Tropical Forest Bryophytes

/1/1'1

iplivtc Ecology

f\1.1),.1,11:111,K. (1935), Zeitschr. Bot., 29,337-75. [\1.,,'.1,1':111,K. (1938), Ann. Bryol., 10,141-50. f\I.",,,,, 1:111 , K. (1969), Vegetatio, 16,285-97, 1\1.1)'.1,11:111, K. (1973), Herzogia, 3, 141-9. f\I.,!'.!, 11:111 ,K. (1975), Acta hist. Leopold., 9,95-111. 1'11.11,.1"11:111, K. and Wutz,A, (1951), Forstwiss. Cbl., 70, 103-17. f\1, 11",111. (19J5), Nova Acta Leopold. N.F., 3,123-277. , 111,11IIW;lIds, 1he sit nation iniproved somewhat. A series of l' '1'" ,1, ..1111" IIIIIJ t lu ,'piphvlic bryophytc communities in the sub""1'" " 1"1' '.1', ,,1 1;1[1:111 :11(' worih Illcntioning (e.g. Hattori and Kanna, 1'1 ,1. 11.11,," I " .rl , 1'1.(,. lw.uxuk i .uul 1la IIori , 1956, 1959, 1968, 1970 and II,. 1111111111'Ill,' 1',11"'1 III lw.uxuk i, 1'){jO). In tropical Asia the most ""1""'1111 \\,,,1. "II II,,· "I'II,I'vll,' C111111.111111' 1I1,)(I"I;II,'lv

,11,,111, I II ,,',

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J':li•• fun'st as a bryophyte

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FOREST

RAJN

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I'

ltryophyte Ecology The rainy period is not interrupted by a dry season or seasons longer than ~. II, months (seasonal rain forests). Exceptionally, the dry seasonmay be as 10llMilS5 months in some seasonal rain forests (Longman and Jenik, 1974) r 11 is compensated for by 5000 mm or more rainfall during other parts of I hl'

ycar.

Tropical ra in forests offer a large variety of habitats for bryophytes, IW\:HlIseof their diverse microclimate and the variety of substrata available 1'01'srnall plants (Richards, 1954). The microhabitats of bryophyte comrunnitics (Fig. 3.1) are determined by the different amounts of direct or rullrcct light and heat radiation and by the availability of water and Illtll'ients from direct rainfall, throughfalI, stemftow, mist and dew and in Ihl' form of ground water. Ali of these are inftuenced by the relative 1IIII1IIdity and by the physical and 'chemical character of substrata such as Illots, stems and leaves of living or dead plants, underlying rock and 11111\111I101' artificially disturbed soil surfaces. In general, the tropical rain IClIl'Nls urc characterized by high temperatures, always adequate for III yuphyíc growth, and by a water supply from a variety of sources. In the 111('IIt)1' of the rain forest water is continually present in adequate amounts l\'I'olllp"nied by high relative humidity but near the upper surface of the i'/llIopy water is of ten deficient. The distribution of light shows a very uuevcn pat tcrn in rain forests both verticaIly and horizontally and is of ten a IhlllllllM ractor of bryophyte végetation. \.1..l Mkrohnbitats (II)

1111,\'/',\' o/large

of bryophyte synusia in tropical rain forests

trees

1'1('(\ IIIISl:S ill (ropical rain forest represent the most shady habitat for 'plphyll1s, uxually combined with the highest degree of hurnidity, I~"pl'dlllly whcrc thcrc arc buttresses (see Fig. 3.2a, b) which sorrietimcs 11'/11'11 IlS Irigh ilS H-IO III up the trunk. The large surface provicles very III111hl!' IIk'!res for corticolous cpiphytic bryophytes (Fig. J.2h), otton II4NIIl'IIIIl\tlwith ülmy ícrns (Hyrncnophylluccnc). The samu ilpplii.:s to trce» wllh NIIII 1'0011'1 nml 10 the unastornosing ncriul root sysll'II1S or stl'lIlIglillg hl,lttll-plpltyl~S, which dcvckip inro indcpcndcnr trcc HI~'IIlN.HWII 011IIH' IIIIUIIIt hllNt'Nof l:rrge II'l'USlIud 011 l!ru lop of 11IOil11'Oots pl'Otl'IJdlllf!,tllloVl' fItl' Noll 1'111I I'un' 111(':1'(,' dl!vuloJls II xhnilnr HciophllollN hygrophlloll III YIII'ltylt' Nynwlllllll. flow fo,' thlN NyIllISilIlII I\I'OWNIIpWllldN 1111 1111' II illik uxuully clt'PI'IIIIN (111[uuuhllty 11I1dlill' plrYNkll1 l'OlllllljOIlN Ilf Iltl' luu k 1111111'1 1\~IIí\IIIí'ly Wt'l ('OlldlIIIIlIH II IltI~1111I-III'h III rlu- !tIlNIl III 11111111 11I1I11d1llN, It III IIIHllllly II IN "'Hldl'll·tI 1111hl'!tIWI'NI 1 °l lIliIlIlIIlIlly 1111"11111 "ltNt'III, III III IíII ly dl'Vlllllpl-" III luwhunl 11111\ llIlI'iIM. IIN 'ntlll'lIlirllty Hit 1IIIItl/ll( 111"11)IltiNHYIIWlltlll1i",i11111111/11 1t1\1I1111111~lvtll\t 1111" tllll\'lIpltVIt 1IlIlIlIlIlftll.Y, ','11111111'11111111111 11111'1,'''' 111111111111/,,111'1' 111111111111111111111111\' I

1,1

HI vophyte

Ecology

Tropical

, 11.11 .utcrized

by the different mat- and weft-forming Thuidium species glaucini associule of Iwatsuki (1960) in Japan; T. ,:/o//{, 'il/"idcs, T. bonianum in Vietnam; T. gratum, T. tamariscellum in c 111';1,T. involvens in East Africa) in which minuscule turfs or solitary ',1",IIIIl'IIS of Fissidens species are intermixed wi th appressed mats of 1 '1' unvaccae. On more porous bark with a decomposing surface many , ,'1/" ,1>/1'111/1 species, Leucophanes (Fig. 3,2b) and Calymperes species '" rur . ludicators of the very shady, wet habitats of tree bases are the ,1'lIdIUid or pendulous Neckeraceae and Pterobryaceae species (in Asia lILIIIV t lomuliodendron and Neckeropsis species; in Africa Porotrichum, ';milllldill. lIildebrandtiella; in America the ubiquitous Neckeropsis .//111, Ilii .uul N. undulata). Pinnatella and Lopidium species are present on ,II" 'II( 1IIl'II1s,Sematophyllum and Taxithelium species are also common in II" .vuuxium l observed about 100 species in Vietnam, 60 in East Africa 111"1>111 :11,';1)on the surface of their living leaves. These are the , 1"1,11\11,'II', 1••\', 'I'''VkS, Some lichcns also occur on the lower surfaces of 11\"'1' 1,,1\',", ;11111:111' s;lid 10 he hypophyllous, while the two groups 1,,,·, II" 11"111111..-Inlíi,'nlnllscpiphylCS,

65

flt,

IIr ophyte Ecology Th ' cpiphyllous life-form is most typical of the tropical rain forests. I \v -n so they are not found everywhere, usually being restricted to the 1 • IV 'S of young trees, shrubs and of long-lived herbs of the lowest layer of 1." ':>t, up to a height of 2-3 m and near streams. In the most humid types of , I II forest they also occur in the canopy. Aceording to Olarinmoye (1974) Ihl'i,' rrowth rate is closely correlated with atmospheric humidity. bl iphyllous Iiverworts occur in extratropical regions only amongst the IlIost occanic climatic conditions, e.g. in Japan (Karnimura, 1939), in 'hillu ( hen and Wu, 1964), in the southern Appalachians (Schuster, II) 9), in the Macronesian islands (Allorge et al., 1938; Sjörgren, 1975, II) IH), in the southern Caucasus Mountains. (Pócs, unpublished), and even I r Ir North as British Columbia (Vitt et al., 1973). ()I -qual importance with climatic factors is the nature ofthe leaves upon wh 'h '1 iphyllae occur. They must have a relative ly long lifespan and, 1" '1 -Iorc, .piphyllae practically never occur on the leaves of deciduous II (' or shrubs. They prefer evergreen, leathery or papyraceous (e.g. pnlm) 1 IIV s (Fig. 3.3a) but are of ten found on ferns, on leaves ofmonocot III'II!" ',', from the ginger family, They do not avoid hairy leaves, but h',I " with 'unwettable', waxy surfaces are unsuitable (Richards, 1932). 'Illlll' '1 iphyllous species show a preference for hard leaves of Podocarpus III ','//('('fJ"olflrfos, e.g. Diplasiolejeunea and Leucolejeunea species, whilst I II /)luII/o/ej unea, Cololejeunea and Hookerioid mosses usuaIly occur tili 111111f .rns (Pócs, 1978). IIhuu ih th high relative humidity that epiphyllous bryophytes require 11111'11:lSSO .iat d with shady localities this does not mean that they avoid 1 I hl. III '0111rast, th y n ver occur on the dark (and dry) lower leaf surface uul, II humkíit pcrmits, they occur in open places su ch as cocoa or Citrus pl "II IIIOIIS, al strearnsides or at the forest edge. Their water and nutrient IIppl i ti 'riv xl fr m precipitation. Rainfall in the tropics often contains 111111\'111111 i '1I1S than in tcmperate regions (Jones, 1960) and rainwater 1 til "l' Ihl Oll~ h Ih anopy is cnrichcd by xudate of leaves, excrement of 111111'111111'1, '1', ( zabó and 50rt05, 1975), Finally, th cpiphyllous III Ilpll ll', 111'1111111 live in ti community on th I af surface consisting of III Vllp" 1'" li .h 'liS, alua , [un i and smHII animals such as Tardi 1!'ada, I 11111111Iltnd A 'Mina, formiri Ih phyllosph r (Mussart , 1898; Ruin n, lilit 1), TIl(' ph 1I01\I1h'r' is tibi' 10 fix fr" nitroa n probably lu' to th' Iwl Iyol' L hlu 'of 1" '11111iul '01111"011'nt (ef'. ulso larinmoyc, 1(74), II1 yophyl '. I'SP 'cillll lív .rwort», IlII Ih, 1l1Os1 importan! rolc in Ih' I plphVlIll1I 'Ollllllllllili 'S ' "pl in dd '1' hnbilalN wh '1" lich 'ilS 'Hn sí ill 111'1111TIH'liVt'lwllfls,livill1' illihis hohitnl, nrc usunll V'I' spcciulizccl il )llIyl!' , "lIrI IId Ipl 'd 10 th, pi ph 1111lll' 'ollditiollS Illd 1110 t '" IIIt I II II' I'Plpllyllll ,For' IIl1pll' , IitI' II l' III ('piph 110\1 Iltll 111111 1 III tlulllt I/()I ,'1 IIti 111'('I IIld 10 11111 111'1'11' 1111111111111 11111,,1 II"

11'11(, J,J (ll) 1~piplryllous community of Radula acuminata on the \eaflets of Caryota (1'111111" ,) in Vietnam, Cuc Phüöng National Park, (b) Dr E.W. Jones, 1'1111 '1'1"l hl oph 1 'S on a rain forest roadcut surface in the Uluguru Mountains, '11111/11111, HIINt1\ fricn, ( ,) Bryophytc synusia of Calymperes richardii and ( J, f/l/llI"I//lII//1/I (I/h/tI//1/I on th' trunk bas of a planted coconut palm in the rain 'tilI I lilit 111'\11\1111('011,'ubu. (d) l,ejelll/eolo,tev;renscoml11unityonthe 111111'111111 1I11111YIIIPItlIII (U(/II,I'IrIIII'fI f(';ii,,) in d 'grad d rain forest near Santiago rit I 111111 1I1I1//(I,\'f(/('!lYfI

/1 rvopltyte

Ecology

Tropical Forest Bryophytes

III '1 \\,OI(S are obligate epiphyllae, the rest facultative (Pócs, 1979). Thc "".\', The hamboos have their centre of evolution in tropical Asia 1\IIt'Il' Ldl woody rcprcscntatives are common in alI vegetation beIts as I 'LI II1 1"'Il'lIl1i:iI I re: c-likc plants or lianas. In tropical America the Bam1.11",· .u ,~,Iq'll'Sl'IIiL"d only hy smalI climbers but in Africa species are '"'1'0/11.1111v"I',"l;lliI'S ill South Vietnam (Tixier, 1966a). At a more developed stage 11"'.1 ·."•.•·I('S !,.ive way to more specialized lignicolous assemblages from 1 , 1,,01,,11;11'( ';Il', Odontoschisma, Lophocoleaceae, Lejeuneaceae and 1;/" ,I/,Ii,1 1(1 Authoccrotalcs, Leucobryaceae, Calymperaceae, Hypna, , .H III'/'"III"/I'gil/III, Hookenaceae and SematophylIaceae. These )" \' '1,I!\,1,.,; ;1II' al It':Jst p:lrt ly mycotrophic or arc able in some other way to IIld •.• IIu : 1IIJI,i('1I1s ol the elecomposed wood. Other species probably 1",1, I II,,· !'.lIlId w:lll'rahsorhing and -retaining properties or the acidic 1" "1"

1,,··.," d,'; 1\ I w' H )d. III ( " d '.1. 1" v' ,v,·!'.'·I:J1i()1I')11dl'l'Olllposillg wood (inclllding dccaying 1

tree 1. III '.1.11)'.) ''1'1'";11(''' III ll(' lill' ,il'heSI ill spcl'ializL'(llignicololls species. \. ·",I//d "'1/,:1/111. /l11''f'III'fI'/'I',t;illlll IIi 11'/11 'l,ftl'/l1 1111 , /11. carinatum, and 1',/. '11/, '\' II/Ill/ti .I cn u d aturn .u « a lwa vx 1)"'~;l'1I1()II rott cu wood and also , II, '"11 lill' 1" IlII... 111/1-//11101'.1/1' "1'1'·'I/III'tI. /ool'.li.l IIlIlill'I/I'I. ("'Idllllo;:i" ,./1 d,I,,'dll!'

II,

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1Ill, '1'11,hVIil' plants and the distribution of an nu al rainfall. The epiphyte 1'1''111.1·.·. 1" Ilii '1l(lll ional to annual rainfall amounts that exceed a monthly i\' 1.11" ,11 IO()111111. Tropical macrovcgetation flourishes increasingly with Ill' l' .1'.1111'nu int hlv rainlall up to ](JO-200 mm per month. Above about 'filill ",11(1111111 r.unl'all ;1 ycar thcrc is no Iurthcr 'improvernent' and . Ill" 111111' ,,1 \'l')'.l'1:lIioll. On the contrnry (ef. Longrnan and Jcnik , 1974), III 1" r l u n un l 1'·)'.111I1S,lu: prOdlll'livily ol lorl'sls is wc ll hclow th.u ol .m 1\' f .11" 1,1111 1",,'c;1 or ('V('1l01;1 s('asollal r.un loresI. 11,,\\ .1"". 111('',IIIpilIS r:lillLtIl othc-rwi:«: alkl'1 Ille IroJlic:d 1'1)\('sl'l \ IIll, ,111,11 Ill,' 1"'I\('I:d I'lIv:;if'!'.IHIIIlVIll'tlll' IIllSI.ou uruu nt v 1!t,,·SIli,l I'h:III)',(' 11111'" \\ ilii 1,11t l ur Illi'l,':IS" ol' r:lilll:dl, ih "I'il'hvli(' hiolll:lc;S .uu l lill' ilii, 1" 1>11\' "'1',IlIIV (ll ('I''1,hvll'S Illi'l,';I';":·; III 11IOI"1l11I1l1 1(1lill' ";1111'111:;' 111111111.1111'>11111 l u I!u- 1\\,(1('.\:11111'1":';1'.1\'('111\('1(' l«: :.1111'111:; 1:11111:1111'.111111 ",.1 1'1111111111 1""I"',II\"'lv III .. 11In 1,1:1''('';I1IIlli' ',;11111' 1",,""1:1111',1111'1,'.u . "1"",,"1.,,11' ('\('11'1''1'11 1,",",1'; .nu l ..tlill 1'"(".1', wIIII 11111'"l.:.... 1'1"'111 I iI""1 .111/'1'111/1

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I'II/'sii

Forest

Bryophytes

In tropical Asia true páramo is not known. In the South American Cordilleras many Espeletia species (Compositae) comprise this vegetation type. Their bryovegetation is briefty summarized by Griffin (1979). In tropical East Africa the giant groundsels (Senecio subgenus Dendrosenecio, Compositae) are the best known and most typical representatives of páramo trees. Although giant Lobelia species are also numerous they never form páramo woodland alone and epiphytes are not known on them. The Senecio stands are of different structure in the different African high mountains aceording to the climatic conditions. In the most humid, the Virunga volcanoes and Ruwenzori mountains they form a continuous belt of closed, 6-8 m high forests between 3800 and 4300 m. On Mt. EIgon and Mt. Kenya they form more open woodlands on the slopes, while on Mt. Kilimanjaro Senecio páramos are restricted to the valley bottoms except for the most humid south-south-west slope along Umbwe Route where its stands occupy large areas, intermixed with Erica and Philippia. In the Senecio stands, most epiphytes live on those parts of the trunks where the dead, shrivelled leaves, which usually form a protecting cover on the stem, are shed and the bark is visible. Bryophytes usually form large cushions on Senecio trunk and branches and are most commonly present at the branching points (Fig. 3,8c). Almost alI are xerophytes, resistant to severe insolation and frost exposure. Some are endemic afro-alpine species, others are disjunct temperate elements (see Table 3.8). Hedberg (1964), studying the afro-alpine plant communities, dealt in dctail with the composition of Senecio páramo woodlands and described their composition, including the bryophytes, It is clear from the literature that there are marked differences between the epiphytes of the humid Scnccio stands of the Ruwenzori Mountains and those of the mu ch drier Mt. Kilirnanjaro stands. The ground layer of Senecio páramos is very rich in bryophytes. Usually ;1 IlIosaic or habitats is observable below the trees composed of spring-bog lIichc-s ;dong streamlets and drier soil or rocky niches. On Mt, Kilimanjaro Illi' ,II lill ill;1I11species of the spring-bog synusia areAntitrichia curtipendula, ( '111/1/'1'/01'1/,\' stramincus and Hygrohypnum hadbergii. On dry soil and Illik:, 1111" .Iomin.mts arc Anastrophyllum auritum, Gynomitrion laceratum ,llltl/,,/,I,,,{,,I/III/I/I/I'I/('rasccns.

1'11 ,"(1'"

,'. 1.111'111,1",1"1,',1111111111111"/111;111111..11''1'11111'''''''''''1111'1'''''1." IIII 1·11 \\·ill, 1IIIt-1ill,II\-/I()II'; \'í",j',í"I:1111111 (II (",It .Ii 1111,,1, \\,1'1 II II. ,","1.,.1 .1111111..1 \ ,111,11'/01", III ,11111.11,(,.1.111\, il 1111,1, .1,,\ 1, III

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III 11)('I 'II/It/ ,/,ill and ill p.irarno belts two further interesting communities ,II. ", IIIil 1I11"1I11l1l1i1lg, Richards and Argent (1968) descrihed the interest"'1' 111\11"'1'.,'1;1111111 ol lava rock caves from Cameroon Mountain. II" ,. ,II. \. 1\ ',lIlId:lII;IV;1 c.ivcs on Mt. Kilimanjaro, between3300-3900 "' \\ 1"" lill 1111'IIIIWI w.ill-, ill dricr places Rhirofabronia perpilosa, ( ",f,d(,' 1, fid, 1 11111III 11I'1I.\'ts , !>i/,IOI,hl'l/I/1II ajricanum and Gymnomitrion ld" I .t t t nt t " •• III lill \\,,'11(111,.1\'II'\V, ;11" lill' l'IÚ'Cls of the dcsert ecosystem as a whole (Ross, I'J(,'II II", ;111':1';"VCI which thcsc conditions obtain inc\ude southern 1.,

-t,

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11" '" , ., l/fff

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IIIV' '1,IIv10'":!IIII'S:!Il' 1101citcd whcrc they follow Index ,/1/1111,