~e ph~sico-chemical envtronme nt, will also have their response on ornic sotl quahty (e. g. SwiFT et al. 1979). ga Concentra tions o~ soluble N, Ca and K also are ~ighest in the canopy of the Lower Montan: Ram forest, as ar~ pH values (Ftg. 10 B). Soluble nutrient concentrat ions m the can<;>P.Y ~ay mfluence tree base vegetation through an ffect on throughfal l preClpttatw n and stem flow quality. Amounts of stem ~ow are small in the Upper Mon_ta~e Rain forests in the transect (VENEKLAAS & VAN EK 199~). ~~spended sods ~n the canopy of the Upper Montane Rain forest are. not stgn~ftcantly poorer m soluble phosphate than in the canopy at }ower aln~udes. Smc~ phosphate concentrat ions of throughfall water were lower at htgher e~evatwns (VENEKLAAS 1990), possibly in the Upper Montane Rain forest leachm~ of phosphate frc;>m living leaves, another potential source, is less. A more raptd uptake of avatl~ble phosphate within the canopy itself offers another explanatiOn. Less leachmg and a rapid uptake within the canopy are consistent wtth t~? more observed processes typical for higher altitudes ¡ ndicating a more eff1c1ent use of phosphate: lower nutrient cycling rates and higher phosphate reallocation (V ENEKLAAS 1991 ).

In the Lower Montane Rain forest tree base suspended soil contains more minerals and is richer in macronutr ients than canopy soil (Fig. 10 A). Splash erosion, highest between altitudes of 1200 and 2500 min the transect (VIs in press) may be of importance . Total content of S and N is not significantly highe~ at tree bases. Sulphate may have primarily a canopy origin. Aerial deposition was shown to be high due to volcanic activity of the nearby 'Nevada del Ruiz' (VENEKLAAS 1990). N values are difficult to interpret, since microfauna! skeletons, present in great abundance (pers. observ. and M.J. SANABRIA, unublished) are included. p Nutrient supply possibly is one of the principal factors limiting the growth of montane forest (GRUBB 1977) and recent studies from Jamaica and Colombia suggest a limited availability of N and P for some forest trees or forest types (TANNER et al. 1990, VENEKLAAS 1990). However, substratum quality is less likely to affect eRiphytic ectohydric bryophytes. and lichens. Both are often considered to b~ mdependen~ from substra_n~m mputs for the_bulk of their nutrients and thetr water reqmremen ts, obtammg both from aenal sources (e. g. ANDERSON et al. 1955, _BRO,W:N ~ BATES 1990). Even though this assumptio n is likely to be an ove.mmphftcat~~n (BROWN & BATES 19?0), apparent correlation s between chemtcal composltlo n of the suspended sod and floristic compositio n of epiphyte communiti es may well be accidental.

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Free-moving animals are likely to show a response to characteristics of the suspended soil. On seven occasions earthworms were collected from soils in the middle canopy of the Upper Montane Rain forest (2550-3370 m), belonging to two species of Glossoscolecidae. Andiodrilus cabalensis is described as a new species to science and one specimen was classified as a juvenile M atiodrilus sp. (G. RIGHI in press). The distribution of earthworms in nature is dependent on a number of factors, among them moisture availability, calcium content of the soil and acidity (SATCHELL 1956). The apparent absence of canopy worms from altitudes below 2550 m may be related to their susceptibility to desiccation. More peculiar, tree dwelling worms were never observed at tree bases or on the oldest branches in the inner canopy. Soil acidity may determine worm distribution patterns. At collection sites pH values were never lower than 3.8 and 4.1 (s. dev. 0.3) on average. In contrast, tree base and inner canopy soils often are more acidic. For example, pH values recorded in the Scapania portoricensis-Plagiothecium novo-granatense tree base community and in the Plagiochila fusccolutea inner canopy community were 3.2 and 3.4, on average.

Experiments with terrestrial earthworms confirm that species actively avoid soil pH values lower than 4 (LAVERACK 1960).

Conclusions Bark type restricted sampling of epiphytic vegetation on 59 host trees in the transect yielded 507 taxa of cryptogams (34 ferns, 187liverworts, 108 mosses, 110 foliose lichens, 30 fruticose lichens and 38 crustose lichens) and 74 taxa of phanerogams. Cryptogams also dominated the vegetation in terms of abundance.

Epiphytic vegetation was highly variable. A particular combination of species was never recorded twice and the cover values of many species fell within a wide range, even within the same community type. The mean similarity ratio, a coefficient of similarity, between releves of a single community type was low, nearly 16 o/o on average.

The phytosociological analysis revealed 22 communities. Particular communities occurred at particular locations within the host tree and in the altitudinal transect.

In the transect a great change in the floristic composition and the structure of epiphytic vegetation was recorded between ca. 2130 and 2460 m. This transition zone coincided with a zone where air humidity increased rapidly with increasing elevation, suggesting that humidity is largely responsible for the altitudinal distribution of species.

A great abundance of Usnea species in the outer canopy of the Upper Montane Rain forest is one of the parameters proposed which may be helpful for the identification of altitudinal forest formations.

Several epiphytic bryophytes and lichens of the Colombian Lower Montane Rain forests appear to occur in the warmer Lowland Rain forest of Guyana in sites which receive more radition energy. These field observations corroborate the results of experimental laboratory studies on the tropical lichen Dictyone-

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Epiphyte communities of tropical montane rain forests. II

101

ema glabratum and suggest that these species require more radiation energy at higher temperatures to attain net photosynthesis.

The chemical composition of suspended soils is highly variable. For example, pH values range from 7.1 in the Hypopterygium tamariscinum community to 3.2 on ~verage in the Scapania portoricensis-Plagiothecium novo-granatense commumty.

Highly significant differences are recorded between suspended soils of the Lower Montane Rain forest and the Upper Montane Rain forest. The latter soils are more organic and generally show lower nutrient contents.

In the Lower Montane Rain forest nutrient concentrations at the tree base are higher than in the canopy. The oc.currence of splash erosion offers an explanation. In ~he Upper Monta~e Ram fores~ the trend was opposite.

An epiphytiC earthworm .sRec1es, new to SCI~nce, appears to be an obligate middle canopy dweller, av01dmg suspended soils with pH values lower than 3 .8 , as commonly present on the bases of trees and in the inner canopy.

Acknowledgements. The assistance with identification is gratefully acknowledged of Drs. B H. ALLEN, A. APTROOT, s. P. CHURCHILL, J.-P. FRAHM, D. GRIFFIN III, R. GROLLE, R. MoBERG, J. Vf...NA, D. H. VITT, K. YAMADA and R.H. ZANDER. As to vascular plants the aid of Drs. J:T. Ar~:ooD, K.l!. KRAMER, H.E. LuTHER, M.T. MuRILLO and J.O. RANGEL-CH. IS appreciated. Special thanks for the identification of numerous collections is forwarded to ~r. H. SIPMAN (lichens), Drs. G.B.A. VAN REENEN (mosses), Dr. S.R. GRADSTEIN (liverworts) and the late Dr. H. INOUE (Plagiochila). I thank Dr. S.R. GRADSTEIN and Dr. A.~. CLEEF f?r t?eir comments on earlier versions of the manuscript and the staff of t~e Instltuto de. S:1enc1as Naturales (COL), Bogota, for their cooperation while visiting their research fac1hty. Drs. ].G.B. OosTERMEIJER jr. provided the illustrations of the communities. Corrections in the English text were made by LORI MATTA. Finally, this study would not have been possible without the tireless and innovative field assistance of J. KLoMP. The study was supported by grant W84-236 of the Netherlands Foundation for the Advancement of Tropical Research.

Summary. In a series of two papers, epiphytic vegetation along an altitudinal transect in the Central Cordillera of Colombia has been described, using the Braun-Blanquet approach in phytosociology. At each of 15 sites, located at altitudinal intervals of approximately 200 meters the vegetation on four canopy trees was sampled. Host trees with a scaling, hard or smooth bark were avoided as were neighbouring trees. Sampling included canopy sampling for which rope-climbing techniques were used.

Twenty-two communities are newly described and their hierarchical position is indicated.

For each community the location within the host tree, the altitudinal range and the chemical composition of suspended soil, when available, is given. As a parameterforthe (diffuse) light conditions at sites, diffuse site factors are estimated from hemispherical photographs.

Of the environmental variables recorded, the complex variables 'location within the host tree' and 'altitude' appear to exert the greatest influence on epiphytic vegetation.

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J.H.D. Wolf

Literature Anderson, L.E. & Bourdeau, P.F. (1955): Water relations in two species of terrestrial mosses.

-Ecology 36: 206-212.

Barkman, J.J. (1958): Phytosociology and ecology of cryptogamic epiphytes.- 628 p. Van Gorcum, Assen.

Bisang, I. (1985): Zur Verbreitung und Okologie der Frullania-Arten der Schweiz.- Botanica Helvetica 95: 247-278.

Brown, D.H. & Bates, J.W. (1990): Bryophytes and nutrient cycling.- Bot. J. Linn. Soc. 104: 129-147.

Deguchi, H. & Inoue, H. (1982): Lake moss-balls found in Tierra del Fuego and Brunswick Peninsula, Southern South America. - Bull. Natn. Sci. Mus., Tokyo, Ser. B, 8 (4): 145-150.

Frahm, J.-P. (1990a): The altitudinal zonation ofbryophytes on Mt. Kinabalu.- Nova Hedwigia 51: 133-149.

- (1990b ): The ecology of epiphytic bryophytes on Mt. Kinabalu, Sabah (Malaysia). -Nova Hedwigia 51: 121-132.

Frahm, J.-P. & Gradstein, S.R. (1991): An altitudinal zonation of tropical rain forests using bryophytes. - ]. Biogeography 18: 669-678.

Gimingham, C. H. & Birse, E.M. (1957): Ecological studies on growth-form in bryophytes; I. correlations between growth-form and habitat. - J. Ecol. 45: 533-545.

Gradstein, S.R. (1992): The vanishing tropical rain forest as an environment for bryophytes and lichens.- In: Bates,J.W. & Farmer, A.R. (eds.), Bryophytes and Lichens in a Changing Environment, p. 232-256. - Oxford University Press, Oxford.

Gradstein, S.R. & Frahm, J.-P. (1987): Die floristische Hohengliederung der Moose en dang des BRYOTROP-Trans ektes in NO Peru.- Nova Hedwigia 88: 105-113.

Gradstein, S.R., van Reenen, G.B.A. & Griffin III, D. (1989): Species richness and origin of the bryophyte flora of the Colombian Andes.- Acta Bot. Neerl. 38: 439-4~8.

Grubb, P.J. (1974): Factors controlling the distribution of forest-types on tropical mountains: new facts and a new perspective.- In: Flenley, J.R. (ed.), Altitudinal zonation in Malesia.- Univ. of Hull. Misc. Ser. 16: 13-45.

- (1977): Control of forest growth and distribution on wet tropical mountains: with special reference to mineral nutrition. - Ann. Rev. Ecol. Syst. 8: 83-107.

Grubb, P.J., Lloyd, J.R., Pennington, T.D. & Whitmore, T.C. (1963): A comparison of Montane and Lowland Rain forest in Ecuador. I. The forest structure, physiognomy and floristics. - J. Ecol. 51: 567-601.

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

Hofstede, R.M., Wolf, J.H.D. & Benzing, D.H. (1993): Epiphytic mass and nutrient status of an Upper Montane Rain forest.- Selbyana 14: 3-11.

Kerfoot, 0. (1968): Mist precipitation of vegetation.- Forestry Abstracts 29: 8-20.

Klinge, H. (1963): Ober Epiphytenhumus aus El Salvador, Zentralamerika.- Pedobiologia 2: 102-108.

Laverack, M.S. (1960): Tactile and chemical perception in earthworms. II. Responses to acid pH solutions. - Comp. Biochem. Physiol. 2: 22-34.

Lis boa, R.C. (1976): Estudos sobre a vegetacao das campinas amazonicas. V. Brioecologia de una campina amazonica.- Acta Amazonica 6: 171-191.

Miigdefrau, K. (1983): The bryophyte vegetation of the forests and paramos of Venezuela and Colombia.- Nova Hedwigia 38: 1-63.

Nimis, P.L. (1985): Phytogeography and ecology of epiphytic lichens at the southern rim of the clay belt (N-Ontario, Canada).- The Bryologist 88: 315-324.

Pike, L.E., Denison, W.C., Tracy, D.M., Sherwood, M.A. & Rhoades, F.M. (1975): Floristic survey of epiphytic lichens and bryophytes growing on old-growth conifers in western Oregon. - The Bryologist 78: 389-402.

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P6cs, T. (1982): Tropical forest bryophytes.- In: Smith, A.J.E. (ed.), Bryophyte ecology, p.

59-104.- Chapman and Hall, London-New York.

Proctor, M.C.F. (1990): The physiological basis of bryophyte production.- Bot. J. Linn.

Soc 104:61-77.

Richards P. W. (1984): The ecology of tropical forest bryophytes. -In: Schuster, R.M. (ed. ), New :Uanual of bryology 2, 1233-1270.- The Hattori Botanical Laboratory, Nichinan.

Richardson, D.H.S. (1973): Photosynthesis and carbohydrate movement.- In: Ahmadjian, v. & Hale, M.E. (eds.), ~he lichens, p. 249-288.- Academic Press, New York-London.

Righi, G. (in press): Colombian earthwo~ms.- In: van der Hammen, T. & dos Santos, A. G. (eds.), La Cordillera Central Colomb1ana, Tr~nsecto Parque los Nevados (Cuarta Parte). _ Stud. Trop. And. Ecosyst. 4. Cramer, Berlm-Stuttgart.

Satchell, J.E. (1956): Some aspects of earthworm ecology.- In: Devan McE, D.K. (ed.), Soil Zoology, p. 180-20~.- Butterwo~ths, London.

1989): Ep1phyte successiOn on Quercus garryana branches in the Willamette Stone, D .F. ( Th B I . e ryo og1st 92: 81-94. . . . .

Valley of western Oregon.Swift, M.J., Heal, O.W. & Anderson,J..M. ~1 ?79): ~eco.mposmon m terrestnal ecosystems.

Studies in Ecology 5.- Blackwell Sc1entlf1c Publications, Oxford-London-EdinburghMelbourne.

Tanner, E.V.J., Kapos, '-:"·: Fr~skos, S., H~aley, J.R. & Theobald, A.M. (1990): Nitrogen and phosphorus fertilization of Jamaican montane forest trees. - J. Trop. Ecol. 6: 231-238.

Van Leerdam, A., Zagt, R.J · & Venek!aas, E.J · (1990): The distribution of epiphyte growthforms in the canopy of a Col?m~1an cloud-forest.- Vegetatio 87: 59-71.

Van Reen~n, G.B.A. (1987): Alu~ud1.nal b~ophyte zonation in the Andes of Colombia: a prelimmary report.- Symp~s1a B10log1ca Hungarica 35: 631-637.

Van Reenen, G.B.A: & ~radste1?, S.R. (1983): A transect analysis of the bryophyte vegetation along an alt1tudmal gradient on the S1erra Nevada de Santa Marta, Colombia. - Acta Bot. Neerl. 32: 163-175.

Veneklaas, E.J. (1990): Nutrient fluxes in bulk precipitation and throughfall in two montane tropical rain forests, Colombia.-]. Ecol. 78: 974-992. _ (1991 ): Litterfall and nutrient fluxes in two montane tropical rain forests, Colombia. - J. Trop. Ecol. 7: 319-336.

Veneklaas, E.J. & van Ek, R. (1990): Rainfall interception in two tropical montane rain forests, Colombia.- Hydrological Processes 4: 311-326.

Vitousek, P.M. (1984): Litterfall, nutrient cycling, and nutrient limitation in tropical forests. - Ecology 65: 285-290.

Westhoff V. & van der Maarel, E. (1973): The Braun-Blanquet approach. -In: Whittaker, R H. (ed.), Handbook of Vegetation Science, part V, p. 617-726.- Junk, The Hague.

Wolf. J.H.D. (1993): Epiphyte communities of tropical montane rain forests in the northern Andes. I. Lower montane communities. - Phytocoenologia 22: 1-52. _ (in press): Dive;sitypatterns and biomass of epiphyti~ bryophytes and lichens along an altitudinal grad1ent m the northern Andes. - Ann. M1ssoun Bot. Gard.

Address of the author: Jan H.D. WoLF, University of Amsterdam, Hugo de Vries-Laboratory, Kruislaan 318, NL-1098 SM Amsterdam, The Netherlands.

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Epiphyte communities of tropical montane rain forests. II

J.H.D. Wolf

value(= average cover value when present) and maxim um cover value are given. The'#' refer to collection numbers.

Table 1. Synoptic table of the epiphyte communities in the Upper Montane Rain forest. In addltlon to the presence degrees of species, their minimal cover value, characteristic cover COMMUNITY 3

NUMBER OF RELEVES IN THE COMMUNITY MEAN RELEVE AREA (dm2) TOTAL NUMBER OF TAXA IN THE COMMUNITY

3. Communities of the Upper Montane Rajn forest Trichocolea tomentosa (Sw.) Gott.

Lepicolea pruinosa (Tayl.) Spruce Lepidozia spp.

Lophocolea trapezoidea Montagne Bazzania hookeri (Lindenb.) Trevis.

Adelanthus pittieri (Step h.) Grolle Campylopus asperifolius Mitt.

Riccardia spp.

Plagiochila bursata (Desv.) Lindenb.

Sematophyllum insularum (Sull.) Mitt.

3 1 Canopy communities of the Upper Montane Rajn forest Jamesoniella rubricaulis (Nees) Grolle s.lat.

Prionodon fusco-lutescens Hampe Trachylejeunea dominicencis Step h.

Frullania convoluta Lindenb. & Hampe Leptoscyphus jackii (Steph.) Grelle Plagiochila echinella Gott. group Dictyonema glabratum (Spreng.) D. Hawksw. (=Cora pavonia Fr.) Hypotrachyna prolongata (Kurok.) Hale Hypotrachyna physcioides (Nyl.) Hale Leptoscyphus porphyrius (Nees) Grolle Hypotrachyna bogotensis (Vain.) Hale Bazzania longistipula (Lindenb.) Trevis.

Drepanolejeunea spp.

Hypotrachyna laevigata ((Smith) Hale) -group Anoplolejeunea conferta (Meissn.) Step h.

Daltonia longifolia Tayl.

Cheilolejeunea spp.

Diogenesia cf. tetranda {A. C. Smith) Sleumer Hypotrachyna imbricatula (Zahlbr.) Hale Hypotrachyna caraccensis (Tayl.) Hale Frullania sphaerocephala Spruce Herbertus pensilis (Tayl.) Spruce Hypotrachyna reducens (Nyl.) Hale Grammitis apiculata (Kl.) Seymour Herbertus juniperoideus (Sw.) Grolle Hymenophyllum sp. A (# 750) Squamidium leucotrichum (Tayl.) Broth.

Herbertus subdentatus (Step h.) Fulf.

25 7.2 107 pres cover(%) Ofo - av + 4 8

32 6.2 133 pres. cover(%) 0 /o - av +

<I <I <I

9 <I I 63 <I 2 <I <I <I 56 <I 2 3 <I <I 9 <I <I

34 6 40 <I I 5 47 3 6

12 28 4 32 8 4 36 24 36 12 64 60 64 12 24 8

<I <I <I 5 5 5 <I <I <I I 3 5 I I I <I 3 10 <I <I I I I I <I 4 25 <I <I I <I 6 45 <I <I I <I <I I <I 4 10 <I I 5 <I <I I <I 2 S

8 4

<I 5

4 45 2 10 3 20

I

5 iS

10 <I I 8 <I <I 44 <I 5 48 <I 19

5 <I 45 80

<I <I <I 13 <I <I I

44 <I 11 19 <I 5 34 <I <I 44 <I 3 47 <I 13 91 I 19 16 <I 5 6 I 6 69 <I 13 69 <I 2 38 <I 6 13 <I 9 56 <I <I 59 <I 4 56 <I <I 3 I I 19 <I I 3 2 2 2 2 2 28 <I 4

16 <I 6 15 34 4 2 2 2 3 34 24 I 4 6 9 6

2 53 <I 13 39 <I 10 26 <I <I 23 <I I 21 <I

<I 9 60 <I <I <I <I 2 10 <I <I <I

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Epiphyte communities of tropical montane rain forests . II

J.H.D. Wolf

56

57

Table 1. (cont.)

COMMUNITY 3

Macromitrium trachypodium Mitt.

Amphilejeunea viridissima Schust.

Harpalejeunea ancistrodes (Spruce) Schiffn.

Anzia leucobates (Nyl.) Mull. Arg. Cladonia cf. ochrochlora Flk. Heterodermia vulgaris (Vain.) Follm. & Redon Frullania apiculata (Reinw. et al.) Nees Zygodon reinwardtii (Hornsch.) Braun Epidendrum sp. B (11609) Sticta cf. damaecornis (Sw.) Ach. Elaphoglossum aff. lindigii (Karst.) Moore Grammitis meridensis (KJ.) Seymour Everniastrum fragile Sipm.

Epidendrum spp. elleantoides group Elaphoglossum sp. A (II 826) Herbertus acanthelius Spruce Hypotrachyna microblasta (Vain.) Hale Lepyrodon tomentosus (Hook.) Mitt.

Hypotrachyna gigas (Kurok.) Hale Everniastrum cirrhatum (Fr.) Hale Pachyphyllum sp. A (II 751) Zygodon squarrosus (fayl.) C. Mull.

Menegazzia sp. A(ll 993) Leptotheca boliviana Herz.

Syzygiella integerrima Steph.

Blepharolejeunea incongrua (Lindenb. & Gott.) v. Slag. & Kruijt Anastrophyllum auritum (Lehm.) Step h.

Dictyonema spp.

Anastrophyllum leucocephalum (Tayl.) Step h.

Hastifera sp.

Brachiolejeunea laxifolia (fayl.) Schiffn.

Lepidopilum haplociliatum (C.M.) Par (incl. L. muelleri) Aureolejeunea fulva Schust.

Adelanthus crossii Spruce Ceratolejeunea grandiloba Jack. & Steph.

Plagiochila sp. B (II 1354) Elaphoglossum sp. A (II 771) Sematophyllum cuspidiferum Mitt.

Leptodontium viticulosoides var. panamense (Lor.) Zander Cyrtolejeunea holostipa (Spruce) Evans 3.1 1 Outer canogy !wig:let communities Oropogon bicolor Essl.

Oropogon loxensis (Fee) Th. Fr. Aureolejeunea paramicola (Herz.) Schust.

Everniastrum catawbiense (Degel.) Hale

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Ep iphyte communi t ies o f tropi ca l mo ntane ra in fo rests . II

j.H.D. Wolf

58

59

Table 1. (cont. ) COMMUNITY 3

Drepanolejeunea andina Herzog Frullania peruviana Gott.

AnaslTophyllum tubulosum (Nees) Grolle Plagiochila sp. C (If 1367) Pertusaria spp.

31.11 Djplasjolejeunea payckertij - Usnea spp. community Diplasiolejeunea pauckertii (Nees) Steph.

Usneaspp.

Graphidaceae Ramalina sp. A (# 793) Frullania cf. paradoxa Lehm & Lindenb.

Colura naumannii (Schiffn.) Steph.

Colura tenuicornis Steph.

Ramalina cochlearis Zahlbr.

3 1 2. Omphalanthus fj!jforrojs commynjty Omphalanthus filiformis (Sw.) Nees (incl. 0 . platycoleus Herz.)

Frullania brasiliensis Raddi Macromitrium guatemalense C.Miiii.M. longifolium Adelanthus decipiens (Hook.) Mitt.

Plagiochila corniculata Dum. Lejeunea sp. A (# 572) Taxilejeunea pterigonia (L. & L.) Schiffn.

Leptoscyphus amphibolius (Nees) Grolle Omphalanthus ovalis (Lindenb. & Gott.) Gradst.

Lejeunea reflexistipula (Lehm. & Lindenb.) Step h.

Plagiochila dominicensis Tayl.

Pireella cavifolia (Card. & Herz.) Card.

31 3. Upper Montane Rajn forest canopy" typjcym" commynjty AnaslTophyllum piligerum (Nees) Steph.

31 .4 Pla~jochjla fuscolytea commynjty Plagiochila fuscolutea Tayl.

3 2. Tree base commynjlies of the Upper Montane Rajn forest Calypogeia peruviana Nees & Mont.

Telaranea nematodes (Gott. ex Arnst.) Howe Lophocolea aff. connata (Sw.) Nees Riccardia andina (Spruce) Herz.

Plagiochila esmeraldana Step h.

Prionolejeunea spp.

Racopilum tomentosum (Hedw.) Brid.

Porothamnium cf. comosum Hymenophyllum sp. B (# 588) Leskeodon cubensis (Mitt.) Ther.

Thuidium delicatulum (Hedw.) B.S.G. var. peruvianum Cyclolejeunea accedens (Gott.) Evans Syrrhopodon lycopodioides (Brid.) C. Miill.

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Epiph yte co mmuni t ies of tropical montane rain forests. II

J. H. D. Wolf

60

61

Table 1. (cont.)

COMMUNITY 3

3 2 1 Hymenophylly m ljndenjj - Bazzanja breyteliana commynjties Hymenophyllum lindenii Hook.

Bazzania breuteliana (Lindenb. & Gott.) Trevis.

3 2.1.2 5capanja portoricensjs- Plagjothedum noyo-I:'J'anatense community Scapania portoricensis Hampe & Gott.

Plagiothecium novo-grana tense (Hampe) Mitt.

Porotrichodendron superbum (fayl.) Broth.

Blepharostoma trichophy!!a (L.) Dum. Riccardia ciliolata (Spruce) Gradst.

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Rhizogonium lindigii (Hampe) Mitt.

Cephalozia crassifolia (Lindenberg & Gottsche) Fulford Calypogeia cyclostipa (Spruce) Step h.

Vittaria remota Fee

2

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Symphyogyna brongniartii Mont.

Hypnel!a pilifera (Hook. & Wils.) Jaeg.

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Lophocolea bidentata (L.) Dum. 3 <I <I <:1 Cyclodictyon rubrisetum (Mitt.) Kuntze Monoclea gottschei Lindh.

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7 <I <I I Taxilejeunea lancifo!ia Step h.

Lophocolea muricata (Lehman) Nees Plagiochila macrifolia Tayl.

Aneuraspp.

10 <I I 5 Taxilejeunea sulphurea (Lehm. & Lindenb.) Schiffn.

3.2 2 2. Riccardja aberrans community Riccardia aberrans (Steph.) Gradst.

Pyrrhobryum mnioides (Hook.) Manuel Plagiochila ruti!ans Lindenb.

4 Commynjties of the Polylepis dwarf forest Zygodon fragili s Robins Blepharolejeunea securifolia (Step h.) Schust.

4 <I <I <I 10 <1 I 3 Frul!ania tetraptera Nees & Mont.

28 <1 <1 I 3 <1 <I <1 2 <I <I <1 Metzgeria metaensis Kuw. Parmelina sp. A (H 1185) Leptogium mandonii Jorg.

2 <I <I <I Bryaceae sp. A (H 1180) Plagiochila sp. A (H 1164) Tortula andicola Mont.

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E piphyte communities of tropical montane rain fores ts. II

].H. D. Wolf

63

Table 1. (cont.)

COMMUNITY 3

Cetrariastrum dubitans Sipm.

4 2 2 2 Cetrariastrum equadoriense (Sant.) Sipm.

Leptodontium cf. viticulosoides (P.-Beauv.) Wijk & Marg.

Hypotrachyna sinuosa (Smith) Hale 28 <I <I 2 Parmeliella nigrocincta (Mont.) Mull. Arg. Hypotrachyna boquetensis (Hale) Hale 4 2 Campylopys pjttjerj commynjty Campylopus pittieri Williams I C. fragilis (Brid.) B.S.G. 13 <I <I <1 2 Peltigera pulverulenta (fayl.) Nyl. Leptodontium wallisii (C. Miill.) Kindb.

Leptodontium capituligerum C. Miill.

Nephromopsis ? sp. A {# 1172) Zygodon pichinchensis (fayl.) Mitt. . .

Accompanying taxa wjth a preference for the Upper Montane Rajn forest lor dwarf forest>.

Chorisodontiu m mittenii (C. Miill.) Broth.

4 <I <I <I 63 <I 4 30 2 Catagonium brevicaudatum C. Miill. ex Broth.

3 <I <I <I Hypotrachyna densirhizinata (Kurok.) Hale 4 <I <I <I 9 <I <I 1 2 Sphaerophorus formosanus (Zahlbr.) Asah 4 1 1 I 25 <I 3 10 Hypotrachyna andensis Hale 4 I I 1 2 Fernandezia sanguinea (Lind!.) Garay & Dunst.

4 <I <I <I 2 Riccardia capillacea 2 2 2 Stelis sp. (# 972) 5 5 Stelis sp. A{# 971) I Bromeliaceae sp. A {# 961) 5 5 Fernandezia sp. A {#941) 3 <I <I <I Leprocaulon albicans (fh. Fr.) Nyl. ex Hue 3 <I <I <I Lepanthus sp. A (#638) 2 Plagiochila sylvicultrix Spruce Plagiochila bogotensis Gott.

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3 Leptoscyphus physocalyx (Hampe & Gott.) Gott.

Cyrtolejeunea sp . A Metzgeria leptoneura Spruce 7 Cavendis hia cf. pubescens (HBK.) Hems!.

10 Disterigma acuminata (HBK.) Nied.

10 Adelanthus lindenbergianu s (Lehm.) Mitt.

Rhynchostegio psis cf. flexuosa (Sull.) C.Miill.

Scaphosepalum sp. A ( # 824) Elaphoglossum squamipes (Hook.) Moore vel valde aff.

Lophocolea quadridentata Spruce Elleanthus kermesinus (Lind!.) R.f.

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Epiphyte communities of tropical montane rain forests. II

].H. D. Wolf

64

65

Table 1. (cont.)

COMMUNITY 3

Hymenophyllum (Sphaerodonium) elegantulum v.d.Bosch # 725 Symphyogyna brasiliensis Nees Bartramia angustifolia Mitt.

Campylium ? sp. A Relbunium hypocarpium subsp. nitidum (HBK) Ehrendf.

Dicranodontium meridionale Bartr.

Plagiochila superba Lindenb.

Symphyogyna marginata Steph.

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8 <I <I I 9 <I <I 2 <I <I I Frullania caulisequa (Nees) Nees 3 <I <I <I 13 <1 2 IS Sticta fuliginosa (Dicks.) Ach. <I <I <I 10 <I <I <1 Hypotrachyna costaricensis (Nyl.) Hale 222711125 Aptychella proligera (Broth.) Herz.

16 <I <I 2 Frullania arecae (Spreng.) Colt.

15 <I I 9 Lejeunea flava (Sw.) Nees • 11 <I <1 I Sticta weigelii (Isert) Ach. • 10 I 3 5 Acanthocoleus aberrans (Lindenb. & Colt.) Kruijt 8 <I <1 d Meteoridium remotifolia (Hornsch.) Broth. • 8 <I I 2 Metzgeria spp. (sect. Metzgeria) <I <I d Marchesinia brachiata (Sw.) Schiffn. • <I I 2 Lejeunea sp. B {# 561) <I 2 5 Ceratolejeunea cf. maritima (Spruce) Steph. <1 <I I Heterodermia obscurata (Wulf.) Trevis.

5 <I <1 I Frullania standaertii Step h. • 5 <1 <I .::1 Squamidium nigricans (Hook.) Broth.

5 <1 <I 1 Lejeunea megalantha Spruce 3 <I 3 5 Lejeunea laetevirens Nees & Mont. <1 <I ..:1 lsopterygium tenerum (Sw.) Mitt. <I <I d Plagiochila subplana Lindenb.

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E piph yte co mmuni t ies of tropi cal montane rain fores ts. II

J.H . D. W olf

66 T able 1. (cont.)

COMMUNITY 3

Metgeria decipiens (Mass.) Schiffn.

Sticta cf. laciniata (Sw.) Ach. Leptogium burgessii (L.) Mont.

Porotrichum spp.

Plagiochila diversispina Step h.

Dimerella spp.

Lejeuneaceae Microlejeunea spp.

Orchidaceae Odontoglossum ramosissimum Lindl.

Leptoscyphus cuneifolius (Hook.) Mitt.

Normandina pulchella (Borr.) Nyl. Tillandsia sp. (sect. inflata??)

Megalospora tuberculosa (Fee) Sipm.

Drepanolejeunea inchoata (Meissner) Step h.

Taxilejeunea pallescens (Mitt.) Steph.

Leptogium spp.

Pannaria rubiginosa (Ach.) Bory Radula tenera Mitt. ex Step h.

Leptogium resupinans Nyl. Erioderma verruculosum Vain.

Radula sonsonensis Step h.

Lepidolejeunea spongia (Spruce) Thiers Epidendrum sp. A (# 645) Stelis spp.

Phyllogonium fulgens (Hedw.) Brid.

Zygodon cf. peruvianus Sull.

Lindigianthus cipaconeus (Gott.) Kruijt & Grads!.

Lobaria crenulata (Hook.) Trevis.

Plagiochila bidens Gott.

Bromeliaceae Plagiochila choachina Gott.

Polypodium angustifolium Sw. complex Plagiochila alpina Gott.

Prionodon densus (Hedw.) C. Miill. var.luteo-virens Heterodermia microphylla (Kurok) Swinsc. & Krog Radula voluta Tayl.

Leptolejeunea elliptica (Lehm. et Lindenberg) Schiffn.

Polypodium glaucophyllum Kunze Plagiochila adiantoides (Sw.) Dum. Prionolejeunea sp. A Ceratolejeunea patentissima (Hpe. et Gott.) Evans

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Epiphyte com munities of tropical montane rain forests. II

].H. D. Wolf

68

69

Table 1. (cont.)

COMMUNITY 3

Lejeunea aff. ambigua Lindenb. & Gott.

Grammitis semihirsuta (KI.) Morton Araceae Elaphoglossum aff. cuspidatum Moore Hymenophyllum nigrescens Liebm.

Plagiochila densispina Steph.

Themistoclesia recondita A. C. Smith Heterodermia comosa (Eschw.) Follm. & Redon Hypotrachyna cf. pulvinata (Fee) Hale Parmotrema subsumptum (Nyl.) Hale â&#x20AC;˘ Pseudocyphellaria aurata (Ach.) Vain.

Epidendrum xylostachyum Lindley Dictyonema sp. A (# 1008) Plutarchia monantha A. C. Smith Hydrocotyle Bonplandii A. Rich Bazzania sp. (aff. chilensis)

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In addition, the following taxa were only recorded once: In community 4.2.: Cladonia subsquamosa Kremp (1 %), Pannariaceae (< 1 %), Rhodobryum grandifolium (Tayl.) C. Mi.ill. (<1 %) . In community 3.1.1.1.: Bacidia sp. (<1 %), Buellia modesta (Krempelh.)

Mi.ill. Arg. (2 %), Catillaria sp. A #984 (10 %), Caloplaca brebissonii (Fee) Haffeln. & Poelt (< 1 %), H ypotrachyna rhabdiformis (Kurek.) Hale (2 %), Lecidea sp. B #805 (2 %), Pachyphyllum hispidulum (Reich b. f.) Garay & Dunst. (1 o/o ), Phyllopsora sp. A # 427 (2 o/o ), Teloschistes flavicans (Sw.) Norm. (<1 %). In community 3.1.1.2.: Coccocarpia erythroxyli (Spreng.) Swinsc. & Krog (< 1 %), Epidendrum cerneum group# 890 (5 %), Macromitrium cirrosum (Hedw.) Brid. (1 %), Myriotrema concretum (Fee) Hale (2 %), Trichosalpinx sp. A# 859 (30 %). In community 3.1.2.: Cheilolejeunea rigidula (Mont.) Schust. (< 1 %), Chiodecton sp. A #687 (1 %), Clusia sp. (3 %), Dioscorea sp. A# 1431 (2 %), Erioderma sorediatum Galloway & Jorgensen(< 1 %), Erioderma sp. A # 1281 (< 1 %), Frulliana cf. moritziana Linden b. & Gott. (5 o/o ), Grammitis jamesonii (Hook.) Morton(< 1 o/o ), Groutiella apiculata (Hook.) Crum & Steere(< 1 %), Harpalejeunea cf. oxyphylla (Nees & Mont.) Steph. (< 1 %), Heterodermia circinalis (Zahlbr.) W.A. Weber(< 1 %), H.lutescens (Kurek.) Follm. & Redon (< 1 %), H. speciosa (Wulf.) Trevis. (< 1 %), Hymenophyllum sp. C # 1432 (5 %), Lepanthes dasyphylla (2 o/o ), Lepidolejeunea involuta (Gott.) Grelle (1 %), Leucobryum polakowskyi (C. Mi.ill.) Card. (1 % ), M etzgeria fruticola Spruce (< 1 o/o ), Oreopanax cf. capita tum (Jacq.) Dene & Pl. (2 %), Pachyphyllum sp. (< 1 %), Peltigera dolichorhiza (Nyl.) Nyl. (5 %), Peperomia angularis C.DC. (2 %), Prionodon densus (Hedw.) C. Mi.ill. var. densus (< 1 %), Pseudocyphellaria intricata (Del.) Vain. (1 %), Pyrrhobryum spiniforme (Hedw.) Mitt. (< 1 o/o ), Syzygiella perfoliata (Swartz) Spruce (1 o/o ). In community 3.1.3.: Dicranoweisia sp.

A # 969 (1 %), Dicranum cf. paramicola C. Mi.ill. (5 %), Dictyonema sericeum (Sw.) Berk.

(30 %), Neckera chilensis Schimp. ex Mont.(< 1 %), Parmotremasancti-angeli (Lynge) Hale (1 %), Peperomia saligna HBK. (5 %), Pleurothallis antenniferae Lind!. (40 %), Pleurothallis sp. (1 %), Polypodium remotum Desv. (1 %), Scaphyglottis bicornis (Lind!.) Garay (5 %), Siphocampylus venosus (Gleason (5 %), Sphyrospermum buxifolium Poepp. & End!. 2 %). In community 3.1.4.: Cladonia squamosa (Scop.) Hoffm. var. squamosa(< 1 %), Fernandezia

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aff. ionanthera ~~chbf. f. & Warsc.) Schltr. (5 %), Grammitis aff.jamesonii (Hook.) Morton (1 %), Grammtlts he~eromorfa (Hook. & Grev.) Morton (1 %), Herbertus colombianus Van Reenen (< 1 %) , Lewmela ecuadorensis Robbins (2 %), Lycopodium sp. A # 1092 (5 %), Myrrhidendron glaucesce:zs (Benth.) Coult. & Rose (5 %), Pleurothallis sp. sect. Corocodilanthe (2% ). In commumty 3.2.1.1.: Cephalozia bischlerae Fulford(< 1 % ), Plagiochila divaricata Lindenb. (5 %), Polypodium aff. mindense Sodiro (1 %), Radula nudicaulis Steph.

(1 %). In community 3.2.1.3.: Kurziaflagellifera (Steph.) Grelle(< 1 %), Pileafallax Wedd.

(5 %). In co mmunity 3.2.2.1.: Plagiochila laetevirens Lindenb. (1 %), Thuidium minutulum (Hedw.) B.S.G. (< 1 %). In community 3.2.2.2.: Radula sinuata Gott. ex Steph. (< 1 %).

The most common taxa exclusive for the canopy communities are Anoplolejeunea conferta,]amesoniella rubricaulis, Leptoscyphus jackii, L. porphyrius, Plagiochila echinella sensu lata (incl. P. hansmeyeri and P. paludosa), Prionodon fusco -lutescens, Trachylejeunea dominicensis and Hypotrachyna laevigata (not separated from H. producta). ~ithin the ~anopy a ~wiglet. community and a? outer/middle branch commumty, both wtth a relatively high abundance of lichens, and three inner crown communities dominated by bryophytes, are distinguished. . . . .

Exclusive species for all five tree base commumues are, among others, Calypogeia peruv iana, Cephalozia crassifolia, Lophocolea aff. connata and Telaranea nematodes. The tree base communities may be divided in a Hymenophyllum lindenii-Bazzania breuteliana group with three communities and in a group of two communities with Symphyogyna brongniartii and Hypnella pilifera as most common exclusive taxa.

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