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EDIDI:r'
SVENSKA V AXTGEOGRAFISKA SALLSKAPET
Dick J ohansson
Resume:
Ecologie des epiphytes vasculaires dans la foret
dense humide d' Afrique occidentale
UPPSALA 1974
ACTA PHYTOGEOGRAPHICA SUECICA 59
Distributor: Svenska vaxtgeografiska sallskapet, Box 559, S-751 22 Uppsala, Sweden
Dick Johansson
Dick J ohansson
Resume:
Ecologie des epiphytes vasculaires dans la foret
dense humide d' Afrique occidentale
UPPSALA 1 9 7 4
Suggested citation : Johansson, D., 1 9 74, Ecology of vascular eopiphytes in West African rain forest.
Acta Phytogeogr. Suec. 5 9 . Uppsala.
© Dick Johansson 1 9 74
Printed in Sweden 1 9 74
Textgruppen i Uppsala AB
Abstract Johansson, D. 1974, Ecology of vascular epiphytes in West African rain
forest. (Resume: Ecologie des epiphytes vasculaires dans la foret dense
humide d'Afrique occidentale.) Acta Phytogeogr. Suec. 59. Uppsala. 1 3 6
pp.
in the Nimba area 1 5, The phorophytes (the tion 87, Succession traced by analysis of humus
Preface
My first contact with a tropical rain forest dates M any other Lamco employees have in one way or
back to August 1 9 64 when I com m enced employ another helped or simplified m y field work.
ment for Lamco J.V . Op. Co. in Liberia. As a high Help and guidance in the determination o f critical
school teacher at the I nternational school in Yekepa specimens have been received from Jean Bowden,
I used my spare time during the following six years F.N . Hepper, Prances Jarrett, M. Sands and P.
to explore the rain forest. In the vicinity of the newly Taylor of the Kew Herbarium , and from L. Garay of
constructed Yekepa community , at 5 00 m altitude , the Oakes Ames Herbariu m , H arvard Univ. , C am
virgin rain forest covered the slopes of the Nimba bridge, M ass. Prof. Sanford, Univ. of Ile Ife, Nigeria,
range up to the peaks at 1 300 m altitude. The rain provided an unpublished m anuscript on the use of
forest was very poor in species in the field layer . On epiphytic orchids in general ecology work .
the trees , however, vascular epiphytes were abun The determinations of the insects were performed
dant and soon caught m y particular interest. While by M . A . Grog an, S . L . Shute and I . H . H . Yarrow of
trying to learn about the ecology of the epiphytes it the Entomological Dept. of the British Museum
soon became evident that the literature on this sub (Natural History).
ject was very scanty. Although the vascular epi Aina A im , Gertie Backlund , Margareta Hellman
phytes are a very characteris tic floristic element in and Barbro Enander h ave typed the m anuscript.
the rain forest inaccessible growing sites have M iirta Ekdahl has prepared the large tables and
resulted in few and o ften superficial reports on their Ulla-Britt Sahlstrom h as copied the photographs.
ecology. The map over the investigated area was drawn by
A fter an i ntroductory period in which to become Per Ake S oderman .
acquainted with the tropical flora a prelim inary plan In the preparation of the m anuscript valuable
for the future investigation was outlined in coopera criticism has been provided by Doe. Orj an Nilsson.
tion with Prof. H ugo S jors at the Institute of Dr Jerk Hellkvist at the Institute of Physiological
Ecological Botany in Uppsala. Botany in Uppsala has commented on the section
A fter returning to Sweden in 1 9 70 m y work was about the water economy of the epiphytes. The
continued at the newly started Afro-Botanical Dept. linguistic revision has been carried out by Andre
of the Institute of Systematic Botany in Uppsala un Brochu and the French resum e was translated by
der the guidance of Prof. Olov H edberg. Renee Perrette.
During an excursion to Kenya and Tanzania in Granges International Mining Division,
January 1 9 7 1 , under the leadersh ip of Prof. Stockholm , has supplied a grant for the photo
Hedberg, I was able to study epiphytes in several en graphic illustrations.
vironments different from those of the lowland Prof. Hedberg, my teacher and tutor, has through
tropical rain forest. As an indirect result of a three his never failing enthusiasm and encouragement ac
months ecological study the same year for Sweco in tively contributed to the completion of this work.
the Kidatu area in Tanzania, a knowledge of Finally , Astrid did a considerable amount of work
epiphytes in dry environments was gained . behind the scenes.
Several persons have contributed to the comple A ll person s and institutions mentioned above I
tion of this study. H arvey Boettcher, M anuel Diaz want to sincerely thank for valuable cooperation.
and Franz Zimsek o ften accompanied my tours of Uppsala, January 1 9 74
the forest. K urt Eriksson helped in the construction Dick Johansson
of the light-measuring equipment and Miss Anne Institute of Systematic Botany
C ron e carried out some laboratory work in Liberia. Box 5 4 1 , S-75 1 2 1 Uppsala, Sweden
tioned below, in parentheses, is take n from Willis What are the p roperties that make a plant successful
(1 966). The majority of the orchids (20 000) are as an epiph yte? Two condi tio ns must be ful filled. (1)
epiphy tes (cf. H u nt 1 9 6 7, S ummerhayes 1 96 8). The The plant or i ts di aspores m us t i n one way or
b romeliads ( 1 400) play an important part in the another be able to reach a posi tion on the phoro
epiph yte flo ra of the New World and to a lesser ex ph yte. (2) It m ust possess the ability to s urvive
tent this also applies to the C ac taceae (2000). The drough t after its establish ment.
latter family also i ncludes a pantropical genus, Rhip- ( 1 ) The weigh t of the diaspores h as been emphasized
Barkman ( 1 9 5 8 :9- 1 5) has given a v al uabl e s um m ary Th e typical epiphytes, h av e also been divided into
of the terminology con cerning non-vascular epi a large num ber of units bas ed on the hab it or
phy ti c cryptogams . d ep enden ce of water, light, a nd s ubstrate.
R egarding non-vascular cryp togams th e term Growth habit
obligate is o ften used for typical epiphytes s en s u S chimper ( 1 888) r ecognized three types : pro to, nest
Oliv er (Table 1 ) . Th e r est of the epiphytes are called or bracket, and tank epiphytes , bas ed on the habit of
facultativ e (Barkman 1 9 5 8 : 1 1) . A s ubdivis ion of the the plant.
facultativ e epiphy tes into three categories, preferen
tial, indifferent, a nd o ccasional h as been used by Water
Hilitz er ( 1 9 2 5 ) . Both v as cular and non-vascular Th e d ependen ce on water is a com mon cri terion for
plants are incl uded in the m i cro and macro-epiphytes a s ubdivision. Oliver (I 930) distinguishes between
of Tixier ( 1 96 6 :45) and in the Sonn en(lich t), hydrophytes, m esophytes and x erophytes. S chnell
S chatten, and Epiphyllen v eg etation of K napp ( 1 9 5 2 :2 8 2) recogni zes the sam e units as L ebrun
( 1 973). ( 193 7) with one addition, 'les epiphytes hygro -m eso
Oliv er ( 1 9 3 0) pres ented a s ch em e that will b e used phi l es'. Th e units used by S ch n el l are th us :
h ere as a base for comparing th e terminology of I. Epiphytes hygrophiles d e la base de troncs
different authors (Table 1). I I . Epiphy tes hygrom esophiles d e la region moyenne
I. Typi cal epiphytes. Species which are h abitually d es tron cs
epiphyti c. Ill. Epiphytes m esophiles des grosses bran ch es et
du sommet d es tron cs d es grand arbres
II. O ccasional epiphytes. H er e are i ncluded those
I V. Epiphytes x erophiles d es bran ch es superieures
plants wh ich are under ordinary circumstan ces
terres trial . O ccasionally they are found on trees
Light
wher e they appear to thrive until maturity.
A division into shade-, s un -, and extreme x ero
I ll. H emi -ep iphytes . These are trees which begin ph ilous epiphytes , has been used by Davis &
th ei r existence on other trees or tree ferns, send down Ri chards ( 1 9 3 3 :3 80-3 8 1 ), and R i ch ards ( 1 9 3 9 :3 1 -
one or more taproots to th e ground and eventually 3 2). R i chards admi ts the diffi culties in disting uishing
es tablish th em selves as indep endent plants. th e latter two gro ups.
their water econo my . The use of epiphy tes as i n vations from roughly 1 500 trees, pal ms, fer ns and
dicators of microclimate and forest destruc tion was lianas.
also i nves tigated (cf.. Sanford 1 9 7 2). O u tside the specially i nvestigated area I have
A more specialized s tudy regarding the phen carried out surveys of the epiphy te flora in every
ology, polli nators a nd special field charac teris tics of cou nty of Liberia. The neighbouring countries of
the orchids was also i ncluded. Gui nea and Ivory Coast have also been visited on
My s tudy i s res tricted to the vascular epiphy tes. several occasio ns e nabl i ng a comparison of the flora.
However, the so-called filmy fer ns, e.g. Trichomanes Epiphytes in e nvironments, different from the rain
and Hymenophyllum, are i n the mai n excluded, but forest types, have been s tudied i n K e nya and
the species observed are listed i n chapter I I (Table 7). Nor thern Tanzania (Jan. 1 9 7 1 ), a nd in Sou thern
In several way s these fer ns resemble the non Tanzania (April -June 1 9 7 1 ). Brief v isits i n Zambia
vascular epiphy tes. Their leaves are usually o ne cell (July 1 9 7 1 ) a nd i n Cameroon (M t Cameroo n, July
l ayer thick and without sto mata and can take up 1 9 7 1 ) have supplied addi tional i nfor mation.
water i n the same manner as mosses. The facultative E fforts have been made to get photographs from
(occasional, hemi- and ephemeral) epiphy tes are also specime ns growi ng i n situ, bu t difficul ties in ob
excl uded but listed (Table 7) and referred to when of tai ni ng good pictures of some of the phorophytes i n
i mportance for the epiphy te flora. the dense forest and o f the epiph y tes i n the crowns,
explai ns why some of them had to be photographed
FIELD W OR K in rather unnatural e nvironments . All pictures, ex
My field work w a s carried out during 1 9 64- 1 9 7 0, cep t Fig. 3 1 , are taken by the author.
wi th the major part of the distribution recordings
fro m the las t two years of thi s period. TER M INOLOGY
The scanty k nowledge of the presence and the dis Epiphyte is here used as defined by Barkman
tribution on the trees of epiph y tes i n the tropics can (1 9 58 : 9). When no thi ng else is stated the term
be traced back to two different causes. The first is epiphyte refers to vascular epiphy tes. The 'host tree'
the very i naccessible growing sites of these plants. (shrub, pal m etc.) of the epiphyte is referred to as the
The second is the difficul ties connected with the phorophyte followi ng Ochsner 1 9 28 ; (cf. Barkman
determi nation of the epiphy tes. Several species are 1 9 58, S jogren 1 9 6 1 , Tixier 1 9 66, Skye 1 9 6 8). This
extremely si milar i n size and shape i n their flowerless term is accordi ng to the defini tio n by Ochsner
s tates (Fig. 3 3). ( 1 9 28). The no me nclature of the phorophytes agrees
To overcome the first obs tacle my research has wi th Voorhoeve (I 96 5) and for species no t treated
been directed i n three different ways: record ing of by him the nome nclature in Hutchinson & Dalziel :
the epiphyte flora, ( 1 ) on felled trees (close obser Flora of West Tropical Africa (F .W.T.A .) 2 nd ed., is
vation), (2) of s tandi ng trees (dis tance observation), followed .
(3) on a mi nor part of a tree by close or distance The term high forest is applied to fores ts with a
methods (occasional observati o n). The two first canopy at a height of 3 0 m or more regardless of the
methods give a rather accurate picture of the dis origi n. Secondary or rege nerati ng fores t refers to
tribution of epiphy tes on the trees while the third is forests in various s tages of regrowth wi th the canopy
utilized for addi tional i nfor mati o n. below 30 m height.
To get acquai nted with the epiphy tes i n various
stages of their developme nt I s tarted fro m the very
beg inning to collect spec i me ns and grow them in
The area of investigation
s mall wooden boxes in a specially cons tructed 'frame
house'. The phenology of the various species could Th e area i nvestigated, called the N i mba area, is
then easily be studied and minor ch arac teristics si tuated in northern Liberia, near the border be
observed. The i nves tigation i ncludes the exami nation tween Gui nea and the Ivory Coas t (Fig. 2) a nd i n
of 22 0 felled trees and 236 trees exami ned accordi ng cludes parts of the Nimba M ts . The cou ntries rele
to the distance method on 4 7 species of phorophy tes vant to this i nvestigation are shown in Fig. 3 .
i n the high forest. The so-called occasional recor Large mountai ns are poorly represe nted i n Wes t
d ings number more than three thousand obser- Africa bu t within the so-called Guinea Highland,
GUINEA
BONA •
lJMT YULLITON
(/.EETON N
I
I
I
�MT TOKAOEH
I
I
I Mf SOUTH NIMBA
I
I
I
I
I
I 'ZlJLO'Wit
I
I
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/
3km
.______.___....___�
�
L.::..:.J 800- 1000m j:::::::::::]1000 -1300 m H��h��H 1300 m
>
stretching from Fou ta-Dj allon in Gu inea to the Dans the Ivory Coast the Dans mass if contains the two
massif in the Ivory Coast, a series of r idges and high peaks of Mount Tonko u i, 1 240 m, a nd M t Dou
peaks r ise to altitudes of more tha n 1000 m. I n 1340 m . The N imba Mountains s tretch fro m Guinea
northern S ierra Leone, the Loma Mountains reach into L iberia, where the r idge has a N E-SW d irection
1 94 7 m at the Bintu mane peak . I n Guinea, Mount w ith the h ighest part s l ightl y above 13 00 m (Fig. 4).
Bal a in the Ziama massif is 13 8 7 m h igh, but eve n Here the range is ris ing abruptl y fro m a tableland at
more imposing are the N imba Mountains w ith a around 5 00 m altitude. I ts h ighest part is b u il t up by
number of h igh peaks (e.g. M t M olard 17 5 2 m). In a narrow ridge or crest w ith s teep slopes on each s ide
'
2
/Po r �'7=""�":---+--'-�=--.,.,..-
Guin
Nigeria
(Fig. 1 1 ). The southernmost part ends in a steep fro m a h igh-p ressure belt above the subcontinent of
mountain, Bele (Bilimu). East of the ridge the two South Africa towards a low-pressure b elt formed
p eaks Mt Gang ra and Mt Yulliton of more than above North Africa. Th is w ind is in W est Africa
1 000 m dominate the landscap e. South of th ese known as the southwest monsoon.
p eaks rises another ridge with th e h igh est part at Mt When th e moist SW -monsoon reach es the
Tokadeh at around 1 000 m altitude. After the dis Lib erian coast h eavy rains start to fall. Th e rainfall
cov ery of iron o re in the Liberian N imba in 1 9 5 5 and in th e coastal strip is v ery h igh, generally mo re than
th e start of mining operations in 1 96 3 , a large influx 4000 mm a y ear. Towards the interior p recipitation
of p eople has tak en place into this p reviously sparse dec reases, and in the northern part the p recipitation
ly populated area. The early stages in the exploration is l ess than half of the amount of the coast.
and th e new dev elopments in this area are d escribed In th e southeast of th e country, a short dry p eriod,
in Gardlund (1 9 6 7). 'th e mi ddl e d ry ', often app ears in Jul y -A ugust, which
in c ertain y ears is even felt in N imba (Fig. 5).
I n periods of v arying length the macroclimate at
CLI M A TE
Y ek epa (5 1 0 m) has been measured, i n a traditional
For L ib eria as a whole the y ear can roughly b e way, w ith th e instruments in a meteorological box,
d ivided into a rainy s eason from March/April t o Oc 1 .5 m abov e the ground. The rainfall has b een
tober/November, a nd a dry s eason during the re measured w ith a standard rain gauge also at 1 .5 m
maining months. These extreme w eather types are above th e ground. The maximum and minimum
c reated by the y early movements of the I nter temperatu res were tak en at s ev en o 'clock for each 2 4
Tropical Front (I .T.F.). This front is formed b e hours p eriod. Th e r elativ e h umidity w a s recorded
tween the relativ ely cool, moist air of the equatorial with a hygrograph, wh ich was calibrated to a psy
zone over the Atlantic Ocea n, the Equatorial chro meter (dry and w et bulb th ermometer) . At the
Maritime A ir M ass, a nd the hot d ry mass of air over p eak of th e ridge, th e mining company , Lamco
th e S ahara, th e Tropical Continental Air Mass. Dur J.V . Op . Co., has s ince 1 95 7 p erformed regular
ing th e d ry s eason the movements of thes e air masses meteorological observations. Unfortunately the
are, s imply speaking, regulated by the h igh-p ressure observ ation site has b een mov ed twice, and the data
b elt formed over the Sah ara and th e low-pressure after January 19 6 7 only includ e rainfall and
b elt ov er the area of South Africa. The w ind blow cloudiness.
ing fro m th e h igh-pressure b elt to th e low is d eflected Th e mic roclimatical effects o n the d istributio n of
to the west owing to the earth's rotatio n. This wind th e ep iphytic plants w ithin the trees are more con
wil l th us appear as a north eastern d ry often hot and spicuous than macrocl imatic effects . (Howev er the
dust-laden w ind, co mmonly k nown as the H armat d ifference between th e epiphytic flora at the c rest of
tan. During the rainy s eason the wind is blowing the ridge and that in the surrounding lowland is no
Yekepa 510 m
500 ..., ... "',
,--,... \
/ \
30 \
21-22.6 1969
,------
25-26.1 1970
r;' \,
-
25
-
-
250
.-- 20
r--
r-- 15
,----
06 12 18 24 06
n 10 12 14 13 18 20 21 15
I Fig. 6. Temperature variations during a day in the wet and
a day in the dry season. Yekepa 5 1 0 m.
� 10 11 15 17 21 20 23 10
temperatures. The lowest temperature recorded is
from February 1, 1 9 7 0 when a readi ng of 12. 5°C
was regis tered during the night. The following day
J A N. DEC.
the m aximum temperature reached 3 2 . 5°C which
Fig. 5. Monthly rainfall at Yekepa 5 1 0 m and at Nimba gives a d ifference of 20. 0°C (Fig . 8). The daily v aria
Range 1 248 m, 1 969. The number of days with rainfall is tion of the te mperature at the higher parts of the
given below each month. ridge is u nk nown but there is a five year (1963-
1967) series of maxim u m a nd minimu m temperature
doubt regulated by the macroclimate.) Unfortu nate recordings from the Geologis t's Camp Nimba. The
ly, m ic roclimatical investigations in the treetops and figures from 1966 are given in Table 2. The
along branches, 30-40 meters above the ground, differences between the mea n maxi mu m and
have been impossible to perform. The microcl imate m inim u m temperatures are s mall. The largest
in a rain forest, discussed later in chapter V, has been differences occur i n the dry season.
investigated in Nigeria by Evans ( 193 9) and in the In a h igh forest, at gro u nd level , the daily variation
Ivory Coast by Cachan (I 963), and Cachan & in temperature, as one would expec t, is s maller than
Duval ( 1963). outside the fores t (Table 3). K u nkel (1966), who h as
carried out occasional temperature recordings fro m
Temperature various pl aces in Liberia, esti mates the temperature
In Yekepa the daily maximum temperature during in an open area to be around 5°C higher tha n inside
the rainy seaso n is close to 3 0°C and the minimum the forest.
Temperature oc
Period Max. Min. Mean
January 22.8 19.2 2 1.0
February 22.4 19.0 20.7
March 21.7 18.7 20.2 30
I I
September 19. 1 17.4 18.2
October 20.6 18.4 19. 5 . I I I
15.6 30. 6
November 22.7 20.3 2 1. 5 1.6
i 6 8 10 1214 16 18Z0 22t� · 6 8 '" '"/ t¥ ' 6 8l12 " 16 18 Z022?Y.· 6 8 10 "" 16 18 Z0 2�· 6 8 10 1214 16 18 Z0 2!� · i 8 10/l 16 1820 22?� ·; 8l ""l 1820 22?� .
mi.J-J-1.1- LU�IIJ_H.)
Hontag Monday Lundi Dit:nsfoq rue:sdoy Mordi Hillwoch Wednesday Hercr�i Donner&tag Thunday Jeudi I Freitag Friday Vendredi I Sonnobend Saturday Som�i I Sonnlag Suttd_
oy Dimonche IMonfaa
. UII 1 I I U_/
1,[ I I
,
u
.
- r e ke p a 5 1 0 m
7 0 I I 'Jj I I
L// /// ril l 1/1/ 1/ l rlllll rt /Tlr/1 ! rUt/ lrt l I TT /ll rt / 11
J an . 2 6- Feb. 1 1 9 7 0: I V" I I •� .,.. I J" A-T 1" I
. �H
1 -H�-1+��
r -+ r V V
/ I
I 1 7
��-��.++ � -l-
t--� ·
�
.· Honlog HOliday Lundi Dit:rutaq lut:sdo Hard(
1 14 16 1820 21�,· 6 8 10l14 16 18Z022f'�i· 6 8 1 /" 16 181 ?�.· n61 10 12 "l "t22?�,·;
6i7""/6 "2flt?� · 6 810/"; ; �¥,· 6 8 10 12 14 16 1820 22?;� ' 6 8 ,0/ / Hillwoch Wednesday Hercredi / Donnerdog Thursday Jeudi / Freilag Friday Vendredi / Sonn !Hnd
a Solurdoy Sam�i f So nlog Sunday Dimonche fHonloo
Fig. 8. Relative humidity during a week in the wet and a imum and minimum temperature (°C) are given for each
week in the dry season. Yekepa 5 1 0 m. Figures for max� 24 hr period.
lowered during the day as the temperature rises. The small (F ig. 9). The weekly average of the absolute
absol ute minim um rel ative h um idity (Table 4), al m in im um relative h um id ity is as high as 84 %.
though only lasting for a short per iod in the after
noon, can be used to ill ustrate the yearly variation in
Table 4. Absolute minimum relative h u m idity. Mean for the
the desiccation effect on the plants.
periods. Yekepa 5 1 0 m.
Naturally large variations are found in the daily
changes in the relative h um idity. This applies to Absolute minimum
variat ions between individ ual days of the same Period relative h u m idity (o/o)
and the dry season (February 2-8, I 9 7 0) illustrates 1 1 .8- 1 7.8 1 969 73. 1
2 2 . 9- 2 8 . 9 1 969 55.7
thi s (Fig. 8). The impact of the dry H armattan winds
4 . 1 1 -9. i I 1 96 9 48.0
is clearly visible in the graph fro m the dry season.
8 . 1 2- 1 4 . 1 2 1 9 69 46.4
During the period 3 1 . I -5.2 I 970 the mean absolute 1 2. 1 - 1 8. 1 1 9 70 4 7.4
minimum relative h um idity was only 2 3 . 2 %. 2.2-8.2 1 9 70 35.7
A c t a phytogeogr. suec. 59
12 Dick Johansson
Monlog Hondoy l.undl Oi�nslaq Tuesday Mardi Hilfwoch Wedn�sday M�rm:di Donn�rslog Thrmdoy J�udl FreUog FrldtJy Vendr�l SonnabMd Saturday Sam�i Sonnlag Sunday Oimanthe Honlag
Fig. 9. Relative h u midity during a week in the wet season. Mt Gbahm 1 300 m.
Cloudiness Wind
The cloudiness has been recorded at the Geologist's In general wind velocities are low. Thunderstorm s
Camp Nimba and Mine Office at 1 248 m . The day are accompanied or rather p receeded by strong
was div ided into two recording periods, 06°0- 1 2°0 winds that usual ly last for only a few min utes.
and 1 2°0- 1 8°0 (Fig. 1 0). The ratio between cloudy Kunkel (1 9 6 6 : 5 9) reports that, at the end of
and sunny days in 1 9 69 reveals that June, July and February 1 962, a very strong thunderstorm passed
August were the cloudiest months, although the John - Davis Town (Putu district) in eastern Liberia.
period September to October h ad the same mean The wind had a hu rricane-like strength and created
rainfall as the three earlier months. January 1 9 70 an alm ost 1 00 m gap in the forest east of the settle
showed only five mornings with more than 50 % ment. Only around 20 % of the trees were still stan
c loud cover and all the afternoons were cloudless. ding after the storm h ad passed. In the N imba area
During the period January 1 2 to Feb ruary 6, 1 9 70 no such storms of s igns of earl ier ones have been
no clouds whatsoever were reported at any period of observed.
the day. The local cloud o r mist system that fre For certain short periods the dry Harmattan
quently can be observed around the highest parts of winds are strongly felt in the N i mba area. The wind
the ridge du ring the d ry season seems to be cor is weak , merely as a breeze, but is characterized by
related with the N E -winds (Fig. 1 1 ). its dust-filled, very d ry ai r. The v is ibility is low (a few
I v o ry C o a st L i b e ri a
DAY S Altitude
30
r-r- r-r- r-r--
r-r--1-r-
-_ -r-_r-
�
-
-r- 1---
�
-r-
I JOO
?
�
\�
= Air sinking
\
wn
�
15 -
�
-
A i r fo rced jA
Clouds diss �
u p w a rds /
�-·
.• 0<g tx'x
�Rx
_.
0 � �R:><�
A M PM
600
J F M A M J J A s 0 N D
VEGETATION
Earlier botanical exploration
Th e N imba Mts in what today is part of Guin ea and
th e Ivory Coast wer e during the French adm in is tra
tion subj ect to a s er ies of studies cover ing s ev eral
aspects of natural h istory (e.g. L amotte et al. 1 9 5 5).
A r emarkabl e work on the vegetation of the N imba
Mountains was pres ented by Sch n ell ( 1 9 5 2).
From th e Liber ian part of the N imba Mountains
botan ical collections have b een p erform ed by W .
Harl ey a t and around M t Bilumi (M t Bele). Und er
th e sponsorsh ip of th e N imba R es earch C o m m ittee,
h eaded by K . C urry-Lindahl, bo tanical collections
w er e carried out by P. Adams ( 1 964) and J. Adam
( 1 964- 1 9 65 and 1 9 70). Based upon h is ex ten s iv e
coll ections from all parts of t h e ridge, Adam ( 1 9 7 1 )
publish ed a work entitled 'Flare d escriptiv e d es
Mon ts N im ba'. During h is study of Lib erian h igh
forest trees, Voorhoev e ( 1 965) v isited th e N imba
M ts and mad e collec tions there.
ed forest in this case could be regarded as a single would be favourable for it throughout most of the
association of fluctuating composition. Richards country.
( 1 964 : 262) states : "The failure of any one species to The Liberian rain forest is p art of a rain forest belt
gain the upper hand in the m ixture may be due, as th at stretches from Ghana to Sierra Leone, and
Au breville suggests, to all the species having very which is a western extension of the huge lowland rain
similar ecological requirements and responding in a forest of the Congo basin (Fig. 1 5). A region with
very similar way to slight variations in the environ dry climate from Western Nigeria to Eastern Ghana
ment. As would be expected on grounds of probabili separates the two forested areas from each other.
ty, under these conditions the composition of these This break is commonly known as the 'Togo
mixed communities will fluctuate in space and prob Dahom ey gap'. The high forests of Liberia exhibit
ably in time as well." several features typical of tropical rain forests such
as a large number of species which are mostly
The Liberian rain forest woody plants, e.g. many tree species (Cooper &
Through human interference only 35 % of Liberia is Record 1 9 3 1 , Kunkel 1 96 5 , Voorhoeve 1 9 65). The
at present covered by high forest (Voorhoeve undergrowth is also mainly composed of woody
1 9 65 : 1 9), although climatic and edaphic conditions plants. The herbaceous plants are mainly represented
Table 5. The number of trees, 40 cm or larger in diameter, in a Table 6. The number of 15 different timber trees, 45 cm or
forest at 525 m altitude east of Grassfield. Total area in larger in diameter in East Nimba National Forest. Total area in
vestigated, 4 ha. (From Adam 1969.) vestigated, 565.2 acres. (Source: Lamco J . V. Op. Co.)
lies at approximately 40-45 m height above the Harungana madagascariensis (Guttiferae). Among
ground (Table 5). the taller species easily observed in older secondary
R em arkable is the dominance by the family forests, Ceiba pentandra, Pycnanthus angolensis and
Leguminosae. It is represented by seven genera, eight Elaeis guineensis m ay be mentioned.
species and 49 individuals in the inventory. The
species of this family constitute nearly 48 % of the Forest at altitudes above 1 000 m
total number of trees. Four fam ilies, Leguminosae, In the higher parts of the Nimba R ange a distinct
Rosaceae, Ochnaceae and Sterculiaceae together change is noticed in the physiognomy and floral
represent 85 % of the trees in the inventory. Adam composition of the forest. These parts, which are
( 1 969) also emphasizes the l arge differences in the very small and m ainly consist of the narrow ridges,
flora composition betw�en the Kitoma and the Nim are covered by a low single dominant Parinari ex
ba forests. The K itom a forest i s located 30 k m south celsa forest, reaching a height of 1 0-20 m . The
of the Nim ba Mts. In Kitoma 1 7 7 individuals of smaller trees have their ramifications close to the
Calpocalyx aubrevillei were found in an area of 1 5 . 2 ground, an d the field layer is usually built up by
ha, which gives 1 1 . 6 trees pe r h a. In N im b a 7 in herbs. At 1 200- 1 300 m altitude the Aframomum
dividual s were found which gives 1 . 7 trees pe r ha. herbs are typical. The single dominant Parinari
The figures for the Piptadeniastrum africanum are forest is by no means something specific for the
0.9 trees per ha in Kitoma, and 8 . 2 trees per ha in Nimba Mts but can be found at similar altitudes at
Nim ba. e.g. Mount Wutivi in western Liberia ( Kunkel
An inventory of the East Nimba N ational Forest 1 9 62 b), Mt Tonkoui in the Ivory Coast and Fouta
and adj acent parts h as been undertaken under the Dj allon in Guinea. From Uganda a rain forest com
leadership of T.A. Gorgla (1 9 6 9). This study in munity occurring between 4500 and 5000 feet i n.
cluded 15 species of trees that could be of commer which Parinari excelsa forms about 80 % of the
cial i nterest (Table 6). Although the study excludes canopy is reported by Eggeling ( 1 94 7).
A cta phytogeogr. suec. 59
Ecology of vascular epiphytes in West African rain forest 1 7
The canopy is not closed except in very minor sec species. When a doubt in the determination of
tions. The light intensity in the lower parts of the species has arisen, the epiphyte flora h as simply been
forest and at the ground is therefore considerably recorded without correlation to a specific
higher than in the forest at the base of the mountains. phorophyte. The list of phorophytes that briefly will
Schnell ( 1 9 5 2: 338) remarks : "On retrouve dans les be presented is dominated by medium to large sized
forets a Parinari excelsa du Nimba et des massifs trees. For each phorophyte a few properties that
voisins, un certain nombre de caracteres deja might be of importance for the epiphyte flora have
signales dans les forets montagnardes, par J. Lebrun, been added. The m ajority of these phorophytes are
H. Scaetta (1 9 7 3) , Schluter, Eidmann : faible hauteur included in the excellent · study of the Liberian rain
des arbres, tronc ramifie a quelques metres du sol, forest by Voorhoeve ( 1 96 5 ) .
reduction du nombre des especes, reduction du nom The properties of the phorophytes refer to tall
bre de s lianes, abondance des epiphytes (surtout specimens and it must be emphasized that m any of
C ryptogames), feuillage generalement persistant." these properties are rather vague and dependent on
On the crests at 1 200- 1 300 m altitude the environmental influence. The roughness of the bark
Parinari excelsa forest is in total dominance, but a refers to the basal parts of large branches, and the
little lower down at 1 000- 1 1 00 m trees of a rather terms low, medium and tall refer to the classes
large number of species can be found (Adam 1 0-20, 2 1 - 35 , 36 m or more, respectively. The
1 9 70 :204-207). phorophytes are presented in alphabetical order
Seasonal changes in the forest without regard to family.
In the Nimba forest, not much seasonal changes in
leaf-fall are noticed. The evergreen trees at the Phorophytes in high forest
various strata appear luxuriant at all times of the A lbizia glaberrima (Schum. & Thonn.) Benth.
(Leguminosae)
year because progressive leaf-fall occurs, while the
Large. Bark smooth. Crown open, spreading. Deciduous.
few deciduous trees appear dormant only for a
(DC.) MacBride (Leguminosae)
A lbizia zygia
rather short time. These deciduous trees are so few Medium. Bark rather smooth. Crown open, spreading.
and scattered that their presence does not alter the Deciduous.
face of the forest. Bernhard-Reversat et al. A lstonia booneiDe Wild. (Apocynaceae)
( 1 972 :2 1 8) reveal that the maximum leaf-fall of Large. Bark rather smooth, soft. C rown fairly open,
various species in the Ivory Coast do no coincide, heavily branched. Evergreen.
and there is no correlation with the precipitation, but A mphimas pterocarpoides Harms (Leguminosae)
rather with the incident radiation-a view that earlier Medium-large. Bark scaly. Crown dense. Deciduous. (Fig.
had been stressed by Hopkins ( 1 970) in a study of 1 6).
the seasonal growth in S Nigeria. Furthermore, the A nthocleista nobilis G.Don (Loganiaceae)
interspecific variation of the start and the length of Small. B ark with thorns. Crown open. Evergreen.
the leaf-fall period at the same locality also tend to A nthonotha fragrans (Bak.f.) Exell & Hillcoat
obscure the presence of these trees. Some of the (Leguminosae)
more common deciduous trees in the forest are .{Jom Medium-large. Bark scaly. Crown dense. Deciduous.
bax buonopozense, Chlorophora regia, Ceiba pen A ntiaris toxicaria(Rumph. ex Pers.) Lesch. var.
tandra, Daniellia ogea, Entandrophragma utile, Pip welwitschii (Engl.) Corner (Moraceae)
Large. B ark smooth, thick. Crown fairly open. Briefly
tadeniastrum africanum, R hodoghaphalon brevicus deciduous.
pe, Terminalia ivorensis, and Triplochiton scleroxy
Berlinia confusaHoyle (Leguminosae)
lon. In a phenological study of Chlorophora regia at Medium-large. Bark smooth or scaly. Crown rather dense.
Njala, Sierra Leone, Cole (1 968 : 76) found not only Evergreen.
an interspecific variation in leaf shedding time Bombax buonopozense Beauv. (Bombacaceae)
between trees growing within a radius of 1 00 yards, Medium. R ather smooth bark. C rown heavily branched.
but also a difference in the length of the leafless Deciduous.
period between male and female trees. Hutch. & Dalziel (Leguminosae)
Bussea occidentalis
Medium . Bark smooth. Crown dense. Evergreen.
The phorophytes (the host trees) Calpocalyx aubrevillei Pellegr. (Leguminosae)
As stated earlier the rain forest is very rich in tree Medium. Bark smooth. Crown small. Evergreen.
A. �. . . -
Fig. 1 6. Trees common in the Nimba area. (A) Heritiera pterocarpoides, (D) Lophira alata, (E) Ceiba pentandra,
uti/is, (B) Piptadeniastrum africanum, (C) Amphimas (F) Entandrophragma utile.
Terminalia ivorensis A. Chev. (Combretaceae) Cultivated for their edible seeds ("Almond tree") or as an
Large. Bark deeply grooved. Crown rather open. ornamental. Low-medium. Bark fissured. Crown
Deciduous. (Fig. I 7). spreading, open. Deciduous.
Large. Bark grooved and scaly. Crown open. Deciduous. The cacao tree. Low. Bark smooth. C rown dense.
(Fig. 1 7). Evergreen.
Composition and geographical distribu 5000 in Malaysia alone (Van Steenis 1 93 8). The
tion general poverty of the African rain forest has been
suggested to be a result of past climatic changes
The flora of Africa · is generally considered to be (Mildbraed 1 922). Stewart (Stewart & C ampbell
poorer in species than those of other tropical regions. 1 970: 1 1 ) explains the relative low number of orchid
The orchids, of which the majority are epiphytes, species in Africa : "Vast tracts of the land mass are
have been estimated to comprise two to three thou unsuitable as orchid habitats."
sand species in Africa south of the Sahara, including However, the poverty of epiphytic plants in the
the island of the Malagasy republic (Madagascar), African rain forest is by no means an exception.
and the islands of the Seychelles, Comoro, and the From Sarawak, Borneo, Richards ( 1 93 6 : 1 5- 1 6)
Mascarene groups (Stewart & Campbell 1 970). This remark s : "One of the most striking features of the
figure may be compared to the more than three thou Sarawak rain forest, especially when compared with
sand species of orchids recorded in the South that of tropical South America, is the poverty of the
American republic of Colombia alone. Another ex epiphytic vegetation both in species and individuals."
ample from the orchid family may illustrate the The vascular epiphytic flora of the investigated
relative poverty of species. In the Zaire (former area includes a total of 1 5 3 species (the facultative
Belgian Congo) there are a total of 4 1 4 species epiphytes excluded). The epiphytes may be divided
(Nihoul et al. 1 964), compared with an estimated into three major groups : ( 1) the Pteridophytes with
Fig. 1 9. Records of
Angraecum birrimense (O)
and Cyrtorchis monteiroae
(e).
Fig. 21. of
Records
Polystachya (0)
dalzielii
and P. leonensis ( •). (A)
Picket Hill (B) Mt
Kakouima (C) Loma Mts
(D) Wologisi Range (E)
Nimba Mts (F) Massif de
Dans (G) Mt C ameroon
39 spp., (2) the Orchids with 1 0 1 spp. and (3) the rare. The flowering time is expressed in two months
remammg species, called the Other vascular periods (when the most frequent flowering occurs).
epiphytes, with 1 3 spp. This grouping is followed in A brief description of the plant is added for the
the list and analyses. orchids only. The particulars of the flowers are, due
The orchid flora seems to be composed of three to their limited duration, mostly of minor importance
different elements : (A) Species of the rain forest, e.g. in the field work. In most cases the determination
A ngraecum birrimense and Cyrtorchis monteiroae, has to be based on vegetative characteristics only.
(B) Species of the deciduous forest, e.g. A nsellia The inflorescences which often remain on the plant
africana and Rangaiiris rhipsalisocia, (C) and for several years have proved useful. (A field key to
species restricted to high altitudes, e.g. Polystachya the epiphytic orchids of Ghana based on vegetative
dalzielii and Polystachya leonensis (Figs. 1 9 -2 1 ). A characteristics has been presented by Hall & Bowl
few species, e.g. Bulbophyllum inflatum, have only ing 1 96 9 , and in a key to the orchids of Zaire, former
been reported from the Nimba Mountains in Liberia. Belgian Congo, Tournay (1 9 55) used a similar
However this endemic character may only result technique.) Some species are difficult to determine
from insufficient knowledge (cf. Schnell 1 9 52 :407 even in a flowering state. The methods used to dis
and Curry-Lindahl 1 96 9 : 23). tinguish such species are discussed in connection
Some species with a ± wide distribution in tropical with the presentation of the species. The size of the
Africa, e.g. A ntrophyum immersum and plants may vary widely due to environment.
Nephrangisfiliformis are in West Africa restricted to In the following list the grouping previously
the Nimba region. Several other species have had presented is followed. Within each of the three
their previously known area of distribution extended groups the names of the genera and their species are
westwards, e.g. A ngraecum classensii, A ngraecopsis given in alphabetical order. The first set of my collec
elliptica, Begonia rubro-marginata, Bulbophyllum tions (with few exceptions), is kept in the Herbarium
magnibracteatum, B. pavimentatum, B. schimpera of the Institute of Systematic Botany at the Universi
num � Diaphananthe densijlora, Elaphoglossum ty of Uppsala (UPS). A second (almost complete) set
isabelense, and Stolzia repens (Figs. 2 2 -24). A total has been given to the Kew Herbarium (K). For each
of thirty-two species were recorded in Liberia for the species a voucher is quoted, e.g. D .J. 43 1 (UPS).
first time.
Pteridophytes
Asplenium L.
A. aethiopicum (Burm.) Becherer
Guin., S.L., Lib., N.Nig., Cam.
Harley { 1 95 5 :84) and Kunkel ( 1 962 :40) report it from
various parts of Liberia. Trunks and branches. Open
shade, humus deposits. Rare. D.J. 782 (K, UPS).
A. africanum Desv. Fig. 26. Drynaria laurentii, sterile and fertile leaves.
Guin., S.L., Lib., Jv.C. , Ghana, S.Nig.
Basal parts of large branches. Open shade, humus
deposits. Rare. D.J. 852 (K), 834 (UPS). The roots hold A. megalura Hieron. ex Brause
humus in a fashion similar to Microsorium punctatum, Guin., S.L., Lib., Jv.C., Ghana, Togo.
which it resembles at a distance. Middle - outer parts of large branches. Full sun, humus
deposits. Common. D.J. 780 (K, UPS).
A . barteri Hook.
Guin., S.L., Lib., Jv.C ., Ghana, C am. A. variable Hook. var. paucijugum (B allard) Alston.
Basal parts of the trunks or moss-covered rocks. Humid Guin ., S.L., Lib., Jv.C., Ghana, C am.
habitat. Heavy shade, varying substrate. Common. D.J. This taxon is often referred to as A splenium paucijugum
5 5 5 (K), 629 (UPS). Fig. 25. Ballard (Harley 1 95 5 :86, Kunkel 1 96 2 :43). Basal parts of
trunks of tall trees or on moss-covered rocks. Wet and
A. dregeanum K unze
humid habitats. Heavy shade, humus deposits. Rather
Guin., S.L., Lib., Jv.C ., Ghana, Togo, S.Nig., N.Nig.,
common. D.J. 5 89 K, 786 (UPS).
Cam.
On trunks in humid habitats. Heavy shade and humus
deposits. Rather common at altitudes above 1000 m. D.J. Da vallia S m .
Elaphog/ossum Schott
E. barteri (Bak.) C .C hr.
Guin., S.L., Lib., lv.C., S.Nig., Cam.
Basal parts of trunks. Heavy - open shade, large humus
deposits. Rare. D.J. 5 5 3 (K), 697 (UPS).
E. chevalieri C hrist.
Guin., S.L., Lib., lv.C.
B asal parts of moss-covered trunks. Heavy - open shade,
humus deposits. Restricted to altitudes above 1 1 00 m.
Rare. D.J. 642 (K, UPS).
E. isabelense Brause
Lib. (new), F.Po.
Middle - basal parts of large branches of tall trees. Open
shade - full sun, bark or minor humus deposits. Common.
D.J. 495 (K , UPS).
E. kuhnii Hieron.
S.L., Lib., Cam.
Basal parts of tall trees, or rotten logs. Humid habitats.
Heavy shade, humus deposits. Rare. D.J. 493 (K), 646
(UPS).
E. salicifolium (Willd. ex Kaulf.) Alston
Guin., Lib. (new), C am.
Widely distributed over the phorophyte, mostly in the
basal parts of the large branches. Full sun - open shade,
bark. Common. D.J. 644 (K), 825 (K, UPS).
Fig. 27. Nephrolepis biserrata.
Lomariopsis Fee
L . guineensis (Underw.) Alston humus deposits. Rather common. Often associated with
Guin., S.L., Lib., Iv.C . , Ghana, Togo, D ah., S.Nig., C am. Drynaria laurentii.D.J. 5 3 5 (K, UPS).
Reported from various parts of Liberia (Harley 1 95 5 :87,
Kunkel 1 9 6 2 : 5 2). Basal parts of trunks. Heavy - open Microgramma Presl
shade, bark. Rare. Rooted at the base of the phorophyte,
M. o wariensis (Desv.) Alston
which it climbs. Very tightly attached to the bark by short
Guin., S.L., Lib., Jv.C., Ghana, Togo, N.Nig., S.Nig.,
rootlets of the rhizome. After some time it normally loses
Cam.
contact with the ground. D.J. 7 7 7 (K), 482 (UPS). Fig.
On trunks and branches. Open shade, minor humus
1 1 6.
deposits. Common on trees and palms around villages,
less frequent in the high forest. From Liberia a new form,
Loxogramme ( B i u m e) Presl f. nana, has been described (Kunkel 1 962:37- 3 8) . D .J.
L. lanceolata (Sw.) Presl 5 6 7 (K, UPS).
Guin., S.L., Lib., Iv.C ., Ghana, N.Nig., S.Nig., Cam.
Mainly on the trunks. Open shade, minor humus deposits. Microsorium L i n k
Rare. D.J. 8 3 3 (K), 742 (UPS).
M . punctatum (L.) Cope!.
Guin., S.L., Lib., Ghana, S.Nig., C am.
Lycopodium L. Basal - middle parts of large branches. Open shade - full
L . mildbraedii Hert. Syn : L. dacrydioides Bak. sun, minor humus deposits. Common. The debris held by
Guin., S.L., Lib. (new), Cam. the root system is often utilized by other epiphytes. D.J.
On moss-covered branches and trunks. Open shade, 794 (UPS).
minor humus deposits. Common at altitudes above 1 000
m. D.J. 503 (K, UPS).
Nephrolepis Sc hott
L . warneckei (Hert.) Alston N. biserrata (Sw.) Schott
Guin., S.L., Lib., Jv.C., Cam. Guin., S.L., Lib., Jv. C ., Ghana, Togo, S .Nig.
In the outer - middle parts of large branches. Full sun, Epiphytic or terrestrial, equally common in both habitats.
As epiphyte mainly at the basal part of the large branches. Tectaria Cav.
Open shade - full sun, large humus deposits. Common in T. angelicifolia (Sebum.) C opel.
secondary forests (particularly at the leaf bases of oil Guin., S.L., Lib., Iv.C., Ghana, N .Nig., S.Nig., C am.
palms). Rare in the high forest. Used by the natives in In the basal parts of tall trees (or on rocks). Frequently on
treatment of snake bites and urinary complaints (Harley the trunks of Cyathea manniana. Heavy shade, on
1 94 1 ). D.J. 765 (K). Fig. 27. various kinds of substrate. Common. D.J. 1 05 9 (UPS).
N. undulata (Afzel. ex Sw.) J.Sm. T. fernandensis (Bak .) C.Chr.
Guin., S.L., Lib., Iv.C., Ghana, Togo, Dab., N.Nig., Guin., S.L., Lib., Iv.C . , Ghana, Togo, S.Nig., C am.
S.Nig., Cam. Basal parts of large trees. Heavy shade, humus deposits.
Basal - middle parts of large branches. Open shade - full Only in very humid habitats. Rare. D.J. 795 (K).
sun, minor - large humus deposits. Occurs in several
epiphyte communities, particularly with Platycerium Vittaria S m .
ferns. Common. D.J. 707 (K, UPS). V. guineensis Desv.
Guin., S.L., Lib., Jv.C., Ghana, Togo, S.Nig., C am.
On all parts of the trees, but mainly in the basal and
Oleandra Cav.
0. distenta Kunze
middle parts of large branches. Open shade, minor humus
deposits. Often associated with Microsorium punctatum.
Guin., S.L., Lib., Iv. C ., Ghana, Togo, S.Nig., Cam.
Common. D.J. 762 (UPS).
Around the basal ramifications of the tall trees. Open
shade, humus deposits. Common in the high forest and on Xiphopteris K a u lf.
the trunks of the oil palm in farmlands. The humus held X. oosora (Bak.) Alston
by this species is utilized by several other epiphytes. D.J. Guin., S.L., Lib. (new), C am., F.Po.
805 (K), 427 (UPS). Fig. 80. Basal - central parts of large branches. Open shade,
humus deposits. At altitudes above 1 200 m. Rather com
Phymatodes Presl mon. D.J. 826 (K), 5 1 4 (UPS).
P. scolopendria (Burm.) Ching.
X. serrulata (Sw.) Kaulf.
Guin:, S.L., Lib., Iv.C., Ghana, Dab., Togo, N .Nig.,
S.L., Guin., Lib., Jv.C., F.Po.
S.Nig., Cam., F.Po.
Middle parts of large branches. Open shade, minor humus
Basal - middle parts of large branches in high forest. On
deposits. R are. D.J. 592 a (UPS).
trunks of trees and oil palms around villages. In high
forest in open shade, bark - minor humus deposits. In X. villosissima (Hook.) Alston
secondary forests it is found in a wider range of habitats. Guin., S.L., Lib., Jv.C., C am., F.Po.
Common. D.J. 5 3 2 (K), 8 3 1 (UPS). Basal and middle parts of large branches. Open shade,
minor humus deposits. At altitudes above I 1 00 m. Rare.
Platycerium Desv. D.J. 5 1 8 (K), 827 (UPS).
P. angolense Welw. ex Hook.
Guin., S.L., Lib., Iv.C., Ghana, N.Nig., S.Nig. Orchids
Middle parts of large branches of emergent trees or on the Aerangis Rchb.
trunks of trees left in farmlands. Full sun, bark. Common A . biloba (Lindl.) Schltr.
on large Triplochiton scleroxylon trees in the Yekepa Sen., Guin., S.L., Lib., Iv.C., Ghana, Togo, N.Nig.,
community. Rare in the high forest. In Iv.C . this species S. Nig., Cam.
occurs mainly in the deciduous forests in the northern part Basal or middle parts of large branches. Full sun - open
of the country (Boyer 1 964 ). D.J. 809 (UPS). Fig. 85. shade, bark - minor humus deposits. �Common. Growing
P. stemaria (P. Beauv.) Desv. tips of roots orangish-pink (Piers 1 95 9, Moir 1 963, San
Guin., S.L., Lib., Jv.C., Ghana, Dab., N.Nig., S.Nig., ford 1 968). Stem short, woody. Leaves crowded at the
C am. apex of the stem, 8- 1 8 cm, oblanceolate, curved and un
In its ecology very similar to P. angolense. Rare in high equally bilobed, dark green. Inflorescence longer than the
forest, more common on tall trees in secondary forest. In leaves, many flowered, somewhat pendulous. Flowers
Iv.C., this species occurs mainly in the secondary white, star-like. June-July. D.J. 5 62 (UPS). Fig. 28.
evergreen forest in the southern part of the country, and is A . laurentii (De Wild.) Schltr.
± absent in the primary rain forest (Boyer 1 964). D.J. 79 1 Lib., Ghana.
(UPS). Fig. 84. Anywhere on the large branches. Full sun, bark. Rare.
Erect, 10- 1 5 cm. Stem short, woody. Leaves strap-shaped,
Pyrrosia M i rb. arranged in a fan-shaped manner, acutely v-shaped in
P. mechowii (Hieron.) Alston cross-section, rather thick and stiff. Inflorescence longer
Guin., Lib. (new), N.Nig., C am . than the leaves, zigzag-shaped rachis. Flowers white, with
Middle parts o f large branches. Open shade - full sun, a long spur. Jan.-Febr. The zigzag rachis distinguishes it
minor humus deposits. Humid habitats, e.g. swamp from Rangae'ris muscicola. Plectrelminthus caudatus has
forests. Almost exclusively on Mitragyna ciliata. Rare. also a zigzag rachis, but leaves of a different shape. D J .
D.J. 808 (K, UPS). 4 4 9 (K), 4 1 1 (UPS).
Ancistrochilus Ro lfe
A. rothschildianus O'Brien
Guin., S.L., Lib., Iv.C., S.Nig., C am.
Basal or central parts of large branches, occasionally on
moss-covered trunks. Open shade, humus deposits.
Rather common. Erect, 20-30 cm. Pseudobulbs crowded,
conical - pyriform, 2-3 cm high, up to 5 cm in diameter,
longitudinally sulcated, green. Leaf 1 5-30 cm, lanceolate
oblanceolate, thin, ribbed, dark green. Inflorescence long,
curved. Flower large (8 cm in diam.). Oct.-Nov. D.J. 703
(UPS). Fig. 29.
Ancistrorhynchus F i net
A. capitatus (Lindl.) Summerh.
S.L., Lib., S.Nig.
Basal parts of large branches or on trunks. Open shade -
full sun, bark - minor humus deposits. Common. Erect,
25-30 cm. Stem short, woody. Leaves crowded, curved,
20-50 x 1 .5-3 cm, with a few teeth at apex. Inflorescences
short, dense, on the stem below the leaves, long persistant.
Flowers small, white. July-Aug. D.J. 5 3 1 (K, UPS).
A. cephalotes (Rchb.f.) Summerh.
Guin., S.L., Lib., Iv.C., Ghana, N.Nig., S.Nig.
Trunks or the middle parts of large branches. Full sun,
bark. Common. Erect or scrambling. Stem long, woody,
twisted, covered with old leatbases and inflorescences.
Leaves 8-20 cm, linear, apex bilobed. Inflorescences on
the stem below the leaves, short, with a very large number
of flowers. Flower white, small. June-July. D .J. 524 (K,
Fig. 28. Aiirangis biloba. UPS). Fig. 1 1 0.
A. clandestinus (Lindl.) Schltr.
S.L., Lib., lv.C., Ghana, N.Nig., S.Nig., C am.
Trunks or basal parts of larger branches. Full sun, bark.
Rare. Erect, in the Nimba area a small to medium-sized
plant, stem short, woody. Leaves 5-8 x 0.5- 1 .0 cm, stiff
and fleshy, leathery, apical lobes of the leaf very acute and
Fig.29. A ncistrochilus
rothschildianus.
minor humus deposits. Rare. Prostrate, 1 5 -20 cm. deposits. R are. Erect, 8- 1 6 cm. Pseudobulb 1 -2 cm, light
Pseudobulb 4-5 cm, conical or ovoid, 3-4 angled, velvety, green, velvety. Leaf 6- 1 2 cm, oblanceolate-narrowly
light green. Leaf 1 2- 1 5 cm, flat, stiff, oblong, green. oblong, acute, green. Inflorescence about the same length
Inflorescence 40-60 cm, bracts less than 1 5 mm, overlap as the pseudobulb and the leaf, arching, densely flowered.
ping each other. Flower small, lip hairy, sepals strongly Flower small, yellow or pink. June-July. D.J. 5 8 5 (K), 5 8 6
reflexed, yellow with purple spots. July-Aug. Difficult to (UPS).
distinguish from B. schinzianum and B. phaeopogon B. imbricatum Lindl.
without flowers. The size of the bracts gives a possibility. S.L., Lib., Jv.C., Ghana, S.Nig., C am.
D.J. 8 5 6 (UPS). In the middle parts of large branches. Open shade, minor
B. cochleatum Lindl. humus deposits. Rare. Erect, 1 5-25 cm, stout. Pseudo
Guin., S.L., Lib., lv.C ., N.Nig., S.Nig., Cam. bulbs 3-4 cm, ovoid, green, scattered along the rhizome.
Basal or middle parts of large branches. Open shade, bark Leaf 1 2-20 cm, linear - oblong. Inflorescence longer than
or in minor humus deposits. Common. Erect, 1 2-24 cm. the leaves, erect stout. Rachis fleshy, dark purple. Flower
Pseudo bulbs scattered, 3 - 1 0 cm, narrowly cylindrical, small with purple spots. Sept.-Oct. D .J. 7 1 0 (K), 678
bluntly quadrangular, brownish-green with brown or pur (UPS).
ple stripes or dots. Leaf 8- 1 4 x 0.5- 1 . 2 cm, strap-shaped, B. injlatum Rolfe
green, sometimes tinged purple. Inflorescence up to 30 cm S.L., Lib.
long, curved. Flowers small, orange-red. Sept.-Oct. D .J. Trunks or in the basal parts of large branches. Heavy -
63 1 (K, UPS). _ open shade, minor humus deposits. At altitudes above
1000 m. Rather common. Pendulous, 8-20 cm. Pseudo
B. cocoinum Batem. ex Lindl. bulb 1 -3 cm, ovoid quadrangular, rhomboid in cross sec
S.L., Lib., Ghana. tion, green. Leaf 7-20 x 1 .5-5 cm, lanceolate - oblong or
B asal and middle parts of large branches. Open shade, elliptical, leathery, green. Flower small, yellow-green.
humus deposits. Rather common. Erect, 1 5-30 cm. Jan.-Feb. Easy to recognize. The plant grow upside down.
Pseudobulbs crowded, 2-4 cm high, 1 . 5-2.0 cm broad, This species is very similar to B. comatum Lindl., in a ster
ovoid, sharply 3-4 angled, pale yellow. Leaf lanceolate or ile as well as fertile state. The latter has not been recorded
oblanceolate, 1 0-20 cm, acute, not stiff. Inflorescence in the Nimba Mts but from Mt Momi in the Ivory Coast.
long, arching, densely flowered. Flowers small, creamy In the field work specimens without flowers that could be
white. Aug.-Sept. D .J. 54 1 (K, UPS). any of the two species have been recorded as B. injlatum.
B. congolanum Schltr. D.J. 7 1 7 (K, UPS).
Guin., S.L., Lib., Iv.C., Ghana, N . Nig., S.Nig. B. intertextum Lindl.
Middle parts of large branches. Full sun, bark or minor S.L., Lib., Ghana, N .Nig., S.Nig., Cam.
humus deposits. Rather common. Erect 8- 1 4 cm. Middle parts of large branches. Full sun, bark. Common.
Pseudobulb 2-4 cm, ovoid - conical, 3-4 angled, light Erect, 2-5 cm. Rhizome slender, forming mats. Pseudo
green-yellow. Leaf 6- 10 cm, strap-shaped, green. bulbs scattered, up to one cm, elliptical - ovoid, green.
Inflorescence erect, somewhat curved, of about the same Leaf 1 -2.5 x 0.5- 1 .0 cm, elliptical, light green.
length as the leaves. Flower, small creamy-white. Oct. Inflorescence 1 4-20 cm, slender. Flower minute, creamy
Nov. D.J. 7 1 4 (K, UPS). Fig. 58. white. Oct.-Nov. A somewhat different form of this
B. distans Lindl. species occurs : Pseudobulb of similar shape and size as
Lib., Jv.C ., Ghana, S.Nig., C am. above, but with a purplish colour. Leaf strap-shaped, 1 -4
Middle or basal parts of large branches. Full sun - open x 0.3-0.5 cm, dark greenish-purple. Inflorescence much
shade� minor humus deposits. Rather common. Ascen shorter, 4-8 cm. Flower minute, purplish. Oct.-Nov. D.J.
ding, 1 0- 1 5 cm. Pseudobulb depressed, 2-3 cm, conical 687, 6 9 1 (K, UPS).
ovoid, yellow-green. Leaf 8- 1 2 cm, oblong, flat, stiff, B. josephii (K unze) Summerh.
green. Inflorescences (often two) erect, slender, 30-50 cm. Guin., S.L., Lib. (new), lv.C., S.Nig., C am.
Bracts 6- 10 mm. Flower small, dark purple, lip densely Basal parts of large branches. Open shade and humus
hairy. June-July. D.J. 8 5 5 (K), 857 (UPS). deposits. Rare. Erect, 1 2-20 cm high. Pseudobulbs
B. falcatum (Lindl.) Rchb.f. crowded, 2-3 cm, narrowly ovoid-conical ovoid, velvety,
Guin., S.L., Lib., lv.C., Ghana, S.Nig., C am., F.Po. redbrown. Inflorescence long, slender, arching. Flower
Middle part uf large branches. Open shade, minor humus purplish-violet. July-Aug. Other species with redbrown
deposits. Common. Erect. Pseudobulb 1 . 5-5 cm long, pseudobulbs are B. winklerii, and B. buntingii D.J. 44 1
ovoid, 3-4 angled, green. Leaves 3- 1 2 x 0.6-2 cm, oblong (K), 699 (UPS).
lanceolate or oblanceolate. Inflorescence very long, up to B. linderi Summerh.
50 cm. Rachis 0.5-0.8 cm broad. Flowers small, yellow S. L., Lib., Jv.C .
purple. Oct.-Nov. D.J. 668, 670 (K), 6 7 1 (UPS). Basal o r middle parts o f large branches. Open shade,
B. jlavidum Lindl. bark. Common. Erect, 1 2-20 cm. Pseudobulb 2-3 cm,
Guin ., S.L., Lib., Iv.C., Ghana, S.Nig., Cam. ovoid, triangular, with a shiny texture, green. Leaf 8- 1 6
Middle part of large branches. Open shade, minor humus cm, oblong - lanceolate, green, often red coloured along
the margin. Inflorescence erect, the apical part' with the B. oreonastes Rchb. f.
white fleshy rachis is held in a horizontal position. Guin., S.L., Lib., Iv.C ., Ghana, N . Nig., S.Nig., C am.
Flowers white. Oct.-Nov. D .J. 6 7 5 (K, UPS). Middle part of large branches. Open shade - full sun, bark.
The most common epiphytic orchid in the Nimba area.
B. lucifugum Summerh.
Erect, 6- 1 0 cm. Pseudobulbs spaced out along the
S . L., Lib. (new).
rhizome, 1 -3 cm, ovoid, quadrangular, yellowish-green.
Basal or middle parts of large branches. Open shade,
Leaves 2-6 cm, oblong, leathery, green. Inflorescence
minor humus deposits or on bark. Rare. Erect, 1 0-20 cm.
longer than the bulb and leaves together. Rachis narrowly
Pseudobulb 2-4 cm, conical - ovoid, obtusely
winged, flowers yellowish with purple lihes, lip purple.
quadrangular, surface wavy or verrucate, greenish with
March-April. This species is vegetatively similar to B.
purple spots. Leaf strap-shaped, green, 1 0-22 x 0.6-0.9
zenkeranum Kraenzl. The latter species has been recorded
cm. Inflorescence recurved, rachis flattened, green with
from the Kitoma Range, south of the Nimba area.
purple dots, crinkled at the edges. Flower small. Feb.
Specimens without flowers that could be any of the two
March. D.J. 744 (K), 730 (UPS).
species have been recorded in the field work as B.
B. lupulinum Lindl. oreonastes. D.J. 640 (K, UPS).
Guin., S.L., Lib. (new), Ghana, N.Nig., S.Nig. B. pavimentatum Lindl.
Middle parts of large branches. Full sun, bark - minor Lib. (new), S.Nig.
humus deposits. Rare. Erect, 1 5-20 cm. Pseudobulb 4-6 Middle parts of large branches. Open shade, bark. Rare.
cm, rectangular, yellow green. Leaf 8- 1 8 cm, oblong, Erect, 1 2- 1 6 cm. Pseudobulbs crowded, about 2 cm long,
green. Inflorescence erect. Bracts large, with numerous ovoid, obscurely 2-3 angled, velvety green. Leaf 6- 1 4 cm,
short black or purplish hairs. Flower small, yellow with oblong- elliptical, dark green. Inflorescence equal with or
red spots. Jan.-Feb. D.J. 45 1 (K, UPS). longer than the leaf, thin, peduncle longer than the rachis.
B. magnibracteatum Summerh. Flower small, purplish. Oct.-Nov. D.J. 679 (UPS).
Lib. (new), Ghana, S.Nig. B. phaeopogon Schltr.
Middle parts of large branches. Open shade, bark. Rare. Lib. (new), Iv.C., Ghana, S.Nig., Cam.
Erect. Pseudobulbs 1 .5 -2.0 cm long, ovoid, 3 -angled. Outer parts of large branches. Open shade, minor humus
Leaves 5-7 x 1 .5 cm, stiff. Inflorescence of about the same deposits. Rare. Erect, 1 2- 1 6 cm. , Pseudobulbs erect,
length as the plant. Rachis 0.4-0. 7 cm broad with bracts crowded, 2-3 cm, ovoid, 3-4 angled, light green. Leaf
as broad as the rachis. When old and dry the rachis has a oblong- lanceo1ate, 7- 1 4 cm, stiff, green. Inflorescence 30-
characteristic obovate shape ending in a short acute apex. 60 cm. Bracts more than 1 5 mm long, overlapping.
Flowers small, purplish. April-May. D.J. 489 (UPS). Flower small, tepals yellow with red spots. Lip with long
hairs, mobile. July-Aug. This species is hardly dis
B. maximum (Lindl.) Rchb. f. tinguishable from B. schinzianum when in a flowerless
S.L., Lib., Iv.C ., Ghana, S.Nig. state. D .J. 8 1 1 (K ), 106 1 (UPS). Fig. 54.
Middle part of large branches. Open shade - full sun, bark.
B. recurvum Lindl.
Rather common. Erect, 1 2-20 cm, stout. Pseudo bulbs 3-7
S.L., Lib., Cam., F.Po.
cm, ellipsoid - conical ovoid, acutely quadrangular, green,
Middle parts of large branches. Open shade, minor humus
scattered along the woody and stout rhizome. Leaf 7- 1 0
deposits-bark. Rare. Erect. Pseudobulbs 1 - 1 .5 cm long,
cm, oblong - elliptical, stiff, dull green. Inflorescence 1 2-40
ovoid or conical ovoid, obscurely 4-angled, green to brow
cm, rachis 1 .0- 1 .5 cm broad, margins markedly undulate.
nish green. Leaves elliptical, 3-6 x 1 -2 cm, acute or obtuse.
Flowers minute, arranged along a central line of the
Inflorescence erect, 8- 1 3 cm, longer than the leaves,
rachis. Sept.-Oct. D.J. 656 (K, UPS).
recurved in the upper flowering � half. Flower small,
greenish-yellow. Oct.-Nov. D.J. 659 (UPS).
B. melanorrachis (Rchb. f.) Rchb. f. ex De Wild.
Guin., S.L., Lib., Ghana, S.Nig. B. rhizophorae Lindl.
Middle part of large branches. Open shade and minor S.L., Lib. (new), Jv.C., Ghana, S.Nig., C am.
humus deposits. Rare. Erect. Pseudobulbs 1 .5-3.5 cm On branches of small trees close to water. Open shade -
long, narrowly ovoid. Leaves 4- 1 0 x 0.7- 1 . 2 cm, oblong full sun, bark . Rare. Pseudobulb 2-3 cm long, elliptical.
oblanceolate, obtuse. Inflorescence 4-6 cm, rachis dark Leaves oblong, ± as long as the pseuodobulbs.
purple. Flowers crowded, small. Oct.-Nov. D.J. 648 (K, Inflorescence 6- 1 0 cm, erect. Flower small, yellowish
UPS). white. Oct.-Nov. D.J. 748 (K).
B. nigritianum Rendle B. saltatorium Lindl.
S.L., Lib., Iv.C., Ghana, S.Nig. -S.L., Lib. (new), Ghana.
B asal or middle parts of large branches. Open shade, Basal or middle parts of large branches. Open shade, bark
minor humus deposits. Rare. Erect. Pseudobulbs 2-4 cm - minor humus deposits. Rare. Prostrate, lying on the sub
long, conical ovoid, velvety, dark green. Leaf narrowly strate, 4-7 cm. Pseudobulbs crowded, 1 -2 cm, ovoid,
strap-shaped, I 0- 1 5 x 0. 7- 1 .0 cm. Inflorescence erect, 10- flattened, obtusely 3-4 angled, green. Leaf 3-5 cm, falcate,
1 5 cm long, flowering almost to the base. Flower small, oblong-elliptical, dark green. Inflorescence shorter than
white. Oct.-Nov. D.J. 693 (K), 669 (UPS). the plant. Flowers small, dark purple, lip hairy, almost all
Habenaria Wi l l d .
H. leonensis Dur. & Schinz
Guin., S.L., Lib., Iv. C .
Basal parts of large trees. Full sun - open shade, humus
deposits. Rare. In the investigated area epiphytic, but in
the grasslands of Guineean Nimba common as terrestrial.
Erect, up to 30 cm high. Leaves 6- 1 2 x 1 -2.5 cm, narrowly
lanceolate. Inflorescence with a few flowers. Flower white.
July-Aug. In a sterile state hard to separate from other
Habenarias. D.J. 5 80 (K, UPS).
H. procera (Sw.) Lindl.
S.L., Lib., Iv.C., Ghana, S.Nig., Cam.
On trunks of trees, but more often of oil palms. Full sun -
open shade, humus deposits. Rare. Erect, up to 60 cm
high. Leaves lanceolate, 1 2-20 x 2-3 .5 cm. Inflorescence
densely flowered. Flowers white. July-Aug. In a sterile
state hard to separate from other Habenarias. D.J. 595
(K, UPS).
Liparis L. C. R i ch.
L. caillei Finet
Guin., S.L., Lib., N.Nig., C am.
Basal parts of trunks. Open shade, humus deposits. Rare.
Pendulous. A minute plant. Total size 3-6 cm, in
florescence excluded. Pseudobulb 1 2-20 mm, globose,
green. Leaves 2-4 cm, ovate or lanceolate, green.
Inflorescence 1 4- 1 8 cm long, pendulous, rachis winged,
many flowered. Flower pale yellow. June-July. The leaf
and pseudobulb disappear among the debris in which the
Fig. 3 7. L iparis caillei. plant occurs. Even when flowering the inflorescence is
hard to observe. D.J. 5 2 1 (UPS). Fig. 3 7 .
L. nervosa (Thun.) Lindl.
Genyorchis S c h ltr. Syn. : L. guineensis Lindl., and L. rufina (Ridl.) Rchb. f.
G. pumila (Sw.) Schltr. ex Rolfe.
S.L., Lib., Iv.C., Ghana. Sen., Port.G., Guin., S.L., Lib., Iv.C., Ghana, D ah.,
Middle or outer parts of branches on small trees or N.Nig., S.Nig., Cam., F.Po.
shrubs. Full sun, bark. Rare. Erect, 2-3 cm. Pseudobulb In the forest only as an epiphyte, elsewhere often
0.6- 1 .5 cm, ovoid, quadrangular. Leaf 1 -2 cm, ligulate - terrestrial. In the lowest parts of trunks. Open shade - full
oblong, green. Inflorescence slender, 5 -6 cm. Flower sun, humus deposits. Rare. Erect, 1 5-25 cm. Stem with a
minute, Polystachya-like, white-pale green. March-April. ± swollen base. Leaves 5-20 cm, lanceolate, thin, with un
This small inconspicuous plant is similar to a Bulbo dulating margins, yellow-green. Inflorescence erect, longer
phyllum sp. D.J. 485 (K, UPS). than the leaves. Flowers yellow. July-Aug. D.J. 5 5 6 (K,
UPS).
Graphorchis Tho u.
G. lurida (Sw.) 0 . Ktze.
Sen., Port.G., Guin., S.L., Lib., Iv.C., Ghana, N .Nig., Listrostachys Rchb. f.
S.Nig., C am. L. pertusa (Lindl.) Rchb. f.
Middle parts of large branches. Full sun, bark. Common. S.L., Lib., Iv.C., Ghana, S.Nig.
Erect, up to 50 cm. Pseudobulbs crowded, 5 - 1 0 cm, con Middle parts of large branches. Full sun - open shade,
ical - ovoid, yellow, longitudinally wrinkled, with black bark . C ommon. Erect, 1 5-25 cm. Leaves arranged in a
transverse stripes. Leaves 4-6, in a dense tuft at the tip of fan-shaped manner on a short, woody stem, 1 5 -20 x 1 -2
the mature pseudobulb, lanceolate, up to 40 cm, thin, cm, strap shaped, unequally bilobed at apex, leathery, v
ribbed. Inflorescence from the base of the pseudobulb shaped in cross-section. Inflorescence 1 0-25 cm long,
appearing before the leaves, 1 5-50 cm, paniculate. Flower often longer than the leaves. Flowers small, white,
small, yellow and brown. Jan.-Feb. The yellow pseudo arranged in two dense rows along the rachis. Aug.-Sept.
bulb, surrounded by a dense tuft of white erect aerial This plant is of a shape similar to A i!"rangis laurentii and
roots, makes identification easy even when leaves or in Rangae'ris muscicola but has much more fleshy leaves.
florescences are missing. D J . 722 (UPS). Fig. 6 1 . D.J. 700 (K), 443 (UPS). Fig. 3 8 .
Nephrangis S u m merh.
N. filiformis (Kraenzl.) Summerh.
Lib.
Middle or outer : parts of large branches. Open shade -
full sun, bark. Common. Erect when small, pendulous
with increasing size, 30-40 cm. Stem slender, often
branched. Leaves 2-8 cm long but only 1 -2 mm broad,
terete, much curved, acute. Inflorescence very short, few
flowered. Flower small, with a white bilobed lip. April
May. Two other species have terete leaves : T. tridentata
and A ngraecum subulatum. The shape of the leaves
separates it from Tridactyle tridentata, and the pendulous
growth separates it from Angraecum subulatum. D.J. 529
(K, UPS). Fig. 5 9.
Fig. 3 9 . Plectrelminthus caudatus.
P/ectrelminthus R afi n .
Polystachya H ook.
P. caudatus (Lindl.) Summerh.
P. adansoniae Rchb. f.
Guin., S.L., Lib., Iv.C., Ghana, N.Nig., S.Nig . , Cam.
Guin., S.L., Lib., Iv.C ., Ghana, N . Nig., S.Nig., C am.
Middle or outer parts of large branches. Full sun, bark.
Middle or outer parts of large branches. Full sun, bark -
Rather common. Erect, stout, 20-30 cm . Stem short,
minor humus deposits. Common. Erect, up to 20 cm high.
woody. Leaves 1 0-30 cm, arranged in a fan-like manner,
Pseudobulb ellipsoid, 2-4 cm, light green. Leaves strap
oblong or elliptical oblong, almost equally bilobed at apex.
shaped, 8- 1 6 cm, light green. Flowers small, yellow,
Inflorescence longer than the leaves, erect, rachis zigzag
arranged in a spike. May-June. D.J . 4 8 5 , 8 5 8 (K), 437
shaped. Flowers large, white, spur long, spirally twisted.
(UPS).
Large capsules. April-May (Oct.-Nov.). D.J. 650 (UPS).
Fig. 39. P. affinis Lindl.
Guin., S.L., Lib., Iv.C., Ghana, S.Nig.
More or less restricted to trunks. Open shade - full sun.
Podangis Sch ltr.
P. dactyloceras (Rchb. f.) Schltr. Rather common. Growing in a vertical position on trunks.
Guin., S.L., Lib (new), Ghana, N .Nig., S.Nig., C am.
.
Pseudobulbs depressed, crowded, 2.5-4.5 cm in diameter,
Middle parts of large branches. Full sun, bark minor -
orbicular or broadly elliptical, green. Leaf 1 0-20 cm,
humus deposits. At altitudes above 800 m. Rare. Erect, 7- oblanceolate, dark green. Leaf and paniculate in
12 cm. Leaves in a fan-like arrangement on top of a short florescence arching. Flower yellow with red markings.
woody stem, 6- 1 0 cm, lanceolate, fleshy, green. Jan-Feb. D.J. 450. (K, UPS).
Inflorescence arises below the leaves, and does not exceed P. dalzielii Summerh.
the leaves in length. Flowers 5 - 1 0, semi-transparent, Guin ., S.L., Lib., Jv.C.
white. Feb.-March. C an be mistaken for a Bolusiella Middle or outer parts of large branches. Full sun, minor
talbotii or a Rangaeris rhipsalisocia. The first, however, humus deposits. At altitudes over I 000 m . Common. A
has a long thin inflorescence that exceeds the length of the small plant. Pseudobulb around I cm high, conical, very
leaves, the latter has sickle-shaped leaves. D .J. 42 1 (UPS). hard, green. Leaf 2-6 cm, ovate-lanceolate, thin, purplish-
... · .•
· �.
9
·•·.. ·· . .··:. < .··. ·
F� . . ..
Fig. 4 7. Tridactyle bicaudata.
i !� / / , �
: •... ·, :
(j CJ D�
S.L., Lib., Iv.C., S.Nig., C am .
Middle o r outer parts o f large branches. Full sun, bark.
Common. Climbing or hanging. A rather large plant,
branching in many directions. Stem stout, woody. Leaf 2-
8 cm, oblong or elliptical oblong, broadly and obtusely
Fig. 46. R hipidog/ossum paucifolium. (A) Flowering bilobed at the apex, flat, somewhat fleshy but stiff,
plant, (B) Flower with lip removed, front view, (C) Dorsal leathery. Inflorescences less than 0.5 cm, carrying 1 -3
sepal, (D) Lateral sepal, (E) Petal . (F) Lip, (G) Column flowers. Flower small, yellow-brown. Oct.-Nov. D.J. 676
with anther cap and pollinia removed, side view, (H) (K, UPS).
Rostellum, front view, (I) Pollinium. (From Johansson T. armeniaca (Lindl.) Schltr.
1 974.) Guin., S.L., Lib., Ghana.
Basal parts of large branches. Open sh ade, bark - minor
humus deposits. Common. Erect, often branching and
creating dense 'stands'. Stem long, woody in the basal
part. Leaf 5- 1 2 x 1 .0- 1 .5 cm, oblong - lanceolate, un
equally bilobed at apex, flat, rather thin. Inflorescence
Rhipidoglossum Sch ltr.
mostly below the leaves. Flowers small, yellow or apricot.
R. paucifolium Dick Johanss.
Feb.-March. T. fusifera Mansf. and T. tridactylites are
Lib. (New).
vegetatively similar to this species. The first species has
B asal parts of large trees and on small trees and shrubs.
not been observed although F.W.T.A. 1 968 3 : 1 , gives two
Open shade, bark. Rare. Stem very short, with a very
records from the Nimba area. The latter species only oc
large root system. Leaves few, 4-6 x 1 .5-2.0 cm, oblong
curs at altitudes above 1 000 m where T. armeniaca is
lanceolate. Inflorescences pendulous. Flowers crowded,
rare. D.J. 452 (K , UPS).
semi-transparent, pale green. Aug.-Sept. This species is
only known from the Nimba area (Johansson 1 974). D.J. T. bicaudata (Lindl.) Schltr.
5 72 (K, UPS). Fig. 46. S.L., Lib., Iv.C., Ghana, N.Nig.
Middle or basal parts of large branches. Open shade, bark
Stolzia Schltr. - minor humus deposits. Rather common. Erect. 30-50
S. repens (Rolfe) Summerh. cm. Stem long, woody. Leaf 1 0-20 x 1 -2 cm, narrowly
Lib. (new), S.Nig., C am. ligulate, unequally bilobed. Inflorescence from the leaf ax
Middle or outer parts of large branches. Open shade - full ils or below the leaves. Flower pale yellow-orange. Oct.
sun, minor humus deposits. R are. Prostrate. Pseudobulb Nov. Vegetatively similar to T. armeniaca. It is easy to
clubshaped, looking like a swollen part of the rhizome. identify by the very strong aromatic, somewhat vanilla
Leaves elliptical or nearly orbicular, two at each pseudo like scent that is emitted from the roots, particularly when
bulb. Never seen flowering in the Nimba area. F.W.T.A. the plant is pulled off it's substrate. D.J. 684 (UPS). Fig.
1 968, 3 : 1 describes the penduncle as stout .. very short, 47.
flower small,orange, ochra-coloured or reddish. Its creep T. crassifolia Summerh.
ing habit makes it hard to detect. It can be mistaken for a Lib. (new), Ghana.
Peperomia rotundifolia, since both grow in similar On the trunk or anywhere on the large branches. Full sun,
h abitats. D.J. 604 (UPS). bark. Rare. Erect, or sometimes hanging, with a stout
Ca/voa H ook. f. ( M elastomataceae) Table 7. Facultative epiphytes and filmy ferns in the Nimba
C. monticola A. C hev. ex Hutch. & Dalz. area.
Guin, S.L., Lib., Iv.C .
The lowest parts o f the trunks. Heavy shade, humus FAC ULTATI VE EPIPHYTES
deposits. Common. D .J. 785 (K, UPS).
Pteridophytes
C. trochainii Jac.-Fei. Asplenium variable Hook var. variable
Guin., Lib. Bolbitis fluviatilis (Hook.) C hing
Basal parts of trunks or middle parts of large branches. Bolbitis salicina (Hook.) Ching
Open shade, minor humus deposits. Common. D.J. 5 73 Lygodium smithianum Presl ex Kuhn
(UPS). Pityrogramma calomelanos (L.) Link
Selaginella blepharophylla Alston
Medinilla Gaud. ( M elasto m ataceae) Selaginella molleri Hieron.
M. mannii Hook. f. Selaginella myosurus (Sw.) Alston
Lib., F.Po. Selaginella versicolor Spring
Middle or basal parts of large branches. Open shade, Selagim:lla zechii Hieron.
humus deposits. At altitudes above 1 000 m, often solitary.
Other vascular epiphytes
At lower altitudes, often associated with Drynaria lauren
Araceae
tii. Common. D.J. 725 (UPS).
Cersestis afzelii Schott
Culcasia angolensis Welw. ex Schott
Peperomia R u izd & Pav. ( P i pera cea e)
Culcasia liberica N .E.Br.
P. fernandopoiana C . DC.
Raphidophora africana N.E.Br.
Guin., Lib., Iv.C., S.Nig., Cam., F.Po.
In all sections of the phorophyte in a wide range of Begoniaceae
habitats. Common. D.J. 475 (UPS). Begonia oxyloba Welw. ex Hook.
Begonia quadrialata Warb.
P. molleri C. DC.
Lib., Ghana, Togo, S.Nig., C am., F.Po. Melastomataceae
Basal parts of trunks. In heavy shade, minor humus Tristemma incompletum Benth.
deposits. Rare. D.J. 882 (K).
Moraceae
P. rotundifolia (L) H.B. &K.
Ficus anomani Hutch. (Fig. 48)
S.L., Lib., Iv.C., Ghana, Togo, C am.
Ficus kamerunensis Warb. ex Mildbr. & Burret.
Basal or middle parts of large branches. Open shade,
Ficus leprieurii Miq.
humus deposits. Rather common. D.J. 498 (UPS). Fig.
Ficus sagittifolia Warb. ex Mildbr. & Burret.
59.
Urticaceae
Preussiel/a G i lg. ( M elasto m ataceae) Urera oblongifolia Benth.
P. chevalieri Jac.-Fel. Urera rigida (Benth.) Keay
Guin ., Lib., Iv.C. FILMY FERNS
Basal parts of large branches. Open shade, humus Hymenophyllum kuhnii C.Chr.
deposits. Rather common. D .J. 7 8 5 (K).
Trichomanes africanum Christ
P. kamerunensis Gilg. Trichomanes chamaedrys Tanton
Lib., C am . Trichomanes erosum Willd.
Basal parts o f large branches. Open shade, humus Trichomanes guineense Afzel. ex Sw.
deposits. At altitudes above 1 000 m. Rare. D.J. 824 (K). Trichomanes mannii Hook.
N o. of
species
20
O rc h i d s
R a i nfa l l Yekepa
5 1 0 m 1 969
_ ... � � "'
_ ..... - -
JA N. N O V.
FE B . DEC.
16 B u l bophyl l u m spp.
Polystachya spp.
Fig. 59. Life forms of some epiphytes. (A) 1 . Rangae"ris Rhipsalis baccifera, (E) Peperomia rotundifolia, (F)
muscicola, 2. Nephrangis filiformis, 3. Tridactyle triden Polystachya a/finis.
tata, (B) A ngraecum podochiloides, (C) A . distichum, (D)
Table 1 1 . Water content in different parts of some observation that a leaf will temporarily continue its
Bulbophyllum species as a percentage of the total water content transpiration after it has been detached. This
in the plant. Mean of five samples. transpiration is noticed as a decrease in weight.
Loss in weight of plant material is often regarded
Root + Leaf Pseudo-
rhizome proper bulb
as a measure of water loss, since changes in water
content involve greater weight changes in the plant
Bulbophyllum cocoinum 7.9 29.3 62.8
Bulbophyllum congolanum 1 3 .9 34. 1 5 2.0
Table 1 2. Transpiration rates and some records of stomata
Bulbophyllum winkleri 3.5 3 1.9 64.6
numbers in the detached leaves of some epiphytes. The
transpiration rates are given as the loss of weight per hour in
percentage of the fresh weight. All figures refer to the mean of
water holding tissues. The proportions between the five samples.
water in the anatomical leaf (leaf proper + pseudo
bulb) and the rest of the plant were roughly 9 : 1 for Species No. of stomata per sq. mm Transpi-
three species examined (Table 1 1 ). Upper Lower ration
epidermis epidermis rates
The fleshy leaves of Calyptrochilum christyanum
hold approximately 60 % of the total water content Polystachya dalzielii 0 70 20.0
of this spec.ies (Table I 0). The Peperomias have Graphorchis lurida 0 72 1 1 .2
fleshy leaves, particularly Peperomia rotundifolia Polystachya puberula 0 58 1 1 .2
(Fig. 5 9). Due to a poorly developed root system and Asplenium geppii 1 0. 7
Polystachya leonensis 0 82 8.6
a thin trailing stem the plant appears to consist main
Begonia rubro-marginata 6.9
ly of the discoid leaves. Medinilla mannii 6.7
Preussiella chevalieri 6.3
Expenditure of water Arthropteris orientalis 5.6
transpiration rates, and the opinion was established Elaphoglossum isabelense 3.5
Phymatodes scolopendria 2.9
that the epiphytes, as most other ecological groups,
Plectrelminthus caudatus 0 1 04 2.9
provide examples of great variation in transpiration Polystachya rhodoptera 0 62 2.8
rates. Short period transpiration tests by Coutinho in Oleandra distenta 2.4
a virgin forest in Brazil, quoted from W alter Angraecum subulatum 2.3
1 929, Willis & Jefferies 1 963, Swanson & Lee 1 966, Nephrangis filiformis 0.3
Calyptrochilum christyanum 0.3
Rutter et al. 1 96 3). This method is based upon the
than do changes in the content of any other sub Table 1 3 . Transpiration rates in the detached leaves of some
stances present. The transpiration rate at the time of orchids. Given as the loss of weight per hour in percentage of
the fresh weight, and in absolute figures (g/dm2/hr).
cutting the leaf m ay be obtained by extrapolation
from the values in weight decrease during the first Species Transpiration (water loss)
three minutes. The effects of abscission and placing % of fresh g/dm2/hr
the leaf in a different environment for the weighing weight/hr
Graphorchis lurida 1 1.21 0. 1 4
make the accuracy of the result questionable
Bulbophyllum linderi 0.72 0.08
(Iwanoff 1 928, Rufelt 1 963, Willis et al. 1 96 3). But
Bulbophyllum winkleri 0.4 1 0.05
to the extent that these effects are similar for C alyptrochilum emarginatum 0.3 6 0.04
different species, the method is valuable as an index Diaphananthe pellucida 0.66 0.03
ing technique (Kochenderfer & Lee 1 97 3 : 1 77). Cyrtorchis arcuata 0.56 0.02
50
Table 1 4. Occurence of epiphytes on the trees ( 1 0 m or taller) in sample plot 1. The number mark the position of the tree in the pro-
file diagram (Fig. 66).
Piptadeniastrum africanum 1 38 2
Chlorophora regia 2 17
Chlorophora regia 3 17
Lophira alata 4 37 3 4 8
Heritiera utilis 5 23 6 1 7
Piptadeniastrum africanum 6 13
Lophira alata 7 39 4 4 2 10
Heritiera utilis 8 26 2 1 3
Mitragyna ciliata 9 29 5 3 2 10
Piptadeniastrum africanum 10 50 4 7 12
Parinari excelsa 11 11
Parinari excelsa 12 19 3 5
Nauclea diderrichii 13 38 6 4 11
Calpocalyx aubrevillei 14 18 1
Erythrophleum ivorense 15 28 3 3 7
Piptadeniastrum africanum 16 19
Lophira alata 17 44 2 3 6
Calpocalyx aubrevillei 18 16
Calpocalyx aubrevillei 19 25 2 2
Erythrophleum ivorense 20 15 2 4
Chidlowia sanguinea 21 13
Parinari excelsa 22 37 9 4 4 17
U apaca guineensis 23 20 2 2 2 6
Heritiera utilis 24 10
Chlorophora regia 25 26 1 2
Parinari excelsa 26 43 9 8 5 22
Chidlowia sanguinea 27 10
Calpocalyx aubrevillei 28 21 2 2 4
Parkia bicolor 29 29 2 2
Total 29 11 22 4 37
50
Table 1 5 . Occurrence of epiphytes on the trees ( 1 0 m or taller) in sample plot ll. The number mark the position of the tree in the
profile diagram (Fig. 6 7).
Total 35 14 19 4 37
Table 1 6. Occurrence of epiphytes on the trees ( 1 0 m or more) in sample plot lll. The number mark the position of the tree in the
profile diagram (Fig. 6 8).
Cryptosepalum tetraphyllum 1 33 2 3 5
Albizia zygia 2 11
Cryptosepalum tetraphyllum 3 32 2 3 5
Terminalia ivorensis 4 15
Triplochiton scleroxylon 5 23 1
Piptadeniastrum africanum 6 45 6 8 14
Anthocleista nobilis 7 16
Elaeis guineensis 8 23 5 5
Anthocleista nobilis 9 12
Elaeis guineensis 10 23 4 1 5
Lophira alata 11 42 3 7 3 13
Lophira alata 12 14
Terminalia ivorensis 13 19
Pentadesma butyracea 14 13
Lophira alata 15 12
Parinari excelsa 16 32 9 4 5 18
Calpocalyx aubrevillei 17 12 z 2
Terminalia ivorensis 18 42
Piptadeniastrum africanum 19 22
Lophira alata 20 16
Cryptosepalum tetraphyllum 21 36 3 3 7
Coula edulis 22 13 1 2
Heritiera utilis 23 18 2 2
Lophira alata 24 45 7 7 3 17
Canarium schweinfurthii 25 24
Piptadeniastrum africanum 26 38 2 3
Cryptosepalum tetraphyllum 27 15
Cryptosepalum tetraphyllum 28 13
Cryptosepalum tetraphyllum 29 14
Cryptosepalum tetraphyllum 30 19
Parinari excelsa 31 33 6 8 4 18
Parinari excelsa 32 12
Total 32 12 26 6 44
The plots were situated on level ground and on Table 1 7. Occurrence of epiphytes on phorophytes (1 0 m or
well-drained soils. It is doubtful if any of these taller) in high and secondary forest. (High forest figures from
Table 1 8. Aspects on the presence and abundance of epiphytes as reported from various parts of the world.
Country and Min. height of Total no. of % of trees with Max. no. of ep. Total no. of
author trees being trees being vase. epiphytes on any one tree ep. in the area
considered considered (est.)
Nigeria. (Richards, 1 93 9) 1 5 ft 67 15 13 35
Sarawak. (Richards, 1 9 3 2) 25 ft 91 13
Sarawak. (Richards, 1 93 2) 25 ft 44 11
Ecuador, Lowland.
(Grubb et al., 1 963) 20 ft 42 60
Ecuador, Mountain
(Grubb et al., 1 96 3) 20 ft 52 96
On cultivated trees
Trees, many of them of exotic origin, that are kept in
cultivation in and around the villages seem to be of
varying suitability as phorophytes. The common
mango tree, Mangifera indica, has a very poor or
non-existent epiphyte flora. Citrus trees (grapefruit,
orange and tangerine) seem to carry epiphytes only
occasionally, their bark, however, is covered with
white crustaceous lichens to such a degree that it
often gives a whitish impression.
Coffee trees may be observed with a substantial
number of epiphytes.
On 'herbs'
No epiphytes have been observed growing on herbs Table 24. Number of epiphyte 'stands' on a 1 4 m tall dead tree.
in the Nimba area. Guillaumet ( 1 967 :5 7) reports Yekepa 500 m.
ceptional specimen plant, a gigant covering some 20 Such a pattern has been observed by several
square metres and growing intermingled with Aloes, authors. Schimper ( 1 903) states that the epiphytic
Euphorbias and other xerophytic terrestrials, grows plants on a tree in a virgin forest are not the same
near the shores of Lake Elmenteita in the Rift V alley from its base to its topmost branches but exhibit a
in soil which is almost pure volcanic ash." well-marked differentiation. Moreau ( 1 943 :8): "A
Rangaeris muscicola which in dry areas of East few small species e.g. A ngraecum viride are found
Africa occurs on rocks is often connected to the always on twigs rather than branches." Holtum
presence of lichens as judged by notes on herbarium ( 1 960) observed that some small epiphytes grow
sheets : "On lic�ens on the huge granite boulders at only on the smaller branches of trees or shrubs.
the bottom of Njombe river cascade, Tanzania" Richards ( 1 964 : 1 1 8) : "Large epiphytic ferns such as
Lynes 1 93 1 (K), "Covering rocks in a mass. C ling Asplenium nidus and A. africanum tend to prefer the
ing to rock surface with lichens" Zambia, Abercorn trunks to the branches, but many epiphytes show the
Distr. H.M. Richards 8404 (K), "on lichen-covered opposite preference." Morris ( 1 9 70:5): "On all trees
rocks" Malawi, Mlanje, Morris 9 1 (K), "Epiphytic there is a pronounced stratification of the epiphytic
or lithophytic orchid, always in association with grey flora." But where an existence of some distribution
lichens" N. Rhodesia, Ika Hills, Morze 1 96 1 (K). patterns has been documented no actual investiga
One might speculate that the lichens with their tion has previously been done to describe or analyze
water-absorbing and water-holding capaCity could them.
create a suitable microclimate for the germination of
orchid seeds. The microclimate could also be of ma Ecological subdivision of the phorophyte
jor importance in the possibilities for the young Previous schemes
orchid plants to survive dry periods. To make it possible in a short time to record ac
The epiphytic orchids, with their slow growth and curately the epiphytes it is necessary to use some
small leaf surface, combined with a root system kind of subdivision of the tree.
adapted to superficial growth, appear suitable for Van Oye ( 1 924 b) divides the oil palm (Elaeis
this kind of habitat. guineensis) into five zones. This zonation is useful
for the oil palms and other palms whose crowns are
built up by large leaves. For the often extensive
Distribution of epiphytes on the phoro crowns of the large rain forest trees this zonation is
phytes less useful. In Europe Ochsner ( 1 928) uses the same
The first impression one gets, when looking at the zonation as Van Oye, excluding the submedian zone.
distribution of epiphytic plants within a tropical rain Richards ( 1 9 3 9) in certain cases used the height
forest, is the lack of order. It seems that the above the ground, in others a more relative descrip
epiphytes are able to grow at any place on the trees. tion, e.g. base of the first branch, in the records of
There are many reasons that help to create this im the growing site of the epiphyte. Sometimes these
pression. One of these is the limited sector of the two pieces of information are combined. The use of a
trees that can be obser.ved from the ground, another large number of relative terms, without strict defini
that many observations are done in areas where tion, is rather confusing. In 28 felled trees, recorded
human activities have disturbed the original as bearing epiphytes, Richards used no less than
stratification, i.e. bright light may penetrate to the eleven such relative 'localities'.
lowest level of the forest. In such cases one often Hosokawa ( 1 954 a) divides the phorophyte into
finds species normally belonging to the topmost four sections. An arbitrary system of 1 5 ft (4.5 m)
layer of the forest growing close to the ground. zones from the ground upwards has been used by
Many other influences may contribute towards an Grubb et al. ( 1 96 3 :592-59 3).
impression of randomness in the distribution of the Tixier ( 1 966) works with three large sections, but
epiphytes. A regular pattern emerges first when one uses 5 subunits within the 'base of the trunk section'.
has the possibility of examining a substantial number The study of epiphytic lichens and bryophytes in
of trees in an undisturbed environment from the out temperate climates has caused many subdivisions of
ermost branches in the crown down to the base of the phorophytes. Barkman ( 1 95 8 :3 1 -3 2) has made a
the stem. valuable compilation of that terminology. The lower
parts of the phorophytes have generally been subject epiphytes appear. This stratification of the rain forest
to many subdivisions, while as a rule the whole is by no means uniform, but rather large variations
crown has been regarded as one section. This can in can be found (Foggie 1 947, Newman 1 95 4, Robbins
many cases be traced back to poor knowledge of the 1 95 9, Holdridge 1 970). As a rule the crowns are
epiphytes of the crown rather than being based upon rather small compared to the length and size of the
real observations. A number of subdivisions used in trunks, but a few species of trees with the ramifica
studies of vascular epiphytes is presented in Table tion close to the ground give rise to crowns of large
25. dimensions.
Usually the emergent (superstory ; Odum 1 970 a)
Biotopes on the phorophytes in the Nimba area trees have more flattened and expanding crowns
In the tropical rain forest there is a pronounced than the trees in the lower ' strata' of the forest.
stratification of the vegetation. The crowns of the The richness in biotopes which can be found in the
trees form one or more 'strata' and it is in the top crowns of the large trees is in sharp contrast to the
most of these 'strata' that the main part of the more uniform environment in the lower 'strata' of the
Fig. 77. The concealment of the epiphytes by the foliage observed. Note the distribution of the large Platycerium
as illustrated by a Triplochiton scleroxylon. This tree is ferns.
deciduous for a short period when the epiphytes are easily
might seem to offer an immense material. Unfor the monopodia] orchids grow out from the branches,
tunately these farming activities are to the largest ex and thus a very minute part of the plant, aside from
tent carried out in so-called secondary forests. Here the extensive root system, covers the sampling area.
the vegetation is after repeated destruction kept in The scale constructed is therefore based on the ac
various young secondary growth stages. tual number of plants rather than the area covered.
When the tree is lying on the ground it is relatively The transects can preferably be drawn along the
simple, at least when one knows well enough the branches in the crown, since it is here that the most
vegetative parts of the epiphytes to make deter substantial changes occur in substrate, light and
minations, to map the· distribution and to study the microclimate. From the point of ramification, where
ecology of the various species. Certain large the branch is relatively broad, the transect as it runs
branches can be used to illustrate the continuous aiong the branch, covers a smaller and smaller area.
changes in the epiphytic flora from the base to the This fact should be considered when reading the
outermost parts, so-called 'transects'. transect results (Fig. 79).
In the description of vegetative change along an
environmental gradient the 'transects' are of a con Distance observation
siderable importance. But since neither the trunks Such investigations have been performed on a stan
nor the branches represent straight lines a transect of ding phorophyte on which it has been possible to in
traditional type is not easy to work with. Therefore a vestigate the trunk and at least one complete branch
modified type of transect has been used. The method (or more seldom the whole phorophyte). This
is simply to let a measuring tape follow the twistings method, described by Went ( 1 940), allows the recor
and bends of the stretch to be examined. This stretch ding of the epiphytic flora from the ground. With the
is later presented as if it was a straight line. Most of aid of field glasses, or rather a telescope on a tripod,
species either by an area on the tree devoid of strate at its growing site were recorded in a three
orchids or occupied by another species. In the case graded scale.
where the same area is occupied by an intermingling
of more than one species, one stand is counted for L ight
each species present." Light intensity was divided into three classes : heavy
During the investigation the same unit (called shade, open shade and full sun. The amount of light
'group'), as described by Sanford, was used. In order available varies of course with the time of the day
not to cause confusion by adding one more name for and cloudiness of the sky, and also somewhat with
the same thing, the term 'stand' sensu Sanford is the time of the year. Since the light intensity was
adopted. only estimated, mistakes cannot be excluded. How
ever, the main bulk of the observations are easy to
Ecological observations group.
In order to get a rough idea of the importance of cer Light intensities regarded as heavy shade normal
tain environmental influences on the distribution of ly appear in the lower strata of the rain forest. In ex
the epiphytes within the forest, attention has been ceptional cases this intensity can be found higher up
paid to light and substrate. For each epiphyte that in the trees due to dense foliage. In most cases there
was recorded, the light intensity and type of sub- is no problem to separate heavy shade from open
Table i6 _ The trees ( 1 0 m or taller) o f the high forest included in the s tu d y . Trees with epiphytes are marked
C lose observation =
D= ta
iso..=
=-on=
ce ob
"---
"--'
-"' s e"---
'--"
rv:. a
:-=:
.:: Lic:
on
.: _
:..:._ ___________
No . % with No . % wi th
Phorophytes epiphytes Sizes epiphytes Sizes
Total 222 24 1
shade intensities. The light in a tropical rain forest is ( 1 940) used a relative scale to judge the light intensi
not gradually increasing from the ground level up to ty at the growing sites of the epiphytes. He estimated
the crowns of the emergent trees. It is rather discon the light intensity at the growing site in per cent of
tinuous, as many author have pointed out, e.g. the light outside the forest.
Richards ( 1 939 : 30). In the crowns of the trees of the
topmost stratum high light intensities are formed by Substrate
open shade. The limit between full sun and open The substrate was divided into three classes : bark,
shade is also easy to observe, since only plants in e.g. bark without humus attached, minor humus
very exposed posttlons receive light intensities deposits, e.g. bark with thin and scattered humus
regarded as full sun. layer, and large humus deposits. Humus is here
In a study of the epiphytic societies in Java, Went regarded as dead organic material in various stages
Table 27a . Total number of records, according to .the close observation method, o f pteridophytes and other vascular epiphytes in the high fore s t .
II Ill IV V A B c A B c A B c
Pteridophytes
T o ta l n o . of observations 503
The dis tribution o f the records between ( 1 ) the di fferent sections of the phorophytes and ( 2) the different light and substrate classes each refer to
a percentage of the total number of records. A = Bark, B ; Minor humus depos i ts , C ; Large humus deposits.
II Ill IV V A B c A B A n
Orchids
of decay. This subdivision of substrates was easy to representing 4 7 species of 2 1 genera. 222 trees were
work with during the recording. investigated according to the close observation
Each epiphyte or group of epiphytes has thus been method and 24 1 by distance observation (Table 26).
recorded in one of the nine possible light-substrate The quantitative figures from this investigation for
classes. The idea was to provide a base for a quan each species of epiphyte is presented in Tables 27, 28
titative judgment of each species' dependence of light and 29.
and substrate. However, it should be emphasized It became evident that most species of epiphytes
that many other influences effect the presence and are ± restricted to one or two particular section(s) of
distribution of epiphytes. the phorophyte, while a few occur more evenly in a
number of sections (Fig. 78). An analysis of the
General distribution pattern results from this investigation will later be presented,
The distribution pattern was recorded on 463 in correlated to certain environmental influences
dividual trees C> 10 m in height) of the high forest, (Chapter VI).
Table 2 8 a. Total number o f records, according t.o the distance observation m e thod, o f pteridophytes and other vascular epiphytes i n the
II III IV V A B c A B c · A B c
Pteridophytes
Total n o . o f observations 3 3 54
I! lil IV V A B A B A B c
Orchids
B . scariosum
B . schimperanum
B . schinzianum 19 73 . 7 26. 3 15. 8 5.3 63. 2 1 5. �
B . winkleri 15 20 . 0 40 , 0 40. 0 60. 0 6.7 33 . 3
C a lyptrochi \urn christyanum 70 11.4 43 , 0 41 , 4 4.3 23. 0 H.3 . �.G 37 . 1 17.1
C . em a rginatum �5.0 75. 0 1 00 . 0
Chamaeangis vcsica t...1. 45 6. 6 4.4 80,0 8. �) 17 . 8 11.1 60 . 0 11.1
Cyrtorchis arcuata 186 1.1 52 . 7 45. 2 1.1 24 . 7 32 . 3 1 5. 6 IG. 1 10. 8 1.6
C . aschersonii 17 11. 8 23 . 6 47 . 1 17. 6 23 . G 5 , !! 5 8 . :S l l . ti
C. monleiroac
D iapha nanlhe bidens 24 8.3 37 . 5 45. 8 8.3 8.3 29. 2 1� . 5 :. n . :J 1 � . :)
D . densiflora
D. pellucida 27 14. 8 55 . 6 2 !J . G 7.4 66 . 7 7. 4 3.7 11. I 3.7
D . rutila
Eurychone rothschildiana
Genyorchis pumila
Graphorchis lurida 2 57 0.8 17. 9 60.7 20 . 6 16.0 3. 5 71.2 8. D 0.4
Habenaria leonensis 2 50 . 0 50 . 0 100. 0
H . procera
Liparis caillei
L. nervosa
Llstrostachys pertusa 455 5.3 24.6 52 . 7 17.4 21. 1 14. 9 0.7 45. 1 17. 6 0.7
Nephrangis filiformis 306 7.5 69 . 3 23 , 2 18. 0 2.0 74 . 2 5. 9
Plectrelminthus c audatus 20 75. 0 25. 0 5.0 5.0 80 . 0 10. 0
Podangis dactyloc eras
Polystachya adansoniae 229 17.0 50 . 7 32 . 3 10.5 23 . 6 1.7 23 . 1 39.3 1.7
P . affinis 1 100 . 0 100 . 0
P. dalzielii
P . elastica
P. galeata 87 58 . 6 37 , 9 3.4 3.4 48. 3 20 . 7 1.1 18. 4 8. 0
P. lru<iflora 10 80 . 0 20 . 0 50. 0 50 . 0
P . leonensis
P. microbambusa
P . obanensis 50. 0 50 . 0 1 00 . 0
P. paniculata 259 37 . 1 50 ' 6 12. 4 32 . 8 13. 9 0.4 37 . 8 14.7 0.4
P. pobeguinii
P. polychaete 1 85 31. 9 47 . 0 21 . 1 9.2 18.4 22. 2 10.3 28. 1 11.9
P. puberula 240 22. 5 54 . 2 23 . 3 13.7 33 . 3 5.0 22.1 23 . 8 2.1
P . ramulosa
P . rhodoptera
P. saccata 13 15.4 76. 9 7.7 69. 2 30 . 8
Polystachya subulata
P . tenuissima 33, 3 66 . 7 1 00 . 0
P. tessallata 244 0. 8 38. 1 51 . 2 9. 8 12.7 29. 9 23 . 8 9. 0 13. 5 11.1
Ranga�ris brachyceras
R . musctcola 1 80 25. 0 63 . 3 11. 7 11.7 8. 9 13 . 3 49.4 16.7
R . rhipsalisocia 19 63 . 2 36. 8 94 . 7 5.3
Rhipidoglossum paucifolium
Sto lz ia repens
Tridactyle anthomaniaca 766 9. 0 50 . 0 41. 0 14 . 6 G . !l 1.3 64 . ;, 11.1 o.n
T. armeniaca 352 2.8 53 . 4 41. 8 2.0 20. 7 4 3 .8 17.G 7.7 9.9 0. �
T. bicaudata
T. crassifolia 12. 5 75.0 12.5 12. 5 62 . 5 25.0
T. tridactylites 50 8.0 52, 0 30. 0 10, 0 10. 0 40. 0 4.0 16. 0 24. 0 6.0
T. tridentata 43 7. 0 7 0. 0 23 . 0 11.6 2.3 79.1 7.0
Vanilla crenulata
The distribution on the phorophytes of the various previously described. A rather clear distribution
epiphyte species showed somewhat different results pattern for most species of epiphytes was obtained
depending on observation method. Compared to the (Fig. 79), no matter if there were few or many
results from the close observation method many species present. The distribution pattern also seemed
species showed a ± marked displacement towards to be similar between various species of phorophytes.
the outer parts of the crown in the distance observa Thus, the transects along the large branches seem
tion. This could be anticipated since the epiphytes in to represent a rather fast and informative way to get
the distance _ observation are more easily noticed a good insight into the distribution pattern for the
when growing on the outer parts of the branches. various species. This method may preferably be used
Large branches from several phorophytes were when there is a limited number of phorophytes to be
examined according to the transect technique examined.
Table 2 9 ta l number of
. To records, according to the occasional observation m e thod, o f some epiphytes i n e
th h igh fore s t . (Explanation in Table 2 7 a . )
I! III IV V A B c A B c A B c
t
P e ridophyt es
1
A rthropteris monocnrpn 46 21 . 7 54 . 3 15. 2 �.7 26. 1 -H . 8 21 . 7 4.3
00 . 0
en u c t
A . pa li so ti 1 00 . 0
A spl i m a h i opic u m 23 4.3 39. 1 52 . 2 4.3 8.7 43 . 5 26. 1 8. 7 13.0
u
A . hemitomum 10 20. 0 80. 0 60. 0 20 . 0 20. 0
A. variable v . pa u c ij gum 54 59 . 3 40. 7 66. 7 25. 9 7.4
Elaphoglossum kuh n i i 26 88. 5 11.5 11 . 5 76. 9 11. 5
Orchids
c
Calyptrochi turn emarginntum 40 30. 0 57 . 5 7.5 5. 0 15. 0 2. 5 77 . 5 5. 0
Cyrtorchis mo n t i rone 14. 3 71.4 14.3 28. 6 71.4
D iaphananthc densinora 43 48. 8 46. 5 4.7 83 . 7 7. 0 7. 0 2.3
D . rutila 120 61 . 7 33 . 3 5.0 23 . 3 0. 8 73. 3 2. 5
E u rychone rothschi ldiana 27 66. 7 33 . 3 59 . 3 40. 7
Genyorchis pum i l a 16 25. 0 25. 0 43. 7 6.2 12. 5 18.7 25. 0 37 . 5 6.2
Habcnaria leonensis 13 15. 4 84 . 6 7. 7 61 . 5 23 . 1 7.7
H . procera 35 11.4 68. 6 17 . 1 2 . !) 22. 9 54 . 3 5. 7 17 . 1
Lipa ris cni l lc i 26 76. 9 11. 5 11.5 7. 7 69. 2 23 . 1
L . nervosa 29 62 . 1 31 . 0 3.4 3.4 6. 9 48. 3 24 . 1 6. 9 13 . 8
Podangis dnctyloccras 12 83 . 3 16.7 33 . 3 8. 3 41 . 7 16. 7
.0
Polys tachyn nffinis 76 27 . 6 63 . 1 6. 6 2. 6 51 . 3 13 . 2 30.3 5. 3
P . e l astica 1 00 . 0 1 00
P . micmbnmbusa 21 38. 1 57 . 1 4 . 8 14, 3 19.0 66 . 7
P . pobeguinii 33 3. 0 24 . 2 42 . 4 15. 2 15. 2 9. 1 21 . 2 18.2 48. 5 3 . 0
P. ramulosa 16 62. 5 37 . 5 12. 5 18.7 50 . 0 18.7
55
P . rhodoptcra 31 74 . 2 25. 8 3.2 22 . 6 35. 5 3.2 9.7 25. 8
P. s ubu l a ta 22. 2 .6 22 . 2 22 . 2 33. 3 44 . 4
P. Lt=1iui s sim a 14 35. 7 57 . 1 7.1 28. 6 21 . 4 50 . 0
Rangallris brachycerns 18 5.6 22.2 66.7 5. 6 16.7 5.6 50. 0 27 . 8
"
Rhipidoglo s s u m paucifolium 27 51 . !J 37 . 0 11.1 37. 0 11. 1 37 . 0 14. 8
Stolzia repens 15 6.7 13. 3 60 . 0 20 . 0 80. 0 20. 0
Tridactyle bicaudaW 57 26 . 3 21. 1 47 . 4 5.3 22 . 8 54 . 4 3. 5 3.5 15.8
Va n i l la c rcnu l a tn 83 . 3 16.7 33 . 3 66 . 7
Terminology
The frequently observed aggregation of epiphytes
('arboreal gardens' according to Oliver 1 930 :2) in
limited areas on the phorophyte is of interest not
only because of the information they provide as to
Polystachya B u l bophyl l u m B o l u s i e l l a Polystachya common environmental requirements for various
ra m u losa b u fo t a l bot i i leonensis
species but also as a source in the analyses of the
colonization and development of the epiphytic flora
in general. The terminology that has been used in the
classification of epiphyte communities is confusing.
The terms 'epiphyte societies' (Oliver 1 93 0 : 1 6)
and ' association' � Went 1 940 :86) have been used.
C a lvoa B eg o n i a R h i psa l i s Pepero m i a Epiphytic association and epiphytic sociation were
monticola rubro- m a rg i nata ba ccifera· sp.
proposed by Omura ( 1 95 3). Hosokawa ( 1 95 3 ,
vl
IV
1 954 a , b , c , 1 968) has created a new terminology
Ill
for the community units of epiphytes. From the
11 viewpoint of the ecosystem concept he does not
I . agree with the use of phytocoenosial units for
____,
Ca lvoa troc h a i n i i
M ed i n i l l a m a n n i i
Pepero m i a sp.
O l e a ndra d istenta -
B u l bo p h yl l u m coc o i n u m -
B u l bophyl l u m i n tertext u m - -
A n g ra ecopsis e l l iptica
P h y m a todes scolopendria -
Ang ra e c u m disti c h u m
Po lystachya tessa l l ata
Aspl e n i u m meg a l u ra
D ry n a ri a l a u re n t i i
P l a t y<:e r i u m a n golense -
Asplen i u m m eg a l u ra
Tridactyle a rm e n i a c a -
1/i tt a r i a g u i ne e n s i s
Polyst a chya g a leata
A n g r a e c u m d i st ic h u m
Cyrtor c h i s a rc u a t a -
Polystachya tessa l l at a
Polystac hya p u b e r u l a -
R h i ps a l i s baccifera
R h i p s a l i s baccifera - =
Dryn aria l a u re n t i i
G ra ph orc h i s l u ri d a
G ra phorc h i s l u rida
B ol u si e l l a ta l bo t i i
O l e a nd ra diste nta - -
16 10 15
������ m �------�--����- m
Fig. 79. Number of epiphytes and their distribution along tadeniastrun africanum, illustrated according to the
a branch of: Left : Parinaria excelsa, illustrated according 'transect' technique. Yekepa 500 m.
to the 'transect' technique. Seka Valley 600 m. Right : Pip-
To avoid adding to the already confusing ter when a group of three or more species form a unit,
minology of epiphyte communities, they will here and when the distance from two of them does not ex
simply be called epiphyte communities (e.c.). To ceed 0.5 m to the third. The root systems and the
separate them from each other they are named by stems are often heavily intermixed, to such a degree
the dominant species ('cover') and the most frequent that the plants often grow on top of each other.
ly occurring species in the community. This unit Epiphytes that colonize the vegetative parts of
�hould be homologous to the term 'epilia' proposed other epiphytes are called hyper-epiphytes by Tixier
by Hosokawa. ( 1 966) and 'secondary' epiphytes by Hosokawa. The
delimitation of an epiphyte community in most cases
Methods is easy to do, since the grouping of plants is rather
The technique used in the investigation and descrip distinct (Fig. 79). There are some spectacular excep
tion of terrestrial plant communities is impossible to tions from this rule. Drynaria laurentii may often
use under the circumstances of growth and oc cover stretches of more than five meters length. This
currence which the epiphytic plants exhibit. The extended distribution will of course connect a larger
knowledge of the epiphytic plant communities is number of plants to this particular epiphyte.
based on records from close and distance ex However, most epiphyte communities occur in a
aminations. The interest has been concentrated on limited area with no other plants nearby. This is the
the presence of the various species rather than on case particularly for certain communities formed by
their number. epiphytes that are attracted to humus deposits.
An epiphyte community is considered to exist Seven different epiphyte communities are recognized
Table 30. The number, distribution on the phorophytes and species composition of the epiphyte communities examined. (The distri-
bution is given as a percentage of the total number of communities and the species composition according to close and distance ob-
servations.)
Angraecum birrimense-
-
Raphidophora· africana 65 43. 1 5 6.9 13 10 19 15 6 5 38 30 39
Asplenium africanum-
Peperomia sp. 13 3 8. 5 1 5 .3 46.2 3 3 0 3 3 6 7 9
Oleandra distenta-
Tridactyle armeniaca 1 04 6.8 70.2 23. 1 13 17 19 20 12 10 44 47 51
Microsorium punctatum-
Vittaria guineensis 63 6.8 70.2 23. J 6 5 13 11 7 5 26 21 29
Drynaria laurentii-
Asplenium megalura 1 90 5 . 3 34.7 48.9 1 1 . 1 13 8 37 25 8 8 58 41 62
Platycerium stemaria-
Nephrolepis undulata 54 7.4 5 1 .8 40.7 7 5 9 7 4 3 20 15 22
Platycerium angolense-
Nephrolepis undulata 50 30.0 1 2.0 58.0 6 4 7 5 3 3 16 12 17
Asplenium dregeanum-
-
Peperomia sp. 42 88. 1 1 1 .9 9 5 5 19 19
Tridactyle tridactylites-
-
Medinilla mannii 60 3 . 3 7 6 . 7 20.0 11 8 8 7 5 3 24 18 26
Bulbophyllum scariosum-
Polystachya dalzielii 25 1 2.0 64.0 24.0 11 8 6 5 3 4 20 17 21
below 1 000 m alt. and three more above that used (Table 3 1 ). There are, however, some excep
altitude. tions. The 0/eandra distenta - Tridactyle armeniaca
The investigation includes a total of 666 'com and A ngraecum biDrimense - Raphidophora africana
munity aggregations'. Their distribution on the communities, built up by more than five species
phorophyte and the total number of species recorded each, were more common than those with a lesser
in the various communities are given in Table 30. number of species. The possible explanation to this
The ten most frequently occurring species in each could be that a more or less simultaneous coloniza
community are presented in Table 3 2 . tion by several species is taking place when the com
munity forming species has created a suitable en
Abundance vironment. Both these communities are found in the
Like epiphytes. in general the epiphyte communities lower parts of the phorophyte, and could possibly
are more abundant in older trees. A total of 25 'com catch seeds or seedlings washed down by heavy rain
munity aggregations' were found on the 1 1 8 trees fall.
that were examined in the three sample plots (Tables
1 4, 1 5 and 1 6). In relation to the abundance of the Epiphyte communities at altitudes below 1000 m
dominant species most communities were scarce. Angraecum birrimense-Raphidophora africana e.c.
0/eandra distenta, however, was found just as fre This community is most common around the first
quent in a community formation as in a single state. ramification on trees which have a rich supply of
As would be expected the most common number of light in that section. This means that the community
species in the individual communities is three, which will appear mostly in secondary forests or on trees
are the lowest possible according to the definition bordering rivers or creeks, or along tracks or roads.
Table 3 1 . The number of species in the different epiphyte communities, expressed as a percentage of the total number of records.
It gives an impression of an irregular jumble of her development of this community certain orchids may
baceous climbers, lianas and trailing epiphytes. grow between the rhizomes, but later as debris is ac
The community is delicately balanced. An in cumulated they become fewer and fewer.
crease in light and evaporation will change this com This community is not subject to the extreme
munity leaving only the Diaphanthe orchids, a variation in microclimate that exists further out on
decreasing light supply will lead to an overgrowth by the branches, and the light intensity is naturally
the vigorous Raphidophora africana. lower. These conditions are also reflected in the com
position of the species in the community. A high
number of pteridophytes and other vascular
Asplenium africanum - Peperomia sp. e.c.
epiphytes (non-orchids) are typical. Note that no
Asplenium africanum that is rather uncommon,
orchid species except Tridactyle armeniaca is found
forms an accumulation of humus around its attach
among the 10 most common species (Table 3 2). This
ment to the substrate in a similar way as
community is composed of a mixture of humus
Microsorium punctatum. Only 1 3 communities have
preferring species and species tolerating different
been found and examined, 8 with telescope and 5 on
kinds of substrate. Orchids with preference for rich
felled trees. With so few observations it is of little use
humus and open shade found in this community are,
to make an analysis of the occurrence of the various
e.g. Brachycorythis kalbreyeri and Habenaria
species. Worth mentioning is the presence of
procera. The majority of orchid species that oc
Peperomia sp. in 69.2 % of the communities ex
casionally occur here normally belong to the central
amined.
or outer parts of the branches. Also the
pteridophytes in this community are a mixture of
Oleandra distenta - Tridactyle armeniaca e.c. species of humid habitats such as filmy ferns and
The fern Oleandra distenta, that forms the base of Nephrolepis undulata, as well as Drynaria laurentii
this community, is common and easily recognized. It and Microsorium punctatum of more sunny and
often starts to grow in or around the ramifications of drier sections of the crown. Begonia polygonoides,
the larger branches in the central part of the crown. Remusatia vivipara, Peperomia sp. as well as more
It expands in all directions but is mainly found at the drought resistent species such as Rhipsalis baccifera
basal parts of the branches that are often completely and Begonia rubro-marginata are also frequent in
overgrown by thick cushions of this fern and species this community.
associated with it. The slender rhizomes often hang
down giving this community a characteristic Microsorium punctatum - Vittaria guineensis e.c.
appearance (Fig. 80). In the earlier stages of the This community has a very limited distribution and
Table 32. The ten most common species in the different epiphy
Close observation Distance observation
te communities. The occurrence is given as a percentage of the
total number of communities examined (Table 30). Plathycerium stemaria - Nephrolepis undulata e.c.
Nephrolepis undulata 58.3 Nephrolepis undu1ata 66.7
Close observation Distance observation Polystachya tessallata 38.9 Begonia rubro-marginata 50.0
Peperomia sp. 27.8 Polystachya tessallata 38.8
A ngraecum birrimense - Raphidophora africana e.c.
Begonia rubro-marginata 27.8 Davallia chaerophylloides 38.8
Raphidophora africana 79.3 Raphidophora africana 83.3
Vittaria guineensis 22.2 Diaphananthe bidens 33.3
Vittaria guineensis 48.2 Elaphoglossum barteri 36. 1
Lycopodium warneckei 1 9.4 Rhipsalis baccifera 33.3
Elaphoglossum barteri 34.5 Vittaria guineensis 33.3
Rhipsalis baccifera 1 6. 7 Vittaria guineensis 27.8
Calvoa trochainii 34.5 Diaphananthe bidens 27.8
Polystachya galeata 1 3 .9 Tridactyle anthomaniaca 22.2
Diaphananthe bidens 24. 1 Ancistrorhynchus capitatus 2 5 .0
Asplenium megalura 1 1.1 Peperomida sp. 1 6. 7
Microgramma owariensis 20.7 C alvoa trochainii 25.0
Polystachya polychaete 1 1. 1 Asplenium megalura 1 1. 1
Nephrolepis undulata 20. 7 Peperomia sp. 1 9. 4
Habenaria procera 1 3.8 Microgramma owariensis: 1 3. 9 Platycerium angolense - Nephrolepis undulata e.c.
Ancistrorhynchus capitatus 6.9 Nephrolepis undulata 1 1. 1 Nephrolepis undulata 47.5 Nephrolepis undulata 60.0
Rhipidoglossum paucifolium 6.9 Diaphananthe rutila 5.6 Polystachya tessallata 30.0 Polystachya tessallata 40.0
Peperomia sp. 25.0 Davallia chaerophylloides 30.0
A splenium africanum - Peperomia sp. e.c.
Davallia chaerophylloides 1 7 .5 Begonia rubro-marginata 30.0
Peperomia sp. 80.0 Peperomia sp. 62.5
Begonia rubro-marginata 1 5 .0 Diaphananthe bidens 30.0
Asplenium megalura 40.0 Preussiella sp. 37.5
Vittaria guineensis 1 5 .0 Rhipsalis baccifera 30.0
Calvoa trochainii 40.0 Begonia mannii 25.0
Polystachya galeata 1 2. 5 Tridactyle anthomaniaca 30.0
Elaphoglossum barteri 20.0 Arthropteris orientalis 25.0
Lycopodium warneckei 1 2. 5 Vittaria guineensis 20.0
N ephrolepis undulata 20.0 Nephrolepis undulata 1 2. 5
Nephrolepis biserrata 7.5 Chamaeangis vesicata 1 0.0
Begonia mannii 20.0 Asplenium megalura 1 2. 5
Diaphananthe bidens 7.5 Lycopodium warneckei 1 0.0
Polystachya tessallata 1 2. 5
A splenium dregeanum - Peperomia sp. e.c.
Oleandra distenta - Tridactyle armeniaca e.c.
Peperomia sp. 69.0
Tridactyle armeniaca 53.8 Tridactyle armeniaca 4 1 .7
Polystachya leonensis 28.6
Peperomia s p . 42.3 Drynaria laurentii 30. 6
Elaphoglossum chevalieri 2 1 . 4
Nephrolepis undulata 34.6 Begonia polygonoides 27.8
Asplenium barteri 1 9.0
Phymatodes scolopendria 30.8 Rhipsalis baccifera 27.8
Medinilla mannii 1 6. 7
Begonia polygonoides 26.9 Phymatodes scolopendria 2 5 .0
Arthropteris monocarpa 1 4. 3
Calvoa trochainii 23. 1 Nephrolepis undulata 1 9.4
Polystachya laxiflora 1 4. 3
Rhipsalis baccifera 23. 1 N ephrolepis biserrata 1 6. 7
Asplenium aethiopicum 1 1.9
Preussiella chevalieri 1 9. 2 Elaphoglossum isabelense 1 6. 7
Begonia rubro-marginata 9 . 5
Asplenium megalura 1 5 .4 Peperomia sp. 1 3 .9
Bulbophyllum inflatum 7. 1
Microsorium punctatum 1 1 . 5 Asplenium megalura 1 2. 5
Tridactyle tridactylites - Medinilla mannii e.c.
Microsorium punctatum - Vittaria guineensis e.c.
Medinilla mannii 58.8 Medinilla mannii 53.5
Vittaria guineensis 5 1 .8 Vittaria guineensis 36. 1
Asplenium dregeanum 4 1 .2 Polystachya laxiflora 27.9
Peperomia sp. 44.4 Davallia chaerophylloides 3 3 . 3
Polystachya leonensis 35.3 Lycopodium mildbraedii 2 5 . 6
Davallia chaerophylloides 3 7.0 Peperomia sp. 30.6
Vittaria guineensis 35.3 Peperomia s p . 23.3
Polystachya tessallata 29.6 Polystachya tessallata 27.8
Asplenium barteri 29.4 Polystachya leonensis 23.3
Diaphananthe bidens 25.9 Asplenium megalura 25.0
Polystachya laxiflora 23.5 Asplenium dregeanum 20.9
Begonia rubro-marginata 2:2. 2 Medinilla mannii 25.0
Polystachya obanensis 23.5 Vittaria guineensis 18.6
Nephrolepis undulata 1 8. 5 Diaphananthe bidens 1 9.4
Rangaeris muscicola 23.5 Cyrtorchis arcuata 1 3.9
Oleandra distenta 1 8. 5 Graphorchis lurida 1 3 .9
Peperomia sp. 23.5 Bulbophyllum scariosum 1 1 .6
Medinilla mannii 14.8 Angraecum birrimense 1 3 .9
Elaphoglossum barteri 1 7. 6 Elaphoglossum salicifolium 1 1 . 6
Bulbophyllum cocoinum 1 4 . 8 Nephrolepis biserrata 1 1. 1
Bulbophyllum scariosum - Polystachya dalzielii e.c.
Drynaria laurentii- A splenium megalura e.c.
Xiphopteris oosora 42.9 Polystachya dalzielii 44.4
Asplenium megalura 53.2 Asplenium megalura 43.7
Polystachya dalzielii 42. 9 Lycopodium mildbraedii 27.8
Tridactyle armeniaca 46.8 Rhipsalis baccifera 29.7
Elaphoglossum salicifolium 28.6 Asplenium megalura 22.2
Rhipsalis baccifera 45. 1 Tridactyle armeniaca 2 5 .0
Bulbophyllum bifarium 28.6 Vittaria guineensis 22.2
Polystachya tessallata 32.3 Graphorchis lurida 25.0
Xiphopteris vilosissima 28.6 Tridactyle tridactylites 22.2
Bulbophyllum linderi 25.8 Bulbophyllum oreonastes 1 8 . 7
Arthropteris monocarpa 1 4. 3 Elaphoglossum salicif. 1 6.7
Ficus kamerunensis 22. 6 Angraecum distichum 1 7.2
Asplenium megalura 14.3 Asplenium geppii 1 1.1
Graphorchis lurida 22.6 Cyrtorchis arcuata 1 4. 1
Lycopodium mildbraedii 1 4. 3 Arthropteris sp. 1 1.1
Polystachya galeata 16.1 Polystachya tessallata 1 4. 1
Polystachya pobeguinii 1 4. 3 Podangis dactyloceras 1 1.1
Medinilla mannii 1 1 .3 Phymatodes scolopendria 1 2. 5
Vittaria guineensis 1 4. 3 Asplenium aethiopicum 1 1. 1
Nephrolepis undulata 9.6 Bulbophyllum linderi 1 1 .7
covered several ferns and one orchid, Cirrhopetalum clearly can be seen through the lower parts of the
umbellatum (Forst. f.) Hook. & Arn., associated newly developed sterile leaves. The fertile leaves are
with Drynaria volkensii. Moreau probably refers to long and pendulous.
the first stages when the Drynaria fern seldom has The rhizome is short. It is not creeping as in most
any other epiphytes associated with it. other epiphytic ferns but is always growing at a 90
degree angle towards its point of attachment.
Platycerium stemaria - Nephrolepis undulata e.c. The occurrence of the Platycerium communities in
and Platycerium angolense - Nephrolepis undulata relation to the total presence of these ferns, is low.
e.c. The low number of epiphytes associated with the
Two species of the genus Platycerium are common Platycerium ferns is also surprising. Platycerium
in the Nimba area, Platycerium angolense and P. angolense had 1 7 species and P. stemaria 22 species
stemaria can often be found together in non-limiting totally. A look at the species compositions shows
environments, but in open or more exposed habitats that the plants associated with the Platycerium ferns
Platycerium angolense dominates. In other parts of commonly appear in other epiphytic communities as
Africa they also grow on rocks, e.g. in the Matombo well. Nephrolepis undulata is the species that most
region east of the Uluguru Mts in Tanzania. Both frequent takes advantage of the substrate or perhaps
species are widespread in tropical Africa. They in a favourable microclimate caused by the large
habit the outermost branches of trees in the primary Platycerium leaves. Boy er ( 1 964) reports that
forest, but in secondary forest or on scattered trees Microgramma owariensis and Phymatodes scolopen
around villages and in farmland they can be found dria sometimes are associated with Platycerium
on any part of the tree. ferns. She also mentions that small plants of cocoa
The fertile and sterile leaves are of a different trees and oil palms were found in accumulations of
shape (Figs 84, 85). This dimorphism is of a par these ferns. Holtum ( 1 960) states that: " Cymbidiella
ticular function in the life of the plant. The sterile rhodochila in Madagascar is invariably found grow
leaves can be divided into two sections. The upper ing in association with a Platycerium. " The same
part is open towards the air on both sides, it is thin observation appears in Encyclopedia of C ultivated
and pergameneous. Their function is to collect the Orchids (Hawkes 1 965 : 1 3 9) with no reference to the
falling debris to be used in the building of humus. source.
The lower part of the sterile leaves is considerably
thicker. This part of the leaf is a deposition area for Epiphyte communities at altitudes above l 000 m
the debris that has been trapped between the upper Asplenium dregeanum-Peperomia sp. e.c.
part of the leaf and the branch or trunk. The space At the base of trunks (from the ground to roughly
between the sterile leaves is utilized by roots that 4-5 m height) Asplenium dregeanum and Peperomia
General observations
Ferns form the base in seven of the ten recognized
communities. In all these communities it is the
substrate-building capacity of the ferns that seem at
tractiv·e to the other species. Interaction and an
tagonism between ferns and orchids, as well as
between mosses and liverworths, have been reported
by Tixier. He found that the 'stations' rich in orchids
has generally no ferns and reciprocally. "Les
Fougeres presentent une axemie assez forte vis-a-vis
des Orchidees" (Tixier 1 966 : 1 24). However,
Fig. 86. Tridactyle tridactylites - Medinilla mannii
epiphyte community. Mt Gbahm 1 300 m. Drynaria rigidula was found to hold numerous
epiphytes. This statement may seem to contradict
the observations in the Nimba area, but the state
ment is more correct for the lower parts of the trunks
sp. dominate. They are so common that nearly every (a section which was particularly carefully examined
tree has at least one of them on its trunk. At these by Tixier). It is, however, hard to asess the amount
altitudes the filmy ferns are also common and par of competition between ferns and orchids. In many
ticularly on the trunks, and thus are often associated cases it is probably the low light intensities close to
with the species of this community . . the ground that limit the presence of orchids, and
The dominating fern, Asplenium dregeanum, mul higher up in the trees the evaporating power of the
tiplies by sending out modified stolon-like fronds air often limits the presence of ferns.
which take root and produce a colony of daughter
plants, expanding the community in many directions.
Since this community occurs on the trunks Epiphyte succession
relatively close to the ground no distance obser As the phorophyte grows it will pass· through several
vations had to be performed. 'environmental stages', e.g. it's place in the forest
alters, the size and the shape of the crown and the
Tridactyle tridactylites - Medinilla mannii e.c. morphology of the bark changes. These are primary
In the central part of the crown and especially changes. Secondary changes, e.g. establishment of
around the larger ramifications the orchid Tridactyle new species of epiphytes will effect the competition
tridactylites forms huge interwoven masses (Fig. 86). between the epiphytes. The epiphytic flora itself also
Here several plants occur, such as Polystachya lax participates in the successional processes through
iflora, P. leonensis and Asplenium dregeanum. active or passive contributions to the formation of
Scattered groups of filmy ferns may occur but very humus.
dominant are the mosses that occupy the major part Direct or indirect methods may be employed to
of the branches in aHd around this community. estimate the changes. The best method would
naturally be to map the epiphytic flora on a number
Bulbophyllum scariosum - Polystachya dalzielii e.c. of branches in the crowns of living trees. These
This somewhat less clearly defined community is branches could then be re-examined after certain
found in the central or outermost parts of the time intervals, thus giving a correct picture of the
branches. It includes a number of small sized development and eventually changes of the flora.
epiphytes, e.g. Xiphopteris villosissima, that are easi This method would be similar to the 'permanent
ly overlooked in a distance examination. quadrate' technique used in terrestrial ecology, ex
Two species of orchids dominate this community : cept for the changes in size of the branch caused by
Bulbophyllum scariosum which is easily seen, while growth or broken off pieces. Unfortunately this
the more tiny Polystachya dalzielii is harder to method is for practical reasons very difficult to use.
observe when not in flower. In this section of the The inaccessibility of the tree crowns has previously
crown the lichens cover the branches or hang down been discussed. The slow development of many of
( Usnea sp.) from the finest twigs. the epiphytic plants, e.g. the orchids, would also
and the plants involved in this process. The nature of In two accumulations each of a Microsorium punc
these deposits is very heterogeneous. They are built tatum and a Platycerium stemaria community no
up of material in all stages of decomposition, mostly traces of an older epiphytic flora could be found.
resembling peat. Occasionally the decomposition has
gone so far that a mineralization has begun. Such Oleandra distenta - Tridactyle armeniaca e.c.
soil-like deposits can be found at the basal parts on Accumulations from this community proved to be
the large branches of old trees. More commonly the most rewarding in the search for remains of an
deposits are built up by material that is partly earlier flora. Four such accumulations were in
decomposed. It is rather surprising that parts of dead vestigated, two of which held remains that could be
plants incorporated in the accumulation several identified. In one of them sterile leaves and rhizomes
years earlier frequently can be identified. This shows of Drynaria laurentii appeared in the bottom layer.
either that the plants are very resistent towards The other accumulation was so rich in old plant
decomposition, or that the micro-organisms that are remains that it will be presented in detail. This com
responsible for the break -down process are munity was located at the basal part of a large
restrained in their activities by the environment, branch I 7 m above the ground on a Lophira alata at
which is regularly subjected to drought. 600 m altitude. The thickness of the humus layer was
Several cross-sections were cut through the ac not less than 1 6 cm. In the bottom part of the ac
cumulations at a right angle to the axis of the cumulation the material was partly mineralized. The
phorophyte, with a sharp broadblade knife. This size of the investigated part was 0.2 x 4.0 m.
gave a glimpse of the 'stratification' in situ. The con The surface vegetation was a dense aggregation of
tent from the bottom layer was sifted through a sieve Oleandra distenta with the rhizomes hanging down
with a mesh of 10 x 1 0 mm. Remains of plants, such from the branch on both sides. Ten more species
as old pseudobulbs and rhizomes, could then much were present: Asplenium megalura, A rthropteris
easier be discovered. A total of 1 3 deposits of five orienta/is, Begonia polygonoides, Bulbophyllum
different epiphyte communities were examined. cocoinum, Brachycorythis kalbreyeri, Habenaria
leonensis, Nephrolepis undulata, Medinilla mannii,
A ngraecum birrimense - Raphidophora africana e.c.
Polystachya polychaete and Rhipsalis baccifera. The
Two communities, both occurring on the trunk 3-4
topmost 10 cm of this accumulation had a felt-like
m above the ground on two tall trees, were ex
structure, and was composed of living and dead
amined. The humus layer was thin and held in a ver
roots and rhizomes. Below this layer large quantities
tical position by a network of roots. Six species of
of partly decomposed plant material were found.
epiphytes were present : A ngraecum birrimense,
Sterile leaves and rhizomes of Drynaria laurentii
Calvoa trochainii, Diaphananthe rutila, Nephrolepis
were accompanied by leaves and rhizomes from
undulata, Polystachya rhodoptera and Raphido
Phymatodes scolopendria and pseudobulbs of
phora africana, with the first and last species
Graphorchis lurida and several unidentified
dominating.
Bulbophyllum spp.
In the bottom layer in one of the communities 3
Traceable remains of an earlier epiphytic flora
groups of pseudo bulbs from Polystachya affinis were
were found in 30.7 % of the accumulations ex
discovered.
amined. It is interesti.ng to note that the species that
Drynaria laurentii - Asplenium megalura e.c. were overgrown were all able to colonize clean bark
Three humus deposits from this community were ex surfaces, so called pioneer epiphytes, e.g.
amined. Two came from the central part of the Polystachya affinis, Drynaria laurentii, and several
crown and one from the outermost part. In one of Bulbophyllum spp.
the deposits from the central part of the crown some The expansive growth of the Drynaria fern will
old pseudobulbs probably produced by Bulbophyl regularly overgrow or surpress other plants of slower
lum lupulinum and pseudobulbs and aerial roots of growth, e.g. the many Bulbophyllum spp. that fre
Graphorchis lurida were discovered. quently occur in the same habitat.
In the 0/eandra distenta- Tridactyle armeniaca com
Microsorium punctatum - Peperomia sp. and munity it looks like several other epiphytes also are
Platycerium stemaria - Nephrolephis undulata e.c. active in the accumulation of humus, and that Olean-
Epiphyte Phorophyte
A B c
Asplenium megalura + +
Drynaria laurentii + + +
Elaphoglossum isabelense + +
Elaphoglossum salicifolium +
Oleandra distenta +
Phymatodes scolopendria +
Vittaria guineensis +
Aerangis laurentii +
Angraecum birrimense +
Angraecum distichum +
Angraecum subulatum +
Bulbophyllum distans + +
Bulbophyllum linderi + + +
Bulbophyllum maximum +
Bulbophyllum oreonastes + + +
Bulbophyllum phaeopogon +
Bulbophyllum saltatorium +
Bulbophyllum schimperanum +
Bulbophyllum schinzianum +
Chamaeangis vesicata +
Cyrtorchis arcuata +
Graphorchis lurida +
Listrostachys pertusa + + +
Fig. 88. Drynaria laurentii on a phorophyte in secondary Plectrelminthus caudatus +
forest. Mt Tokadeh 700 m. Polystachya polychaete + +
Polystachya saccata + +
Tridactyle anthomaniaca + + +
Tridactyle armeniaca + +
Tridactyle crassifolia +
Medinilla mannii + +
Peperomia rotundifolia +
Rhipsalis baccifera +
A B
dra distenta under certain conditions can utilize this
substrate. The fact that this community is restricted
to the basal area of the crown indicates that a large
accumulation of humus is not enough to promote a
successful colonization by Oleandra distenta.
which may be compared to the ratio in the dense had only four orchid species (36.3 %' in common
forest that was exactly 1 :2 ( 1 9 :38, Table 1 7). This is with the phorophyte from the most protected en
hardly surprising, since the higher light intensities vironment (A) (Fig. 89). This may indicate that most
will favour the orchids which also are able to with of the orchids are adapted to rather narrow en
stand the much drier environment of more exposed vironmental conditions, which also could explain
trees. Drynaria laurentii is the only fern that can their richness in species.
take advantages of these changes in the environment, The study of the changes during a two years
and occurs here in sizes and numbers never found in period in the epiphytic flora on a branch that had
the dense forest (Fig. 8 8). Rhipsalis cassytha is the fallen down from a large tree may serve as an exam
only species of the other vascular epiphyte group ple of changes that occur when the environment
that seems to thrive on these exposed phorophytes. becomes darker and more humid. The branch had
Comparison of the epiphytic flora on three fallen down over a small creek and was lying about
phorophytes of the same species (Lophira alata) of 2 m above the small stream. A stretch of six meters
approximately the same sizes, from the same altitude was examined, in November 1 96 7 and re-examined
but growing in three different habitats, may ex at the same time during the years 1 968 and 1 969
emplify the effect of the environment in more detail. (Table 34).
One phorophyte (A) was 42 m in size and grew in One must remember when analyzing the changes
a high forest surrounded by several other trees of the in the flora on this branch that it does not represent
same size. The second phorophyte (B) was 40 m and the same substrate as a living branch. The effect of
was left in a forest subjected to logging operations the decomposition of the bark remains unknown.
that had created a rather broken canopy. The third Anyway, the changes may give an idea of the
phorophyte (C) was 39 m and grew in a very ex tolerance or adaptability of the various species in
posed environment with several dead trees around it. volved.
A total of 3 2 species of epiphytes were recorded The original species composition shows a number
from these three phorophytes (Table 3 3). A more of sun-demanding orchids, pteridophytes and other
open environment seems mainly to limit the presence vascular epiphytes. Only after some months the
of non-orchid epiphytes. The number of orchid leaves of the orchids started to turn yellow. After one
species was about the same on all of the phorophytes year the Bulbophyllum species were dead and so
( 1 2, 1 2 and 1 1 ), but it should be noticed that the were the A ngraecum distichum and Polystachya sac
phorophyte (C) in the most exposed environment cata specimens. After one year three species of
Table 34. Changes in the epiphytic flora on a dead branch during a two years period. (See text.)
1 96 7 1 968 1 9 69
tree instead of falling down to the ground. This The prompt recovery of the host plant when the
creates the impression that the trees carrying this epiphytes are removed was experimentally proved.
epiphyte have more dead branches than others. Plants artificially infected with the fern Drymo
glossum showed a rapid decline within a year. An
Direct effects anatomical investigation revealed that in the infested
Cook-Melville ( 1 926) considers the epiphytic orchids branches the vessels were partly blocked by gum and
to be parasites on citrus trees in Puerto Rico. fungus and the cortex contained many hyphae. The
The generally accepted belief that epiphytes as a resultant deficient nutrient and water supply would
whole are limited to a non-parasitic life has been amply account for the inhibitory effect.
carefully examined by Ruinen ( 1 95 3), whose opinion The invading hyphae were traced microscopically
and interesting results will be summarized here. She to the place of exit from the epiphyte in which it
states that the typical epiphytes preferably develop formed a mycorrhiza. The inference drawn from this
on young, living tissue and that plants declining in is that the mycorrhizal fungus of the epiphyte is
health usually recover, if the epiphytes are removed. potentially parasitic on the supporting trees. It
Contrary to Went, Ruinen confirms the observation depends on the constitutional conditions presented
by planters on Java that cleaning their coffee, tea by the supporting tree, whether or not it will appear
and citrus trees from epiphytes causes the trees to as such.
become healthier subsequent to this treatment. It would seem hardly possible to speak of a sym
The early symptoms of decline under epiphytic biosis between epiphyte and supporting plant, were it
growth are difficult to define. They suggest a general not for the failures in cultivating some orchids, e.g.
deficiency, an aspect of ill-health, of failing. The Taeniophyllum sp. on inorganic substrate and,
leaves are no longer lush, still green but less so than moreover, for the marked preference shown by
usual, not as turgescent as the non-infested branches, epiphytes for some definite phorophytes under
but nevertheless not wilted. In this stage the develop otherwise identical environmental conditions. These
ing buds are slightly smaller. The leaf fall is conditions are undoubtedly inherent in the suppor
premature, therefore the vegetative period is ting plant itself, which obviously of(ers favourable
shortened. The noxious effect is localized at first to symbiotic relations with the epiphytes, and
the parts directly under the epiphyte, the other parts therefore, from the point of the support the sym
are only secondarily affected, viz. show the influence bioses should be qualified as antagonistic.
at a later date. Ruinen concludes that the generally accepted
The phenomenon, which is termed epiphytosis, belief that epiphytes as a whole are limited to a
may be described as a slow exhaustion of all partners saprophytic habit, seems not only open to discussion
of the biocoenosis one after the other (Ruinen but as such has been proved to be actually
1 95 3 : 1 08). As the epiphytes are known to be erroneous. (Cf. Harvais & H adley 1 967.)
mycotrophic, the most simple form in which the In a study of structural connections between
biocoenosis in its relations and deterioration might epiphytes and host plants, Furman ( 1 95 9 : 1 27) has
be represented is illustrated in Fig. 92. made observations similar to those of Ruinen. In a
Honduran cloud forest the interpenetration of root
hairs or root mycelium with host tissues was ex
amined. Vascular epiphytes, chiefly orchids,
bromeliads and ferns, were found to penetrate living
e p iph yte + fu n gu s + sup p o rt hosts' stem tissues with root hairs, or share a com
mon fungus mycelium with them. Frequently the
L
J
sy m biosis _j host tree stems had very thin, lightly suberized bark.
Schimper ( 1 935 : 3 1 1 ) have earlier stressed the fact
( antibiosis
that many epiphytes belong to those families known
for their mycorrhiza, e.g. Orchidaceae, Ericaceae,
Melastomaceae and 'perhaps' even other ones and
Fig. 92. Interactions between epiphyte, fungus and sup that the fungus might play an important role in the
port. (From Ruinen 1 95 3.) metabolism of the epiphyte.
A cta phytogeogr. suec. 59
Ecology of vascular epiphytes in West African rain forest 93
AUTHOR'S OBSERVATIONS
Any direct effect by the epiphytes on their
phorophytes has not been noticed. However, certain
species of phorophytes, e.g. Mitragyna ciliata and
Parinari excelsa (Fig. 1 8) with a rich epiphytic flora
often give the impression of suffering from
'epiphytosis' as described by Ruinen.
The 'preference' of certain epiphytes for a sub
strate of young bark or living tissues, as reported by
R uinen and Furman, should be reflected in the dis
tribution pattern. Species with such preferences
would have their main location in the outermost
parts of the crown, on the youngest branches and
twigs. Only one species, Bolusiella talbotii, shows
such a distribution pattern (Fig. 7 8).
There are, however, certain species of orchids that
seldom appear on the tall trees, but seem to prefer
younger ones and even shrubs. Such habitats are
found around creeks and small rivers, or in
'farmlands'. Five species are ± restricted to these
habitats : A ngraecum classensii, Diaphananthe den
siflora, D. rutila, Eurychone rothschildiana and
Rhipidoglossum paucifolium.
parts of the branches. It has generally been assumed genera of orchids throughout the world has generally
that the photosynthesis taking place in the roots of been considered to be an adaptation for preventing
the leafless orchids is sufficient for the needs of the water losses through the leaves (Kerr 1 972 : 307,
plant (Dycus & Knudson 1 95 7). However, there Teuscher 1 97 2 :497).
seems to be no experimental proof of this assump If one looks upon the distribution of the species
tion. The rate of photosynthesis in roots of orchids among e.g. the West African leafless orchid genus
with green leaves (Erickson 1 95 7, Dycus & Knud Microcoelia Lindl., they all appear in areas with con
son 1 9 5 7) clearly shows that these roots do not even siderable amounts of rain and with such minor
produce material that can support their own respira climatic disadvantages for epiphytes that a very
tion. Difficulties for leafless orchids growing on dead large number of leaf carrying epiphytic orchids with
substrate (Kerr 1 972 :307) (Teuscher 1 972 :497) or green leaves occur in the very same area (Fig. 9 5).
when transplanted (Stewart, 1 970 :90, 1 97 3 :803) can The possibility that the leaves have become obsolete
be used as argument that these orchids to a certain due to a partly parasitic life, should be carefully ex
extent are dependent on living tissues. amined.
The reduction of leaves that occur in several
Environmental influences can be referred to three altitudes (500-700). Comparisons will also be made
main groups : climatic factors, substrate and biotic with the epiphytic flora at altitudes higher than those
factors. Although environmental influences result in the investigated area, as observed on the high
from a combination of these factors, they will be ex mountains in East Africa. From Tanzania, the
amined separately. Their possible combined effect epiphytic flora in dry environments will exemplify
will be considered later (Chapter VI). climatic influences not represented in the investigated
area.
500-700 33 94 1 :3
Fig. 96. Parinari excelsa densely overgrown by epiphytes.
1000- 1 300 28 27 1:1
Nimba Range, 1 3 50 m.
Pteridophytes
Arthropteris monocarpa
Asplenium aethiopicum
Asplenium barteri
Asplenium geppii
Asplenium monanthes L. Mant.
Asplenium rutifolium (Berg.) Kunze var. bipinnatum (Forsk.)
Schelpe
Asplenium theciferum (Kunth) Mett. var. concinnum (Schrad.)
Schelpe
Asplenium erectum Bory var. usambarense (Hieron.) Schelpe
Drynaria volkensii Hieron.
Loxogramme lanceolata
Lycopodium ophioglossoides Lam.
Oleandra distenta
Pleopeltis excavata (Bory) Sledge
Pleopeltis macrocarpa (Bory) Kaulf.
Orchids
Bulbophyllum stolzii Schltr.
Polystachya zambesiaca Rolfe
Stolzia nyassana Schltr.
I No epiphytes
ll Asplenium nidus L.
Ill
Pteridophytes
Orchids
Ancistrorhynchus refractus Kraenzl.
Bulbophyllum plathyrachis Schltr.
Cirrhopetalum umbellatum (Forst.f.) Hook. & Arn.
Oberonia disticha (Lam.) Schltr.
Polystachya adansoniae
Rangaeris muscicola
Tridactyle anthomaniaca
Tridactyle bicaudata
Fig. 98. Simplified drawing showing the epiphytes utilizing
the humus held by the root system of Asplenium
IV
theciferum on a Nuxia congesta R.Br. (Loganiaceae).
Ngong Hills 2400 m, Kenya. (A) Asplenium theciferum, Pteridophytes
The effect of the altitude on the epiphytes has been Other vascular epiphytes
compared with either the water balance, e.g. drought Rhipsalis baccifera
correlated to low temperatures (Went 1 940 :94, Tix
ier 1 966 : 3 6) or the temperature. V Orchids
Bolusiella iridifolia (Rolfe) Schltr.
The lower the temperature (Table 2) the higher the
Tridactyle anthomaniaca
relative humidity (Fig. 9) and the presence of local
cloud systems or mist, at the highest parts of the
Nimba Range (Fig. 1 1), will naturally influence the
evapotranspiration and favour less drought resistant
species such as filmy ferns, and certain pterido Temperature influence on the occurrence of
phytes. Mosses and lichens are also favoured by the orchids has been stressed by Rupp ( 1 969 : 1 ) : "The
higher light intensities that exist at all levels in the epiphytes reach their greatest development in the
more open forest type at the crest of the range. tropics, decreasing in numbers towards cooler
regions. This fact is well illustrated in Eastern Table 38. Occurrence of epiphytes in two different forest types.
Australia, where Queensland h as approximately 1 00 (Trees < 5 m are excluded). Great Ruaha River 400 m, Kidatu,
known species of epiphytes, New South Wales 5 2, Tanzania.
l
(Baker) Summerh., Cyrtorchis arcuata subsp.
variabilis and C. arcuata subsp. whytei (Rolfe)
Summerh., Microcoe/ia exilis Lindl., Polystachya Evaporation
tessallata. The only epiphytic fern was Phymatodes
sco/opendria (one 'stand'), which was partly growing
on some wet rocks in a shady habitat.
Trees along two parallel strips 50 m long and 10
Fig. 99. Simplified drawing showing the distribution of
m wide were examined ('distance' method) (Table epiphytes on a Combretum tree close to the Great Ruaha
3 8). The first strip was situated in the riparian forest River. Kidatu 3 90 m, Tanzania.
\
/
I
/ 1 m 60-80 % 90- 1 00 %
·..
· ....
· · · · · · · · · · · · · ··.
Saturation deficit 46 m 1 2- 1 6 mm 6- 1 2 mm
\\
,, 1 m 6 - 1 0 mm 0-4 mm
··-·· · · · · · · · · �-�:-:':'::.
Hours with rei. humidity 46 m 1 0- 1 4 hrs 8- 1 2 hrs
less than 9 5 % 1m 1 0- 1 2 hrs 0-2hrs
700 goo 1 100 13 o o 1 500 1 700 1 9 oo
with a maximum in the morning and also in the after altitudes, given as a percentage of the light in the open.
taken in a high forest with the canopy around 40 m and was managed by an assistant. The recording
above the ground in the Seka Valley. Common was performed by holding the cell at the growing site
species of trees in this forest were : Parinari excelsa, of the plant to be investigated. To reach the plants
Heritiera uti/is, Calpocalyx aubrevillei and growing in more inaccessible places, the light cell
Chidlowia sanguinea. The ground was covered with was attached to a four m long aluminium pipe. At
dead leaves and the herb flora was sparsely the end of the pipe, where the light cell was attached
represented, e.g. : Selaginella vogelii Spring and at a 90-degree angle towards the pipe, a V -shaped
Selaginella versicolor Spring. On the lower parts of metal piece was connected to a ball bearing on the
the trunks a few ferns were noticed, Hymenophyllum pipe. When the metal piece with its open end was
kuhnii and A splenium barteri. held against the substrate, the whole pipe could be
At 1 300 m the samples were taken at the crest of turned around on its axis, and thus moving the light
the main ridge 300 m SW of the point where the cell at the same time. By placing the light cell at the
borders of Liberia, Guinea and the Ivory Coast actual growing site of the epiphyte, and then turning
meet. The forest in this locality almost exclusively it, the maximum light intensity was found.
consisted of Parinari excelsa with a somewhat The tree chosen was a Parinari excelsa growing in
broken canopy at 1 0- 1 2 m level above the ground a typical section of the forest that covers the narrow
which was irregularly covered with herbs. crest of the main ridge at 1 300 m altitude from the
The high light intensities in the montane forest is mark of the three national borders down to the min
remarkable, but judging from the abundance of ing area. The canopy in this single dominant
plants on the ground it was not totally unexpected. Parinari forest was at 1 0- 1 5 m above the ground
In a montane forest in Ecuador, Whitmore and partly open. There were a few trees and shrubs
( 1 968 :238) found that the canopy was more of smaller dimensions. e.g. Ochna membranacea
transparent than in a lowland forest. Oliv., reaching up to the lower parts of the crowns of
Particular interest has been paid to the illumina the emergent trees. The tree investigated had a size
tion of the undergrowth and the effect of the sun of 23 m and a circumference of the trunk at breast
flecks on the forest floor, e.g. Evans ( 1 939) in height of 1 62 cm and was heavily branched, with the
Nigeria, Gusinde & Lauscher ( 1 94 1) in Congo, Eid first ramification starting 3 . 5 m above the ground.
mann ( 1 94 1) on Fernando Po, Evans et al. 1 960 and The foliage was concentrated to the outermost
C achan ( 1 963) in the Ivory Coast. The shade il branches which made movements easier in the cen
lumination was found by all of them to be 1 % or less tral part of the crown. At the basal part of the trunk
of the outside light. The light intensity in the sun the cover of epiphytic mosses and filmy ferns was
flecks in British Guiana varied between 1 0.4 and 1 00 %. Higher up on the trunk ( 1 -3.5 m) the
72 % (C arter 1 934). The spectral composition of the epiphytes were more or less concentrated to one side
light reaching the forest floor as examined by Evans (S), with the plants appearing in small patches with
( 1 939) showed approximately 40 % portion of the clean bark surface in betwe(;!n. In the basal parts of
wave-lengths above 7000 A (infra-red light). Similar the crown Tridactyle tridactylites formed dense, un
observations have been reported from Puerto Rico tidy 'stands'. In the central part of the branches the
(Johnson & Atwood 1 970). This indicates a con mosses e.g. Porothamnion hildebrandtii (C.M.)
siderable increase in the transmission of the forest Fleisch, were very abundant, forming large cushions.
canopy just beyond the red end of the visible spec In the apical part of the branches the lichen cover
trum. The effect of this portion of light on the plants was pronounced especially on the twigs, where an
is unknown but it can probably not be used in the Usnea sp. with hanging growth habit was noticed.
photosynthesis, ·since it is outside the range of Bulbophyllum scariosum and Polystachya dalzielii
chlorophyll's absorption. As the forest became were also abundant.
denser a decrease in the proportion of blue light was A total of 1 5 species of epiphytes were growing on
also noticed. the tree, 7 pteridophytes, 5 orchids and 3 other
vascular epiphytes. Thus the pteridophyte/orchid
A t the growing sites of the epiphytes ratio was 1 .4 :1 . A total of 1 1 9 recordings, which
The same photoelectric cell as described earlier was were estimated to cover 80 % of the plants present,
used. The recording equipment stood on the ground were performed during a 3 hours period in the mid-
Table 43. Light intensities as a percentage of the light in the between maximum and minimum intensities should
open at the growing sites of epiphytes on a Parinari excelsa. be noted with caution, since sun-flecks with a very
Nimba Ridge, 1 300 m. short duration may be responsible for the high
figures.
Epiphytes No. of Light intensity
(\
60
parison (Table 43).
It is interesting to note that the arbitrary division
of the trees into 5 sections that has been used
throughout this study, was in this tree not correlated I �
to the light intensities (Table 44).
No marked difference in the light intensities could �J �
\
30
1 I n the open
2 D i apha n a nthe pel l u cida
3 V a n i l l a c re n u l a ta
4 Art h ropte r i s m o n o c a rpa
18 . 3
06. 0 0 12.00 1 8 .0 0
The minor variations that can be observed Bulbophyllum inflatum, growing on a trunk I .5 m
throughout the day are probably caused by haze. above the ground at 1 300 m altitude, give the varia
The more gentle peaks in the curve for tion in light intensities for the orchid which grows in
Diaphananthe pellucida probably indicate that this the most shady habitat of all the orchids in the Nim
species recieves a more indirect type of light. The ba area (Fig. I 07). These two curves can also serve
series of distinct peaks with a very short duration as an example of the 'micro light climate' that
that occur between 1 2°0- 1 4 °0 on the Vanilla curve is probably is of great importance for the epiphytes.
a result of direct sunlight that has been able to Note the similarity between the two curves of March
penetrate down to this level. In denser shade closer 1 8 and 20, 1 970 (March 1 9 was a cloudy and rainy
to the ground the penetration of direct light is less day) when the light that was registered reached the
likely. The light level tends to be more uniform plant between 1 4°0- 1 530• For many plants that live in
throughout the day as can be seen for A rthropteris a superficially dark environment these short periods
monocarpa. of higher light intensities may be of critical impor
In the montane forest the light regularly penetrates tance in their survival. It may also influence the oc
the lower parts of the vegetation. The curves for currence of small groups of several species that com-
1 A s p l e n i u m m e ga l u ra
2 Aspl e n i u m afri ca n u m
3 Aspl e n i u m b a rteri
1 P o l yst a c hy a pa n i c u l a t a
2 P o l ysta chya l a x iflora
3 Po l ysta chya ra m u l os a
J.
genera A splenium and Polystachya were chosen for
study. One species in each genera represented a
specific level in the rain forest. The lowest level (sec
tion I) was represented by Asplenium barteri and
Polystachya ramulosa, the intermediate level (11) of
Asplenium africanum and Polystachya laxiflora, and
the crown stratum (Ill + IV) of A splenium megalura B ra c hyc o ryt h i s Po lyst a c hya B o l u si e l l a
and Polystachya paniculata. ka l b reye ri i l a xifl o ra ta l bo t i i
Substrate
Fig. 1 09. Records of epiphytes distributed between the
different light classes (from Table 27, Brachycorythis
PROPERTIES OF THE SUBSTRATE AND kalbreyeri from Table 29). (A) heavy shade, (B) open
THEIR INFLUENCE ON THE EPIPHYTES shade, (C) full sun.
epiphytic flora frequently were found among species Bark pH and the epiphytic flora
with smooth or hard bark, and fast growing species. The samples were taken from representative areas in
Boyer ( 1 964) concluded that it probably was the the vicinity of epiphytes in various sections of the
smooth nature of the bark that was the cause of the phorophyte. They were all taken from trees felled the
absence of Platycerium ferns on Musanga Smithii. same day, except for a living Parinari excelsa tree at
In the Nimba area trees with a defoliating b,ark (e.g. 1 300 m altitude which was climbed. The bark pieces
Distemonanthus benthamianus), seem to be very dif used in the survey had a size of 6- 1 0 cm2• They were
ficult for the epiphytes to colonize. This has also pulverized and dried in 24 hrs in 5 5 % relative
been observed elsewhere (Dudgeon 1 923, Pessin humidity. Distilled water twice the dry weight of the
1 925 :34, Oliver 1 930: 14, Went 1 940 :95, Rupp sample was added. The measurements were per
1 969 :XI). Voorhoeve ( 1 96 5 :200), however, states formed 24 hrs later on an electric pH-instrument.
that colonization of flaking bark does occur, and in The sampling procedure, e.g. cutting a piece of bark,
some cases the epiphytes even prevent the shedding involves a great deal of subjectivity, since the actual
of the bark, e.g. on Gilbertiodendron preussii. The choice of bark pieces may influence the result. It was
absence of epiphytl c orchids on the oil palm is hard found that dead, corky pieces gave higher pH values
to correlate to any morphological characteristics of than living bark, a fact that is well known. There are
the phorophyte, but may possible be attributed to also difficulties in utilizing a completely identical
germination difficulties of the orchid seeds. When method of sampling due to the variation in the bark
full-grown epiphytic orchids are transplanted on oil morphology. Some species have a thick easily
. . . • .•
I 1
H e r i t i e ra 11
.. alkalinity of the substratum does not influence the
.I • •
Ill
uti l i s :
IV
V
I
11 •• •
occurrence and distribution of Polypodium
Loph i ra : 1: I
.. .. . polypodioides."
Ill
a l a ta ..
IV
V
I I
11
M itra g y n a .
. . ..
Ill
c i l i a ta . I Exudates
IV
V
1••
P a r i n a ri
I
11 I
.. . .
"Orchids seem to dislike trees which exude a latex
..
Ill
excel sa IV
V : I• juice such as most Ficus species and tree Euphor
I•
I
P a ri n a ri 1 3 00 11
.. bias." (Piers 1 968 :4).
... .
Ill
excel sa m
..
IV
V
I
I
I ..
In the Nimba area several species of phorophytes
..
11 11
P i pt a d e n iastru m Ill
:i which · exude a latex juice have a rather rich epiphyte
africa n u m ..
IV
V
flora, e.g. Chlorophora regia, Uapaca guineensis and
even the rubber tree, Hevea brasiliensis, while others
Fig. 1 1 1 . pH in water extracts of bark from different show an opposite tendency, e.g. the figs.
phorophytes. Yekepa. The toxic effect of certain exudates must also be
detachable bark while others have a thin flaky, corky considered. Barkman ( 1 95 8 : 1 34) suspects the 'robin'
or deeply fissured one. Differences in pH value for in the bark of the R obinia tree to be the cause of the
samples of large and small pieces of bark have also poverty of non-vascular epiphytes of this tree.
been reported (Sjogren 1 96 1 : 1 02). Large differences in the presence of orchids
The pH values exhibit a ± significant pattern between various species of trees of the genus Quer
within the phorophytes (Fig. 1 1 1 ). The lowest values cus were observed by Frei ( 1 9 7 3 a : 3 1 1 -3 1 2) in a
were obtained from the upper part of the main trunk cloud forest in Mexico. Two species in the genus had
and the outer branches. only a few species of orchids, two. others carried
The pH of the bark has been studied by many many orchids, and one was never observed with any
authors using various methods. It has been observed orchids whatsoever.
that the pH values decrease from the bases of the When analyzed, the bark from the species with
trees and upward, probably due to decreasing dust many orchids showed no presence of inhibitory sub
content. The pH has also been found to vary accor stances. The bark from the phorophytes devoid of
ding to the time of day (Pessin 1 925 :34). orchids, contained gallic and egallic acids, which are
Du Rietz ( 1 945 : 1 98- 1 99) distinguishes three known to be growth inhibitory substances
groups of bark based on the pH values, 'rikbark' (hydrolyzable tannins). (See also Frei 1 973b : 70 1 -
(rich bark) pH 5-7, 'overgimgs-fattigbark' 708.)
(intermediate bark) pH 4-5 and 'extrem fattigbark' B ark from several tropical species of trees e.g.
(extremely poor bark) pH <4. When trying to Ficus pumila contain strong growth inhibitors
evaluate the influence of the pH value of the bark on (Bovey & Diaz-Colon 1 959 :256).
the epiphytic flora one should remember that the pH The importance of the leaching of growth
value might have changed since the time when the regulating substances from the vegetation has also
seedlings were established. The long span from ger been stressed by Tukey Jr ( 1 970 : 1 60).
mination to a fully developed plant exemplified by Went ( 1 940 :90) assumed that the rich epiphytic
the orchids emphasizes this problem. flora on Castanopsis argentea could be connected to
The epiphytes themselves influence, in a complex the high amounts of tannin in the water on the bark
and imperfectly known way, the acidity in their en surface. The excretion of water-soluble substances
vironment. This can be done by collecting dust and by the bark or the leaves has seldom been measured.
forming humus, through photosynthesis, by release Schweitzer (quoted from Ruinen 1 95 3 : 1 5 1 ) . found
of lichen acids and exchange of ions (Kolkwitz 1 9 3 2, that a total leaf surface area of one sq. m of coffea
Skye 1 968 : I 07). The acidity also regulates the leaves, leached in distilled water during one hour,
nitrification since this process is strongly pH depen- yielded 2 1 -3 5 mg mineral substances, Erythrina
their excretions.
An efficient adaptation to intercept leachates from
the overhead canopy is shown by certain bromeliads.
It has been shown experimentally that metabolites
fall in the well (formed by the diverging leaf bases)
from which they are absorbed and utilized in growth
(Tukey Jr 1 9 70 : 1 5 5 - 1 60).
Humus deposits
Origin and composition
The accumulation of humus starts with the decom
position of the outer layers of the bark. The bark is Fig. 1 1 2. O rigin of substrate and nutrients of importance
for the epiphytic flora.
continuously renewed from within, so that it
automatically provides a steady supply of new organic in origin. This can be compared to the soil
material for decomposition. This is m ainly carried under the trees containing 3 .0- 3 . 9 % of organic
out by fungi, which in many cases are connected to matter. I 0-23 % of the material belonged to a frac
the roots of the epiphytes. tion less than 2 mm in diam. The rest consisted of
To the material from the bark external material is living, or dead and more or less decomposed parts of
added. Dead leaves, twigs, small branches, etc., from leaves and roots.
the phorophyte itself can be assumed to form the The accumulation of organic material through
bulk of the deposits. In this more or less accidental wind, animals, and water running along branches
process the growth habit of the phorophyte and and stem amounted to 5- l 0 % of the total weight.
structure of its bark are of importance. The root The bulk of the organic material was derived from
systems of the epiphytes also help to catch debris in the tissues of the fern itself. The addition of organic
the many niches they form. Certain epiphytes play a material from sources outside the fern was con
more active role in the catching of falling debris. sidered to play a minor role in the nutritional supply
Usually a niche between the plant itself and the sub for this epiphyte.
strate is formed in which debris from above can be The formation of the humus was slow. It was es
caught, e.g. Drynaria laurentii, Platycerium spp. tim ated that it takes 1 0 years to form a layer of
(Fig. 1 1 2). The term 'sole epiphytique' has been used I 0 cm thickness. It was also noticed that the material
by Boyer ( I 964) for the humus collected by that was incorporated was only slowly decomposed
Platycerium ferns. She is of the opinion that these ac despite the rich microflora. The external contribution
cumulations of organic material can hardly be com to the assemblage of humus may be small in volume
pared to the soil in its proper sense. These ac but important in its nutritional effect on the epiphytic
cumulations have attracted the interest of several flora.
authors, e.g. Miehe ( I 9 1 1 ), Paulian ( 1 945) and Dust carried by the winds may be Jf considerable
Klinge ( I 9 6 3). importance in certain areas. Every year in the winter
In the Ivory Coast Boyer ( 1 9 64) found that in the months the Harmattan winds fill the air with a fine
humus of the 'leaf basket' of Platycerium angolense dust in vast regions of West Africa. It is easily
and P. stemaria 6 3 .4-90. 9 % of the dry weight was observed that trees close to dusty roads often show a
+�
�
855
shorter rainfall and the slow release of it is naturally
of great importance for the epiphytes.
30 Cs5
0
pH of humus deposits
0 15 30
Thirteen different humus deposits uti lized by
DAYS
epiphytes were examined. The methods used in these
Fig. 1 1 3. Water holding capacity o f humus from different determinations are the same as described in the in
epiphyte communities, in o/o of the wet weight. vestigation of the bark, i.e. the bark piece has simply
been replaced by a piece of humus (50 g dry weight).
l u xu riant growth of epiphytes (Thorold 1 9 5 2 : 1 3 9, Rather high pH values were obtained, con
Richards 1 964 : 1 1 4 ). Ants and termites carry siderably higher than those of the bark, Table 45.
material of inorganic as well as organic origin up Schnell ( 1 95 2 :5 8-5 9) gives the pH in the humus of·
into the trees. Termite nests built from earth, and the a Bulbophyllum sp. as 4.8 and for a Listrostachys sp.
earth-covered tracks of these insects are common on to 5 . 7. Boyer ( 1 964) reports the pH of humus from
the trees. Richards ( 1 964: I l l) states that the roots Platycerium stemaria as 4. 6-6.4 (6.0 for P.
of flowering epiphytic plants and ferns are one of the angolense).
chief nesting places for arboreal ants. Adamson
( 1 943) suggested that certain epiphytes in the rain Correlation between substrate and epiphytic flora
forest of Trinidad m ay be at least partly dependent Many epiphytes show a high frequence in the recor-
on soil carried up into the trees by termites. In an
Table 45. pH in water extracts from humus deposits with gro-
analysis of the mineral content in the humus from a wing epiphytes. Median value of five samples.
Platycerium fern in the Nimba Mountains Schnell
( 1 95 2 :58) found that 3 8 % of the mineral content Phorophyte Section pH Epiph. utilizing the humus
elaborate on the origin of the larger sized particles Heritiera utilis V 6 .0 Drynaria laurentii
Nephrolepis undulata
that can hardly be transported by the wind, but it
Lophira alata Ill 5.8 Vittaria guineensis
can be mentioned that the earth in this particular Lophira alata IV 7.0 Oleandra distenta
region is very rich in hematite. fridactyle armeniaca
Lophira alata V 6.8 Drynaria laurentii
samples were taken. The samples were divided in two africanum Microsorium punctatum
Triplochiton scleroxylon IV 5.5 Platycerium angolense
groups, one was placed in an air-conditioned room at
Triplochiton scleroxylon IV 4.8 Platycerium stemaria
5 5 % relative humidity, the other placed outdoors
A cta phytogeogr. suec. 59
Ecology of vascular epiphytes in West African rain forest 1 1 1
major groups of epiphytes (Fig. 1 1 4). Total, light 1 5 .4 7 7.0 7.7 1 00. 1
Most pteridophyte species (5 1 %) grow in minor
humus deposits. The orchid species occurred on bark
(50 %). The other vascular epiphytes were found in will be used (Table 4 7). Phorophytes with a high
large humus deposits (54 %) (Table 46). number of species of epiphytes are :
Regarding the interrelationship between epiphytes A mphimas pterocarpoides Cryptosepalum tetraphyllum
and phorophytes three case may be distinguished in A ntho_notha fragans Entandrophragma utile
Calpocalyx aubrevillei Heritiera uti/is
the area studied (denoted A, B and C below).
Canarium schweinfurthii Lophira alata
Ceiba pentandra Mitragyna ciliata
A. Phorophytes with an abundance of epiphytes Chlorophora regia Parinari excelsa
In the Nimba area a 'preference' by epiphytes for Combretodendron macrocarpum Piptadeniastrum africanum
Table 47 a. Records of pteridophytes and other vascular epiphytes growing on different phorophytes in the Nimba a r e a .
<f)
"'
H
.s
.s:::
bl)
:;:;
"'
.s:::
"'
"0
� �]
�- 3 .g
E o �
_g � �
0.. C.) U
Pteridophytes
X • + X " 0 0 " 10
A n trophyum i m m e rsum
. • •
A. n1anntanun1 0 + 7
A r thropteris monoca rpa + X + + + + + X + + + 20
A . orienta l i s + · o + + + x + + X x · x · + x x + o + X + 29
A . palisoti I
A splenium :lethiopicum X " 4
A. nfricanum X + • + + X 0 · 14 . 3
A. barteri + + x · x + + X 20
A. drege:mum + + 7
A . geppii + X + X 0 + " 23 + 2
A. hemitomum 4
A. megalura + • X 0 X 0 0 x x + x x o + X X + 34 + + 4
• • • 0 0 • • •
A. variable var. paucijugum
e
Da va Ilia cha rophy \ loides • + 0 X
•
+ + 0 • • + 0 • 0 • 21 + + 4
Drynaria \au re n t i i + x x x + x x x o x x x x x x 0 X X X X X X X X X + X X X X X X X + + X + + 43
Elaphoglossum burteri X ' • + + X ' ' ' ' X • • 0 10
E. cheva lieri • • X • 4
E. isabelense X + x + o · + · + o x + x x + x x x x x o X + · + 2� r 3
E. kuhnii • + . 2
+ •
E . salicifolium x + + + · o + o + o o x x • + X X + + 3� 2
Lomariops;s guinecnsis 8
Loxogramme lanceolata · x x + · x x · · · + + + X + X + 0 • + X 27 + .
Lycopodium m i ldbraecli i 2
L. warneckei x · o x + · · o + · o o x o o o x x o x · -t o · o + 0 + 0 X 0 X + 33 + + 4
Microgramma owa r i e n s i s x · x x o o o o · · · o + · x x · x + • o + · · + u x x x t + 0 X 30 . + 4
Microsorium punctatum x x o x o x x x · x o o x x x x · o o x x + x x x x x x x x x x x · x 38 + + 5
• • X • • • • 0 • • • 0 X ·t· • • •
Nephrolepis bi serrata X 0 + X • • + X • 0 ' X + + + + • X · • X • + 0 + X + 24
N. undulata o x x · x + x · + + x o o x x x x x + x x x x x 'l: x x x x · x · · o · x x 0 X 0 X 38 r + 3
0\eandra dis Le n ta + x x o + x · x • x x x • o x + + + + + 0 X • + X X X X + • 34 + + 4
Phymatodes scolopendria x o x o · + · + x + X + ' X X X ' + X ' X X + 30
Plat.vceriom angolense 0 X 17
P . slemari::t • 0 X + 0 23
•
Pyrrosia mcchowii 3 2
• •
Tectaria angelicifolia X 6
T. fernandensis
Vi ttaria guineensis -t o x + x + x x x x + x x x x + + x + x x x x + x + X + X X X 42 + + + 5
Xit:.hopteris oosora 2
X . serrulata
X . villosiss ima
0 9 N ffi 0 00 0 � M O � � � M m N 9 9 0 N N � M� 9 - � ffi • M m - � • M � m 9 � ,......., ......, M ':"l :" I L::'
Total no . of pteridoph.vtc s pp. M 1"""1 � !"""' .-1 1"""1 1"""1 M ....-! C\1 J""-4
..-( ,..... ,....; 1'""""4 r-t .....-t C\l .--t C'? N ...-! � .....-t C'. J M C\1 .--t �
,....; C'\1 1"""1
....... M CO 1'"""4 .....-1 CO N CO 1"""1 ,....; M t-- C\1 .......
....... 7'1 ,.......; .....-1 - .....-I
Total no . of other 3 4 5 8 3 5 2 5 2 3 7 8 7 6 4 4 5 5 6 4 1 D 5 1 2 9 2 9 3 1 9 1 3 S 8 3 4 3 7 G 3 3 2 - 9 5 4 5 :2 3 - s 1
vas e . epiphyte spp.
x = close observalion, o = d i s L1.n<"e obse r v a t i o n , + o occasional observation. Only one k i n d o f record is g i v e n , i . e . when a c l ose observation record occu r , d i s La nce and
occasional observations a re excluded. If a distance observation is given no close record ex i s t and onlv when close and distance records a re lacking, occasional records
are show n .
i
0 0 0
ll
0
c ::' i� "' "' c: 0 > >
�
1
"'
1 �! �-�;
�
�� �� �
i
"'
:r. ..;,.
""
I !
"
� �
:;;; =
i
�
i l
.:::: :r.
2 c
�
�� �
!i �
�1 l
X c
�
"'
� ·�
��
2-: T.'
� c
�� :r.. "'
�
t
¥. -:;,
;:
lf 2 -g � §
J � 1 3 �
-;:
· "
3 �
�· �g �
- -§
7.
�
·7.
�- �
a;· �
�
�- c
�
�
� ==
� � �7
high fo re s t
��
Phoroph\·tcs o f the
t\lbi7.i:."t glaberrim:1
Albizia 7..\·gia
so l e �
u ��
20
A l t n i : boon i
=. � A m phimas pleroc:tl'poidt':-;
Anthocleista nobi l i s
+ 0
�
A nthonnth:l f l':tgan s
. 0 A n L i a r i s toxic<l l'i;J
:: .- ')')
� iG
5·f 5·a ��
l3el' l i n i a confus:1
Bomba:x buonopozense
llusse:1 occidcntalis
%
C a l pocal.\'X aubre\·illci
Can;trium schwe i n l u r l h i i
42 CC'ib:l pentandra
C' h itl lowi a s a ngu i nc a
� � !� C h l o r·ophora regi:l
� g. 2 6
. C h r' vsoph_,· l lum perpulchrum
'"'
� � 29 C o m h rctodendron macroca rpum
:J � -!.) 'L
3
C o u l a cdulis
5 =. 24
g-
��
C r·,,·ptns(•p:llum '..C li'aph, · l l u m
i7 Uani e l l i n ogea
� � 26 39
Distcmonantbus benlh:rmianus
· E n t a nd l'Ophr·agma u t i l e
f':l 0
[ � �5
Et·_vthrophleum ivorcnse
F:t�·a r:r t c s s m a nn i i
a. � ��
Gua 1·ea ced rata
HeJ"iticra u t i l i s
Khav:1 a n t h o theca
i i �; � \'
Lophira :.1 la w
I
Lo oa trichi lioidcs
0 o lH
!\'Jammca africana
=.. 2. 47
• 0 /.
�
Mi t ra g.vna c i l i a t:l
c;;· 23 + . N:IUclca d id c rr i ch i i
[� � �� P:n inari :1uhre v i l l e i
:-·
Purin:lri excclsa
�� 13
Pa rina ri glabra
g 1 :.:!
('tl f':l
Parki;I bicolor
''0
�
. i4
Pen t.a c le thra m a c r u p h v l l : l
::: Pentatlesm;1 blltytacca
Q. g- 47 PipL;tdcn i : t s u·um a!'ric:rm1m
;;:· ::: 3
5� 20
P\Tnanthus angolcnsis
Rhodognaphalon b re v i cu s �
�5
g 0 Sacoglo l t i s gaboncnsis
., ;g !)
9
Tcrminalia i\·ot·e n s i s
� 0 Termin:tlia supe rba
� � 41
g. 3 26
Triplochiton sclct·ox\'lon
l':.�paca gu incen s i s
::,:1 t.l 2 :1
� g·
Vilex micrantha
- w !...: X t � _. 1..;. -1 � _. t' �� W ;: -1 ,_. '.ZJ � � I'<J ;:., .;.,, 1..;. � I'<J 1..:1 -1 =": ,_.
�!
,:': :.Jl � � ::': .;; ::: ,_. _. � � ,_. ,_. 0) � :j ; � �-=; � .l::o. � -; ::; :-: - ::;:: _. I � ::; t.;. � 1...;1 ':.1. t .;.,; �1 .l::o. �J. :: � ,_. :.a C': ;: ;:; � � � � � � _. :.;1 =t; � o;..;: .t:.. :,.,) ::::: ¥1 C: � ::-: 1 w 1.:.. rv <x :::t. 1..; l Total no. o f phorophv1c spp
to the construction of a list of favourite 'host trees' from all over the world. Piers ( 1 968 :4) reports that
for certain popular orchids, for aiding the collector in Acacia and Ficus trees are avoided by most
field-work (Sulit 1 950 :6-8). Morris ( 1 970 :5) uses the epiphytic orchids. The same applies to tree Euphor
term 'orchid-prone' for trees that are attractive to bias, exotic conifers and Eucalyptys trees. Moreau
epiphytic orchids. The following species of ( 1 943 : 8), however, remarks that conifers such as
phorophytes have been reported to carry an abun Juniperus bermudiana are colonized by orchids with
dance of epiphytes : a surprising frequency.
From Australia Rupp ( 1 969 :XI) confirms that
Liberia: Cola nitida (Harley 1 95 5 :7 1 ), Mitragyna ciliata
(Voorhoeve 1 965 :324), Parinari excelsa (Voorhoeve Eucalyptys trees are usually barren of orchids. The
1 965 :3 1 9) absence of epiphytes on figs has been observed by
West Africa: Parinari excelsa (Jaeger et al. 1 968 :270, several authors. Schnell ( 1 9 70 :309) mentions that
Schnell 1 970 :309) Ficus mucuso lacks epiphytes and Went ( 1 940) notes
Malawi : Brachystegia spiciformis, Bridelia micranta, /lex that humus epiphytes are lacking on Ficus in
mitis, Parinari mobola, Podocarpus milanjuanus, volucrata Bl.
Syzygium cordatum, Uapaca kiekiana (Morris 1 9 70 :5)
Java: Castanea argentea, C. javanica (Went 1 940)
C. Spe�ific relationships between phorophyte and
West Borneo : Vitex sp. (Schuitem�ker, in Went 1 940 :95) epiphyte
Florida, U.S.A. : A nnona glabra L., Taxodium distichum In the Nimba area, no · specific relationship between
L. (Frei 1 9 73b : 70 1 )
phorophyte and epiphyte has been observed, even if
Haiti : Eugenia jambos L . (Curtis 1 946), .Euphorbia
the total dominance of Parinari excelsa at the
lactera Haw. (C urtis 1 94 7)
highest parts of the mountains has resulted in a near
S. America: Crescentia cujete (Schimper 1 888 :95)
ly absolute correlation between this tree and some
B. Phorophytes with few epiphytes epiphytes naturally occurring at these altitudes (Ta
In the Nimba area epiphytes are poorly represented
ble 47).
on a number of species of phorophytes (Table 4 7), in
There are several reports of a relationship between
particular on Fagara tessmannii, Terminalia ivoren
a particular species of phorophyte and a particular
sis, T. superba and Musanga cecropioides, con
species of epiphyte. Piers ( 1 968 :4) states: "Some
sidering the abundance of full-grown individuals of
epiphytes have marked idiosyncrasies regarding their
these phorophytes. Less frequent phorophytes with a
host tree, e.g. A nsellia nilotica has a strong
sparse epiphytic flora are e.g. A lbizia glaberrima, A .
preference for the Doum Palm (Hyphaene thebaica),
zygia.
the Baobab tree (A dansonia digitata) is the typical
The absence of epiphytic orchids on the trunks of host of A ngraecum dives and Polystachya adan
Elaeis guineensis is obvious. However, they support soniae, while Aiirangis thomsonii attaches itself
a rich pteridophyte flora (Fig. 7 1). Only two almost exclusively to the rugged stems of the gigant
epiphytic orchid species have been observed on this ceder (Juniperus procera) of the Kenya highlands."
palm : Graphorchis lurida and Habenaria procera. Rupp ( 1 969 :XI) gives a similar observation :"one or
(The Kew Herbarium. has 26 collections of the latter two of the ironbark eucalypts are favoured by Den
species. The growth habitat is given for fifteen of drobium aemulum, which for this reason is often
them. Eighty per cent of these were epiphytes, all of callen 'Ironbark Orchid' ". More examples of this
them on Elaeis guineensis.) kind are listed by Richards ( 1 964 : 1 1 9). Morris
In a study of the epiphytes on Elaeis guineensis in ( 1 9 70 :5) found few specific host/orchid relationship
the former Belgian Congo (Zaire), Van Oye ( 1 924) in Malawi, but observed : "There are however a few
did not mention any orchids, but several orchids which would appear to associate with
pteridophytes and figs. In the Philippines it is even specific hosts . . . Bulbophyllum intertextum on New
stated that "Orchids will choose any other tree in the tonia buchananii, Polystachya johnstonii on Vellozia
wild than the palm" (Sulit 1 950 :8). spendens come to mind."
Ruinen ( 1 9 5 3 : 1 02) has suggested the term 'axeny' Moreau ( 1 943 :8) is a little more hesitant : "there is
for this condition of general inhospitality of trees little evidence of specific relation between orchid and
which never or only occasionally bear epiphytes. tree host, except that certain aberrant Polystachyas
Trees belonging to this group have been reported that are otherwise terrestrial also grow on the curios-
l y looking fibrous stem o f Vellozias, but no other leaves, seem to be of minor importance for the rain
host." (See also Richards 1 95 7 :5 6 8.) forest species in general (Odum 1 970 b). Voute
B oyer ( 1 964) states that the occurrence of the ( 1 946, 1 9 64) states that insect outbreaks do not oc
epiphytic ferns Platycerium stemaria and P. cur in tropical forests because of the stability of
angloense on various phorophytes in the Ivory checks and buffers that act on any species that
Coast, does not permit any conclusions regarding becomes overabundant.
their preference for particular species of
phorophytes. Nutritional importance of animals and micro-
Whether a relation between a certain species of organisms
phorophyte and epiphyte really exist is doubtful. In The presence of a rich fauna in the crown of the rain
many cases the epiphytes have for various reasons a forest trees naturally influences the nutritional supp
very limited choice of host trees. Given the oppor ly of the epiphytes through excreta from m ammals,
tunity, many of these epiphytes now occurring on a birds, snakes, lizards, frogs and insects. The effect of
specific host tree, might very well thrive on other bird excreta was early recognized. Sernander ( 1 9 1 2)
trees. Another fact that favours the theory of a coined the term 'ornithocoprophily' for such plants
specific relation between host and epiphyte is the that were favoured by bird excrement (or rather their
poor knowledge of the presence of the epiphytes. In nitrogen content). Du Rietz ( 1 93 2) showed that bird
creasing information on the ecology and presence of excrements also increased the pH of the substrate.
the epiphytes will probably decrease the number of Trees with nesting birds, edible berries, as well as
'specific relationships'. those used as sleeping trees and observation posts,
are known to attract nitrophytic lichens. It is im
possible to even try to estimate the total effect of
Biotic factors nutrition provided by the animals in the diversified
rain forest. It should at least be kept in mind during
Several of the biotic factors have already been the analysing process.
presented and discussed in Chapter Ill. The role of the microorganisms in the nutritional
balance is unknown, but Ruinen ( 1 95 6) found
Human influence bacteria of the genus Beijerincka on leaves on nearly
The human influence on the rain forest effects the all trees and shrubs tested in Java and Sumatra.
epiphytes indirectly. In the Nimba area patches of These nitrogen-fixing bacteria obtain their minerals
scattered trees that remain after forest destructions and some organic material from substances excreted
are the particular growing sites of several orchid by the leaves. Blue -green algae of several kinds are
species. These are species of the deciduous forest, always abundant on the bark of trees in the wet
e.g. A nsellia africana, Plectrelminthus caudatus, tropics but their nitrogen-fixing capacity is unknown.
Polystachya paniculata, P. puberula and Rangaeris (cf. Ruinen 1 96 1 ).
rhipsalisocia. Bond ( 1 959) reports : "There are also nitrogen
fixing micro-organisms present on leaves and
Animal influence branches of trees, which doubtless adds to the
Direct effects, as judged by signs of being eaten or nitrogen available to the roots of epiphytes, as
used by animals, on the epiphytes is . very seldom similar microorganisms do in the soil."
observed. Some species, however, show signs of fre The high amounts of nitrogen in the humus from
quently being eaten (probably by insects), e.g. Polypodium polypodioides puzzled Pes sin ( 1 925 : 3 2) :
Pyrrosia mechowii, A ncistrochilus rothschildianus, "It is indeed remarkable that the nitrogen o f the
A ngraecopsis elliptica and Polystachya ramulosa. humus on the bark of the tree is frequently as high
Effects on the leaves by herbivores or other and at times even higher than that of the most fertile
agents, as can be judged by holes in the falling soils."
V F i l my ferns
4.0
Fig. 1 1 5. General di stribution of epiph ytes on tall
phoroph ytes.
Ecology of vascular epiphytes in West African rain forest 1 1 7
branches, (A, for species in section Ill, B, for species Xiphopteris oosora Begonia mannii
in section IV and C, for species in section V). The A ncistrochilus rothschildianus B. polygonoides
tion of light intensity and substrate composition is with differences in the evaporating power of the air.
not accurate for the majority of epiphytes. This in It is more probable that low light intensities are the
vestigation showed that the different epiphytes often limiting factor since substrates of similar composi
have a marked preference for a certain kind of sub tion and evaporating power of the air equal to the
strate and light intensity. The almost total absence of conditions in the crowns also exist at various places
epiphytes from the crowns of the trees in the lower in the lower parts of the forest.
strata of the forest cannot be satisfactorily explained
Ecologie des epiphytes vasculaires dans la foret dense humide d'Afrique occidentale
repartition des epiphytes a ete enregistree. Pour Colonisation et effets sur les phorophytes
simplifier l'enregistrement le phorophyte a ete divise Les differentes methodes indiquent que ce sont par
en cinq sections (Fig. 76). On a applique trois ticulierement les phorophytes les plus grands qui
methodes differentes de denombrement : les obser sont de preference colonises par les epiphytes. Les
vations proches, les observations a distance et les pteridophytes semblent etre les epiphytes pionniers
observations occasionnelles. L'observation proche tandis que les orchidees sont les dernieres a s'etablir
implique des investigations sur des arbres abattus, (Fig. 8 7). Certains arbres ay ant une riche flore
!'observation a distance est une etude du tronc et epiphytique par exemple Mitragyna ciliata et
d'une branche complete (ou plus rarement un arbre Parinari excelsa, souffrent apparemment
entier) d'un arbre sur pied a l'aide de jumelles ou "d'epiphytosis" (Ruinen 1 95 3). Les effets des
d'un telescope sur trepied. orchidees sans feuille sur les phorophytes peuvent
L'observation occasionnelle est reduite aux in aussi indiquer un parasitisme partiel.
vestigations obtenues a partir seulement d'une partie
limitee du phorophyte. Les resultats de cette derniere Influences de l'environnement sur les epiphytes
methode ne sont utilises que pour completer ceux des Climat. Le rapport pteridophytes epiphyti
deux autres. ques/orchidees se modifie avec !'altitude. A la base
On a egalement examine le substrat et l'eclaire de la zone (500-700 m) la proportion
ment au lieu de croissance de !'epiphyte individuelle. pteridophytes/orchidees est approximativement de
Le substrat a ete divise en trois classes : ecorce, 1 :3 mais, a 1 000- 1 300 m, elle est de 1 : 1 . Des
ecorce avec depots faibles d'humus, et depots impor variations similaires ont egalement ete observees sur
tants d'humus. La lumiere a aussi ete divisee en trois les hautes montagnes d' Afrique orientale.
classes : plein soleil, demi-obscurite et obscurite. Dans les milieux secs, ainsi qu'on l'a observe en
Tanzanie, le nombre des epiphytes decroit rapide
ment (specialement les pteridophytes) mais l'abon
Occurence des epiphytes dance de certaines orchidees epiphytiques peut sur
Dans la haute foret 50,4 % des arbres hauts de dix prendre. D ans les montagnes meridionales de Tan
metres ou plus portaient des epiphytes, par contre zanie on a observe que tous les arbres (� 4 m) cVune
dans une foret "secondaire" on descendait a 1 4,8 %. terre boisee Brachystegia portaient l'orohidee I
Il etait done evident que particulierement les Tridactyle tricuspis.
orchidees etaient rares dans une foret "secondaire".
Certaines especes d'arbres peuvent abriter un grand Substrat. La valeur pH d'echantillons d'ecorce
nombre d'especes d'epiphytes par ex. Heritiera uti/is provenant du meme phorophyte presentait d'assez
(30 spp.) Mitragyna ciliata (28 spp.) (Tableau 42). grandes variations. C 'est dans les parties exterieures
Aussi le nombre des individus ("stands") d'epiphytes des grandes branches et la partie superieure du tronc
sur certains arbres etait-il tres grand, par exemple un principal que l'on trouvait les valeurs les plus basses.
seul Heritiera uti/is ne portait pas moins de 1 85 7 On n'a cependant pas pu trouvef de corrClation per
epiphytes et un Parinari excelsa 1 1 7 1 . tinente entre la valeur pH de l'ecorce et la flore
epiphytique. Les valeurs pH des depots d'humus
etaient beaucoup plus elevees que celles de l'ecorce.
Groupements epiphytiques La majorite de l'espece pteridophyte "preferait"
On dit qu'il y a groupement epiphytique quand trois les depots mineurs d'humus tandis que les orchidees
especes au minimum d'epiphytes forment une unite poussaient surtout sur l'ecorce et les autres epiphytes
et quand la distance entre deux de ces epiphytes et la vasculaires dans les grands depots d'humus.
troisieme ne depasse pas 0,5 m. On distingue sept
groupements epiphytiques differents au-dessous de Correlation entre l'abondance d'epiphytes et l'espece
1 000 m et trois autres de plus a des altitudes de phorophyte
superieures (Tableau 30). Les grands specimens d' Heritiera uti/is, Lophira
Les fougeres forment la base de sept des dix alata, Mitragyna ciliata et Parinari excelsa sont
groupements. Il semble que ce soit leur capacite de riches en epiphytes tandis que par exemple
formation de substrat qui attire les autres epiphytes. A nthocleista nobilis Fagara tessmannii, Terminalia
ivorensis et T. superba sont frequemment plus ou La distribution des epiphytes dans le phorophyte
moins depourvus d'epiphytes. On n'a pas observe de Chacune des cinq sections du phorophyte est
correlation specifique entre }'epiphyte et le dominee par un groupe d'epiphytes (Fig. 1 1 5).
phorophyte. La plupart des especes d'epiphytes ont une dis
tribution plutot limitee sur le phorophyte, c'est-a-dire
Lumiere. L'intensite de la lumiere au sol dans une dans une ou deux sections. La majorite des especes
haute foret a la base de la zone etait de 0,36 % de la poussent dans la couronne. On distingue six modeles
lumiere exterieure mais dans une foret Parinari a differents de distribution sur les grandes branches
1 300 m d'altitude elle etait de 6,65 % (Tableau 42). (Fig. 1 1 9).
Dans les parties de base et les parties centrales des La distribution des epiphytes ne depend pas seule
grandes branches d'un arbre Parinari excelsa a 1 300 ment de !'evaporation (Pessin 1 925) mais elle est
m la lumiere etait reduite de 70-85 % par rapport a egalement influencee par la preference marquee
celle de la demi-obscurite (Tableau 43). La variation d'une certaine sorte de substrat et d'eclairement. Les
quotidienne dans l'intensite de la lumiere au lieu de pteridophytes presentent une distribution qui va a
croissance de certains epiphytes a revele des l'encontre d'un accroissement possible de !'evapora
differences tres significatives entre les differentes es tion. La distribution des orchidees semble toutefois
peces (Fig. I 06- 1 08). En general, on trouve les etre le resultat d'un equilibre entre le desir d'une
pteridophytes dans la demi-obscurite tandis que les grande intensite de lumiere et la capacite de resister a
orchidees sont plus egalement reparties entre une plus grande evaporation d'air qui s'ensuivrait.
!'habitat en demi-obscurite et )'habitat en plein soleil.
Adam, 1.G. 1 96 6 - La vegetation du Mont Nimba au Bernhard-Reversat, F., Huttel, C., & Lemee, G. 1 9 72 -
Liberia et sa protection. - Notes africaines, I. F.A.N. Some aspects of the seasonal ecologic periodicity and
1 1 2. plant activity in an evergreen rain forest in the Ivory
1 969 - Etude comparee de quelques forc�ts ouest Coast. - In Golley, P.M. & Golley, F.B. (ed.)
africaines. - Bull. I.F.A.N. 3 1 : 3 4 1 -409. 1 97 2 :2 1 7-2 1 9.
1 9 70 - Etat actuel de la vegetation des monts Nimba Blum, G. 1 93 3 - Osmotische Untersuchungen in Java. I.
au Liberia et en Guinee. - Adansonia 1 0 : 1 9 3 -2 1 1 . - Ber. Schweiz. Bot. Ges. 42 : 5 5 0-680.
1 9 7 1 - Flore descriptive des Monts Nimba. - Mem. Bond, G. 1 9 5 9 - The incidence and importance of
M us. N at. Hist. N at. B, 20, 22. biological fixation of nitrogen. - Adv. of Sci. 1 5 :3 82-
Adamson, A.M. 1 943 - Termites and the fertility of soil. 3 86.
- Trop. Agric. Trin. 20(6) : 107- 1 1 2. Boyer, Y. 1 964 - Contribution a la etude de
Ake Assi, L. 1 963 - Contribution a la etude floristique de l'ecophysiologie de deux fougeres epiphytiques :
la Cote d'Ivoire et des territoires limitrophes. 2, Platycerium stemaria et Platycerium angolense. -
Monocotyledones et Pteridophytes. - Lechevalier, Ann. Sci. Nat. 1 2(5): 8 7-228. Paris.
Paris. Bovey, R.W., & Diaz-Colon, J.D. 1 969 - Occurrence of
Allee, W.C. 1 926 - Measurement of environmental fac plant growth inhibitors in tropical and subtropical
tors in the tropical rain forest of Panama. - Ecology vegetation. - Physiol. Plant. 22:253-259.
7 :273-302. Burgeff, H. 1 936. - Die Samenkeimung der Orchideen.
Alien, S.E., C arlisle, A., White, E.1., & Evans, C.C. 1 968 Fisher, Jena.
- The plant nutrient content of rainwater. - 1. Ecol. - 1 95 9 - Mycorrhiza of orchids. - In Withner, C .L.
5 6 :497-504. (ed.) 1 95 9 : 36 1 -3 9 5 .
Alston, A.H.G. 1 95 9 - The Ferns and Fern-Allies of Cachan, P. 1 963 - Signification ecologique des variations
West Tropical Africa. - Crown agents, London. microclimatiques verticales dans la foret sempervirante
Ames, 0. 1 922 - Observations on the capacity of orchids de basse Cote d'Ivoire. - Annales Fac. Sci. Univ.
to survive in the struggle for existence. - Orch. Rev. Dakar 8 :89- 1 5 5.
30:229-234. C achan, P. & Duval , J. 1 963 - Variations microclimati
Anonymus. 1 968 - How old is an orchid plant? - Amer. ques verticales et saisonnier dans la forc�t semper
Orch. Soc. Bull. 3 7 :405 . virante de basse Cote d'Ivoire. - Ibid. 5 -98.
Arditti, 1. 1 966 - Flower induction in orchids. - Orch. C arter, G.S. 1 934 - Reports of the C ambridge expedition
Rev. 74 :208-2 1 7. to British Guiana, 1 9 33. Illumination in the rain forest
- 1 96 7 - Factors affecting the germination of orchid at ground level. - J. Linn. Soc. (Zool.) 3 8 : 5 79-589.
seeds. - Bot. Rev. 33 : 1 -97. Childers, N.F., Cibes, H.R., & Hernandez-Medina, E.
Arditti, 1. & Dueker, J. 1 968 - Photosynthesis by various 19 5 9 - V anilla - The orchid of commerce. - In
organs of orchid plants. - A mer. Orch. Soc. Bull. 3 7 : Withner, C .L. (ed.) 1 95 9 :447-508.
862-866. Coe, M . & Curry-Lindahl, K . 1 96 5 - Ecology of a
Arditti, 1., Healy, P., &· Ernst, R. 1 972 - The role of Mountain : First report on Liberian Nimba. - Oryx
mycorrhiza in nutrient uptake of orchids 11. - Ibid. (1ourn. of the fauna preserv. soc.) 8(3).
4 1 :503-5 1 0. Cook-Melville, T. 1 926 - Epiphytic orchids as an
Ashton, P.S. 1 969 - Speciation among tropical forest epiphytic pest on citrus trees. - 1ourn. Dept. Agr.
trees. - Bioi. J. Linn. Soc. 1 : 1 5 5 - 1 96. Puerto Rico 1 0 : 5 -9.
Atlas of Tanzania, 1 967 - Ministry of lands, settlement & Cole, N.H.A. 1 968 - The vegetation of Sierra Leone. -
water development. Dar es Salaam. Njala, Sierra Leone.
Aubreville, A. 1 93 8 - La foret coloniale; Les forets de Coombe, D.E., & Hadfield, W. 1 962 - An analysis of the
l'Afrique occidentale fran�aise. - Ann. A cad. Sci. growth of Musanga cecropioides. - 1. Ecol. 50 :22 1 -
colon. 9 : 1 -245. Paris. 234.
Bainbridge, R., Evans, G.C., & Rackham, 0. (ed.) 1 968 - Cooper, G . P . & Record, S.J. 1 93 1 - The evergreen
Light as an ecological factor. - The British ecological forests of Liberia. - Yale Univ. Forestry Bull. 3 1 .
society. Blackwell Scientific Publications, Oxford and Coutinho, L.M. 1 964 - Untersuchungen tiber die Lage
Edinburgh. des Lichtskompensationspunkts einiger Pflanzen zu
Barkman, 1.1. 1 95 8 - Phytosociology and ecology of verschidenen Tageszeiten mit besonderer
cryptogamic epiphytes. - Van Gorcum & Co, Assen. Berticksichtigung des 'de-Saussure-Effektes' bei
Sukkulenten. - Beitr. zur Phytologie (Walter Focke, W.O. 1 893 - tfber epiphytische Gewiichse. Nach
Festschr.) 1 0 1 - 1 08 . Briefen von Fritz Muller. - Abh. naturwiss. Ver.
Curtis, C . H. 1 93 3 - How orchid seeds are dispersed. - Bremen 1 2 :5 62.
Orch. Rev. 4 1 : 1 1 2- 1 1 6. Foggie, A. 1 947 - Some ecological observations on a
- 1 943 - Pseudo-bulbs. - Ibid. 5 1 : 1 3 7. tropical forest type on Gold Coast. - J. Ecol. 3 4 :88.
Curtis, J.T. 1 93 9 - The relation of specificity of orchid Frei, J.K. 1 973 a - Orchid ecology in a cloud forest in the
mycorrhizal fungi to the problem of symbiosis. - mountains of Oaxaca, Mexico. - Amer. Orch. Soc.
Amer. J. Bot. 26 :390-399. Bull. 4 2 :307-3 1 4.
1 946 - Nutrient supply of epiphytic orchids in the 1 973 b - Effect of bark substrate on germination and
mountains of Haiti. - Ecology 27 :264-266. early growth of Encyclia tampensis seeds. - Ibid.
1 94 7 - Ecological observations on the orchids of 4 2 : 70 1 -708.
Haiti. - Amer. Orch. Soc. Bull. 1 6 :262-269. Freise, F. 1 936 - Das Binnenklima von Urwiildern in
19 5 2 - Outline for ecological life history studies of subtropischen Brasilien. - Petermanns Mitt. 8 2 : 30 1 -
vascular epiphyte plants. - Ecology 3 3 :5 50-5 5 8 . 307.
Curry-Lindahl, K. 1 969 - Report t o t h e government of Furman, T.E. 1 95 9 - Structural connections between
Liberia on conservation, management and utilization epiphytes and host-plants. - Proc. 9 :th lnt. Bot.
of wildlife resources. -,--- I.U.C.N. Morges, Switzerland. Congr. 2 : 1 27. Toronto.
(Stencil). Garay, L.A. 1 972 - On the origin of Orchidaceae. 11. - J.
Davies, T.A.W. & Richards, P.W. 1 93 3 , 1 934 - The Arnold. Arbor. 5 3 :202-2 1 5.
vegetation of Moraballi Creek, British Guiana. An Gessner, F. 1 9 56 - W asserausha1t der Epiphyten und
ecological study of a limited area of tropical rain Lianen. - In Ruhland, W. (ed.) 1 95 6 :9 1 5 -9 50.
forest. Parts I and 11. -J. Ecol. 2 1 : 3 50-3 84 ; 22 : 1 06- Goebel, K. 1 888 - Morphologische und Biologische Stu
1 55 . dien. I. Ueber Epiphytische Fame und Muscineen. -
Dirmhirn, I . 1 96 1 - Light intensity at different levels. - Ann. J ard. Bot. Buitenz. 7 : 1 - 7 3.
Trans. Roy. Ent. Soc. London 1 1 3(2) :270-27 5 . 1 922 - Erdwurzeln mit Velamen. - Flora (N.F.) 1 5 : 1 -
Dudgeon, W. 1 923 - Successions o f epiphytes in the 26.
Quercus incana forest at Land our, Western Golley, P.M. & Golley, F.B. (ed.) 1 972 - Tropical
Himalayas. - Journ. Ind. Bot. Soc. 3 :2 70. ecology with an emphasis on organic productivity. -
Dunn, H.A. 1 94 1 - The major orchids of Panama. The Papers from a symp. on trop. ecology, in New Dehli
physiography of their habitats. - Amer. Orch. Soc. 1 97 1 . Athens, Ga.
Bull. 9 :25 1 -267. Gore, A.J.P. 1 968 - The supply of six elements by rain to
Du Rietz, G.E. 1 9 32 - Zur Vegetationsokologie der os an upland peat area. - J. Ecol. 5 6 :483-495.
tschwedischen Kustenfelsen. - Beih. Bot. Centralbl. Gorgla, T.A. 1 96 9 - Inventory results, East Nimba
49 :6 1 - 1 1 2. National Forest and parts adjacent. -_Report to Lam
1 945 - Om fattigbark- och rikbarksamhiillen. - eo J.V. Op. Co. Yekepa, Liberia. (Stencil).
Svensk Bot. Tidskr. 3 9 : 147- 1 50. Greendale, D.J. & Nye, P.H. 1 964 - Organic matter and
Dycus, A.M. & Knudson, L. 1 95 7 - The role of the nutrient cycles under moist tropical forests. - Proc.
velamen of the aerial roots of orchids. - Bot. Gaz. 1 0 :th Int. Bot. Congr. :248. Edinburgh.
1 1 9 :78-87. Grubb, P.J., Lloyd, J.R., Pennington, T.D. & Whitmore,
Eaton, J.S., Linkens, G.E. & Bormann, H.F. 1 97 3 - T.C . 1 963 - A comparison of montane and lowland
Throughfall and stemflow chemistry in a northern rain forests in Ecuador. - J. Ecol. 5 1 :567-60 1 .
hardwood forest. - J. Ecol. 6 1 :495-508. Guillaumet, J. 1 967 - Recherches sur la vegetation e t la
Eggeling, W.J. 1 947 - Observations on the ecology of the flore de la region du Bas C avally (Cote d'Ivoire). -
Budongo rain forest, Uganda. - Ibid. 3 4 :20-87. O.R.S.T.O.M., Paris-Orsay.
Eidmann, H. 1 94 1 - Meine Forschungsreise nach Gusinde, M. & Lauscher, F. 1 94 1 - Meterologische
Spanisch Guinea. - D. Biologie 1 0 : 1 - 1 3. Beobachtungen im Kongo-Urwald. - Sitzungs. Akad.
Erickson, L.C. 1 95 7 - Respiration and photosynthesis in Wiss. Wien. 1 50 :2 1 8-347.
Cattleya roots. - Amer. Orch. Soc. Bull. 26 :40 1 -402. Gardlund, T. 1 96 7 - Lamco i Liberia. - A.-W.,
Evans, G.C . 1 939 - Ecological studies on the rain forest Stockholm.
of Southern Nigeria. 11. The atmospheric environmen Hall, B.J. & Bowling, J.C. 1 96 9 - Field key to the
tal conditions. - J. Ecol. 27 :43 6-482. epiphytic orchids of Ghana based on characters of
1 968 - Model and measurement in the study of shoots and infructescences. - Adansonia 9( 1 ) : 1 39-
woodland light climates. - In Bainbridge, R., et al. 1 55 .
(ed.) 1 96 8 : 5 3-76. Harley, G.W. 1 94 1 - Native african medicine, with
Evans, G.C., Whitmore, T.C . & Wong 1 960 - The dis special reference to its practice in the Mano tribe of
tribution of light reaching the ground vegetation in a Liberia. - Cambridge, Mass.
tropical forest. - J. Ecol. 48 : 1 93 -204. Harley, W.J. 1 95 5 - The ferns of Liberia. - Contrib.
Exell, A.W. & Launert, E. (ed.) 1 9 70 - Flora Gray Herb. Harvard Univ. 1 77 :5 8- 1 0 1 .
Zambesiaca. X, Pteridophyta. - Crown agents, Lon 1 95 6 - Plants of the tree trunks i n Liberia. - Amer.
don. Fern J. 46 :65-96.
- 1 9 5 9 - Handbook of Liberian ferns. - Ganta, Liberia. Hutchinson, J. & D alziel, J . M . 1 95 4 - Flora of West
Harris, J .A. 1 9 1 8 - On the osmotic concentration of the Tropical Africa. 2 :nd ed. Vol. 1 , part I. Revised by
tissue fluids of phanerogamic epiphytes. - Amer. J . Keay, R . W .J . � Crown agents, London .
B ot. 5 :490- 506. 1 9 5 8 - Ditto . Vol. I, part 2. Revised by Key, R . W . J .
1 9 34 - The physio-chemical properties of plant saps in - Ibid.
relation to phytogeography. - M inneapolis. 1 96 3 - Ditto. Vol. 2. Edited by Hepper, F.N. - Ibid.
<2.
H arvais, G. & H adley, G. 1 9 7 - The relation between 1 96 8 - Ditto. Vol. 3 , part 1 . Revision edited by
host and endophyte in orchid m ycorrhiza. - N ew Hepper, F . N . - Ibid.
P hytol. 66 : 205-2 1 5 . Iwanoff, L. 1 9 28 Zur M ethodik der
H awkes, A.D. 1 965 - Encyclopaedia of cultivated Transpirationsbestimmung am Standort. - B er.
orchids. - Faber & Faber, London. Deutsch. Bot. Gesell. 43 :306-3 1 0.
Heagarty, C . P. 1 9 5 5 - O bservations on the germination Jaeger, P., H alle, N . & Adam, J . G. 1 9 6 8 - Contribution a
of orchid seeds . - Amer. Orch. Soc. B ull. 24 :45 7-464. la etude des Orchidees des Monts Loma (Sierra
Hilitzer, A . 1 9 25 - La vegetation epiphyte de l a B oheme. Leone). - Adansonia 8 :2 6 5 - 3 1 0.
- P ubl. Fac. Se. Univ. Charles, Prague 4 1 : 1 -200. Jaeger, P. & Adam, J . G . 1 9 72 - Contribution a !'etude de
Holdridge, L . R . 1 9 70 - A system for representing struc la vegetation des Monts Loma (Sierra Leone). -
ture i n tropical forest associations. - In Odum, H .T. & Compte rendu des seances de la societe de
Pigeon, R . F . (ed . ) 1 9 70, C h apter B - 1 2. biogeographie 422-424 : 7 7 - 1 03 .
Holtum, R . E . 1 960 - The ecology of tropical epiphytic Jarvi s, P . G. & J arvis, M . S . 1 96 3 - The water relations of
orchids. - Proc. Third World Orch. C onference. Lon tree seedlings. I V Some aspects of the tissue water
don. relations and drought resistance. - P hysiol. Plant.
Hopkins, B. 1 9 70 - Vegetation of the Olokemeji forest 1 6 : 5 0 1 -5 1 6.
reserve, N igeria. VI, The plants of the forest site with Johansson, D. 1 9 74 - Rhipidoglossum paucifolium, a
special reference to their seasonal growth. - J . Ecol. new African species of Orchidaceae. - B ot. N ot.
5 8 : 7 65 - 7 9 3 . 1 2 7 ( 1 ).
Hosokawa, T. 1 943 - Studies o n t h e life-forms o f Johnson, P . L. & Atwood, D . M . 1 9 70 - Aerial sensing
vascular epiphytes a n d t h e epiphyte flora o f Ponape, and photographic study of the El Verde rain forest. -
M icronesia. - Trans. N at. Hist. Soc. Taiwan 3 3 : 3 5 - In Odum, H.T. & Pigeon, R . F . (ed.) 1 9 70, C h apter B
5 5 , 7 1 -8 9 , 1 1 3 - 1 4 1 . 5.
1 94 9 . Studies of the life-form of vascular epiphytes and Jones, E . 1 9 60 - Contribution o f rainwater to the nutrient
the spectrum of their life-forms. - J . J ap. B ot. 24 :4 1 - economy of Northern N igeria. - Nature 1 8 8 :4 3 2.
45. K amerling, Z. 1 9 1 2 - De verdampin·g van epiphyte
1 9 5 0 - Epiphyte-quotient. - B ot. M a g . Tokyo 63 : 1 8- Orchideen. N atuurk. Tijd schr. N ed.-Ind.
20. 7 1 :5 4-72.�
1 9 5 2 a - A plant-sociological study in the mossy Karsten, G. 1 8 95 - M orphologische und biologi sche
forests of Micronesian i sl ands. - Mem. F ac . Sci. Untersuchungen iiber einige Epiphytenformen der
Kyushu Univ. E ( 1 ) :6 5 - 8 2 . Molukken. - Ann. J ard. B ot. B uitenz. 1 2 : 1 1 7- 1 9 5 .
1 9 25 - Uber mantelformige organe bei Epiphyten und
1 9 5 2 b - A synchrological study in t h e swampforests
Wurzelklettern. ·- Flora 1 8- 1 9 ( N . F.) :3 00- 3 1 I.
in the Micronesian Islands . - I bid. : 1 0 1 - 1 23 .
K eay, R . W.J. 1 9 5 9 - Explanatory notes to vegetation
1 9 5 3 - O n the nomenclature of aerosynusiae. - P roc.
map of Afric a published on behalf of A . E.T.F.A.T.
7 :th. lnt. B ot. Congr. :69 1 -694. Stockholm.
Oxford Univ. Press, London.
1 95 4 a - On the structure and composition of the
Kerr, A . D . 1 9 72 - The leafless Orchids. - Amer. Orch.
C ampnosperma forests in P alau, M ironesia. - Mem.
Soc. Bull. 4 1 : 3 07-309.
Fac. Se. Kyushu Uniy. E( 1 ) : 1 99-2 1 8 .
K linge, H. 1 963 - Vber Epiphytenhumus aus El
1 9 54 b - On the C ampnosperma forests of Yap,
Salvador. - Pedobiologia 2 (2) : 1 02- 1 07 .
Ponape and K usaie in M icronesia. - I bid. : 2 1 9-243.
Knapp, R. 1 9 73 - D ie Vegetation von Afrika. - Fisher,
1 95 4 c - On the C ampnosperma forests of Kusaie in Stuttgart.
M icronesia, with special reference to the community Knudson, L. 1 9 22 - Non-symbiotic germination of
units of epiphytes. - Vegetatio 5 - 6 : 3 5 1 -3 60. orchid seeds. - B ot. Gaz. 73 : 1 - 2 5 .
1 95 5 - On the vascular epiphyte comm unities in Kochenderfer, J. & Lee, R . 1 9 73 - Indexes t o transpira
tropical rainforests of M icronesia. - Mem. Fac. Sci. tion by forest trees. - Oecologogica Plantarum 8 : 1 7 5 -
Kyushu Univ. E(2) : 3 1 -44. 1 84.
1 95 7 - Outline of the mangrove and strand forests of Kolkwitz, R . 1 9 3 2 - Urwald und epiphyten. - Ber.
the M icronesian i sl ands. - Ibid. (2) : 1 0 1 - 1 1 8 . Deutsch. Bot. Gesell . 50 : 1 1 0- 1 1 6 .
1 9 6 8 - Ecological studies of tropical epiphytes in K unkel, G. 1 9 62 a - Uber die F arne Liberias ihre
forest ecosystem. - In M isra, R. & Gopal, B. (ed.) V orkommen und Verbreitung. - Ber. Schweiz. B ot.
·
1 968 :482-50 1 . Gesell. 72.
Hunt, P . F . 1 9 6 7 - Size of the orchid family. - Orch. Rev. 1 9 62 b - Aufzeichnungen iiber die Vegetation des Mt.
75 :229. Wutivi-Gebietes (Nordwest-Liberia). - Wildenowia
Rupp, H.M.R. 1 96 9 - The orchids of New South Wales. Stewart, J. & C ampbell, B. 1 9 70 - Orchids of tropical
- National herb., Sydney. Africa. - Allen & Co., London.
Rutter, A.J. & Whitehead, F.H. (ed.) 1 9 63 - The water Stacker, 0. 1 929 - Eine Feldmethode zur Bestimmung
relations of plants. - British Ecological Soc. Symp. der momentanen Transpirations- und
Ill. Blackwell, Oxford and Edinburgh. Evaporationsgrosse. I, Il. - Ber. Deutsch. Bot. Gesell.
Sanford, W.W. 1 96 7 - Orchids of West Africa. - Amer. 4 7 : 1 26- 1 29, & 1 30- 1 3 6 .
Orch. Soc. Bull. 3 6 :963-969. Sulit, M.D. 1 9 5 0 - Field observations on trees hosts of
1 96 8 a - The Genus Aerangis. - Ibid. 37: 490-4 93. orchids in the Philippines. - Philip. Orch. Rev. 3 : 3 - 8 .
1 96 8 b - Distribution of epiphytic orchids in semi Summerhayes, V.S. 1 96 8 - Orchidaceae, Part 1 . - In
deciduous tropical forest in Southern Nigeria. - J. Milne-Redhead, E. & Polhill, R.M. (ed.) 1 9 68.
Ecol. 5 6 : 6 9 7- 705 . Swanson, R.H. & Lee, R. 1 96 6 - Measurement of water
1 96 9 - The distribution of epiphytic orchids in Nigeria movement from and through shrubs and trees. - J.
in relation to each other and to geographic location Forest. 64 : 1 8 7- 1 90.
and climate, type of vegetation and tree species. - Teuscher, H. 1 9 7 2 - Microcoelia guyoniana and other
Bioi. J. Linn. Soc. 1 : 24 7 - 2 8 5 . leafless epiphytic orchids. - Amer. Orch. Soc. Bull.
1 97 1 - The flowering time of West African orchids. - 4 1 :497-5 0 1 .
Bot. J. Linn. Soc. 64 : 1 6 3 - 1 8 1 . Thornton, I. 1 9 6 5 - Nutrient content o f rainwater i n the
1 9 7 2 - Epiphytic orchids as characterizers of vegeta Gambia. - Nature 205 : 1 0 2 5 .
tion in West Africa. - A paper presented at the World Thorold, C .A. 1 9 5 2 - The epiphytes of Theobroma cacao
Orchid Conference at Mendelin, Colombia 1 9 7 2. in Nigeria in relation to the incidence of Black Pod dis
Sanford, W.W. & Adanlawo, I. 1 9 7 3 - Velamen and ex eases (Phytophthora palmivora). - J. Ecol. 40 : 1 25 -
odermis characters of West African epiphytic orchids 1 42.
in relation to taxonomic grouping and habitat Tixier, P. 1 9 66 - Flore et vegetation orophiles de l'Asie
tolerance. - Bot. J. Linn. Soc. 66 : 307-3 2 1 . tropicale. - Soc. d 'edition d'enseignement superieur,
Scaetta, H . 1 93 7 - L a genese climatique des sots mon Paris.
tagnards de 1' Afrique Centrale. Les formations Tournay, R. 1 9 5 5 - Orchidaceae africanae. 1 . Les
vegetates qui en caracterisent les stades de degrada orchidees congolaises : caracteres vegetatifs et floraux,
tion. - Inst. Roy. Colonial Beige, Bruxelles. types biologiques, clef de determination des genres
Schelpe, E.A.C .L.E. 1 9 70 - Pteridophyta. - In Exell, sympodiaux, apen;u les genres. - Bull. Soc. Roy. Bot.
A.W. & Launert, E. (ed.) 1 9 70. Belg. 8 7 : 5 7 - 7 9.
Schimper, A.F.W. 1 88 8 - Die epiphytische Vegetation Tukey, H.G., Jr. 1 9 70 - Leaching of rnetablites from
Amerikas. - Bot. Mitt. Trop. 2. foliage and its implication. - In Odum, H.T. & Pigeon,
1 903 - Plant geography upon a physiological basis. R.F. (ed.) 1 9 70, Chapter H 1 1 .
Oxford. Vanderyst, R.P.H. 1 9 2 2 - Nouvelle contribution a !'etude
1 9 3 5 - Pflantzengeographie auf Physiologischen des Ficus epiphytes sur l'Elaeis. - Rev. Zool. Afric.
Grundlage. - Revised by Faber, A.F., Jena. 1 0( 2 ) .
Schnell, R. 1 9 5 0 - La foret dense. - Lechevalier, P aris Wallach, A. 1 9 3 9 - Beitrage zur Kenntnis der
VI. Wasseraufnahme durch die Luftwurzeln tropischer
1 9 5 2 - Vegetation et flore de la region montagneuse Orchideen. - Zeitschr. f. Bot. 3 3 : 443-468.
du Nimba. - Mem. I.F.A.N. 22, D akar. Waiter, H. 1 9 7 1 - Ecology of tropical and subtropical
1 9 70 - Phytogeographie des payes tropicaux. I -11. - vegetation. - Oliver & Boyd, Edinburgh.
Gauthier-Vitlars, Paris. Visser, S.A. 1 9 64 - Origin of nitrates in tropical
Sernander, R. 1 9 1 2 - Studier ofver lafvarnes biologi. I . rainwater. - Nature 20 1 : 3 5 .
Nitrofila lafvar. - Syensk Bot. Tidskr. 6 : 803 - 8 8 3 . Voorhoeve, A.G. 1 9 65 - Liberian high forest trees. -
Sharp, A.J. 1 95 7 - Vascular epiphytes in the Great Pudoc, W ageningen.
Smoky mountains. - Ecology 3 8 : 6 3 5 - 6 54. Voth, P.H. 1 9 3 9 - C onduction of rainfall by plant stem in
Sjogren, E. 1 9 6 1 - Epiphytische Moosvegetation in a tropical rain forest. - Bot. Gaz. 1 0 1 : 3 2 8-3 40.
Laubwaldern der Insel Oland (Schweden). - Acta Voute, A.D. 1 9 46 - Regulation of the density of the in
Phytogeogr. Suec. 44. Uppsala. sect populations in virgin-forests and cultivated woods.
Skye, E. 1 9 68 - Lichens and air pollution. - Ibid. 5 2. - Arch. Neerl. Zool. 7 :4 3 5 -4 70.
Smith, J.J. 1 9 2 5 � Ephemeral orchids. - Ann. Jard. Bot. 1 9 64 - Harmonious control of forest insects. - In
Buitenz. 3 5 : 5 5 -70. Romberger, J.A. & Mikola, P. Vol. I 1 9 6 4 : 3 25 -3 8 3 .
Spanner, L. 1 9 3 9 - Untersuchungen iiber den Warme Went, F.A.F.C. 1 8 9 5 - Ueber Haft- und Nahrwurzeln bei
und Wasserhaushalt von Myrmecodia und Hyd Kletterpflanzen und Epiphyten. - Ann. Jard. Bot.
nophytum. - Jahrb. Wiss. Bot. 8 8 : 24 2-283. Buitenz. 1 2 : 1 -7 2 .
Steenis, C .G.J.J., van 1 93 8 - Recent progress and Went, F.W. 1 9 3 1 - Over de sociologie der Epiphyten.
prospects in the study of the M alaysian flora. - Handel . 6 Ned. Ind. Naturw. Congr. : 3 8 1 -3 8 7. Ban
Chron. Bot. 4 : 3 92-3 9 7. doeng.
Stewart, J. 1 9 73 - More about Microcoelias. - Amer. 1 9 40 - Soziologie der Epiphyten eines tropischen
Orch. Soc. Bull. 42 :80 1 -805. Urwaldes. - Ann. J ard. Bot. Buitenz. 5 0 : 1 -9 8 .
Whitmore, T.C. 1 9 6 8 - A study of light conditions in Koopowitz, H. 1 964 - Aerangis- an extrapolation from
forests in Ecuador with some suggestions for further ecology to cultivation. - Amer. Orch. Soc. Bull.
studies in tropical forests. - In Bainbridge et al. (ed.) 3 3 : 1 026- 1 02 8 .
1 96 8 : 2 3 5 - 24 7 . Kunkel, G. 1 9 63 - Die einfliisse von Kultur und Siedlung
Wiesner, J. 1 89 7 - Zur Physiologie von Taeniophyllum auf das Waldbild Liberias. - Zeitschrift fiir
Zollingeri. - S.B. Akad. Wiss. Wien, Math.-Naturw. Wirtschaftsgeogr. 7 :4 5 - 5 1 .
C l . 1 06 : 7 7- 9 8 . Leakey, C . L.A. 1 968 a - The orchids of Uganda. -
Willis, A.J. & Jetferies, R.L. ( 1 9 6 3) - Investigations of Amer. Orch. Soc. Bull. 3 7 : 8 8 3- 8 8 6.
the water relations of sand-dune plants under natural 1 96 8 b - The orchids of Uganda. The major
conditions. - In Rutter, A. & Whitehead, F. (ed.) angraecoid genera. - Ibid. 3 7 : 9 7 8 - 9 8 3 .
1 963. 1 9 6 8 c - The orchids of Uganda. Various species of
Willis, J.A., Yemm, E.W. & B alasubramaniam, S. 1 9 63 - horticultural merit. - Ibid. 3 7 : 1 05 2 - 1 0 5 5 .
Transpiration phenomena in detached leaves. - Morton, J.K. 1 9 6 1 - West african lilies and orchids. -
Nature 1 99 : 2 6 5 - 2 6 6 . Longmans, Green & Co ... London.
WiJlis, J. C . 1 96 6 - A dictionary of flowering plants & Onochie, C .F.A. 1 9 6 2 - The use of field characters in the
ferns. - Seventh ed., revised by Airy Shaw, H.K., tree identification. - C omptes Rendus, IV Reunion
C ambridge Univ. Press, London. pleniere A.E.T.F.A.T. 1 9 6 1 . Lisboa.
Withner, C . L. (ed.) 1 9 5 9 - The orchids, a scientific sur Piers, F. 1 96 8 - Orchids of Africa. Angraecum distichum
vey. - Ronald Press, New York. and A. aporoides. - Amer. Orch. Soc. Bull. 3 7 :9 9 8-
- 1 9 5 9 - Orchid physiology. - In Withner (ed.) 999.
1 9 5 9 : 3 1 5 - 3 60. Richards, P.W. 1 9 5 7 - Ecological notes on West African
Yarranton, G.A. 1 96 7 - An instrument for measuring the vegetation. - J. Ecol. 45 : 5 6 3 - 5 7 7 .
microrelief of the bark. - C an. J. Bot. 4 5 (8) : 1 1 73 - 1 9 6 3 - Ecological notes on West African vegetation.
1 1 78. Ill. The upland forest of Cameroon Mountain. - J.
Ecol. 5 1 : 5 29-544.
Sanford, W.W. 1 9 70 a - Plectrelminthus caudatus. -
Literature not cited in the text but related to the sub Amer. Orch. Soc. Bull. 3 9 : 224-226.
1 9 70 b - Practical conservation of orchids in Nigeria.
ject
- Nig. J. Se. 4 :49-5 7 .
Briscoe, M.S. 1 95 2 - The relation of insects and insect 1 9 70 c - Orchids of the Bamboutos and Manengouba
borne diseases to the vegetation and environment in highlands in East Cameroun. - Amer. Orch. Soc. Bull.
Liberia. - Ecology 3 3 : 1 8 7-2 1 4. 3 9 : 6 8 1 -6 8 5 .
Bryant, M.T. 1 9 66 Some of our palms. - The Liberian
-
1 9 70 d - A selection o f West African orchids. - Ibid.
naturalist 4( I ) : 9 - 1 3 . Monrovia. (Stencil). 3 9 : 8 90-8 9 5 .
- 1 96 6 Trees
- roadsidP.s . - Ibid. 1971 - The orchid flora o f Equatorial Guinea i n rela
4(2) :9-20.
tion to that of West Africa. - Mitt. Bot. 1 0 : 2 7 8-298.
Hopkins, B. 1 96 5 Forest and savanna. - Heinemann,
-
Schelpe, E.A.C .L.E. - 1 9 66 - An introduction of the
London.
South African orchids. - MacDonald, London.
Harrison, E.R. 1 9 72 - Epiphytic orchids of Southern Strong, R.H. (ed.) 1 9 30 - The African Republic of
Africa. - The Natal branch of the Wildlife Preserv. Liberia and the Belgian Congo. I, 11. - C ambridge,
and Conserv. Soc. of South Africa. Durban. Mass.
Jaeger, P. & Summerhayes, V.S. 1 94 9 - Note sur Warner, R. 1 94 5 - Vegetation and agriculture of Liberia
quelques Orchidees recoltees dans les Monts Loma and adjacent West Africa. - Iowa A cad. Sci. 5 2 : 1 7 1 -
(Sierra Leone) et les contrees limitrophes. - Kew. Bull. 1 89 .
3 :4 7 5 - 4 8 3 .
Kloppers, J. 1 9 7 1 - Die Suid-afrikaanse epifiete. - S.
African Orch. J. March 1 9 7 1 : 20-23.
1. E. A lmquist, Upplands vegetation och flora. (Vegeta mossflora i Uppsalatrakten. (Summary : Moss flora
tion and flora of Uppland.) 1 9 29. 4 0 : - ( 2 7 :-). I S B N and moss vegetation in the neighbourhood of Uppsa
9 1 - 7 2 1 0-00 1 -X . la.) 1 9 4 5 . 2 7 : - ( 1 8 :-). I SB N 9 1 - 72 1 0-0 1 9-2.
2. S . Thunmark, D e r See Fiolen und seine Vegetation. 20. N. A lbertson, Gsterplana hed. Ett alvaromrade pa
1 9 3 1 . 2 1 :- ( 1 4 :-). I S B N 9 1 -7 2 1 0-002-8. Ki nnekulle. (Zusammenf. : G sterplana hed. Ein Alvar
3 . G.E. Du R ietz, Life-forms of terrestrial flowering gebiet au f dem Kinnekulle.) 1 94 6 . 2 7 :- ( 1 8 : - ). I S B N
plants. I. i 9 3 1 . 1 5 :- ( I 0 :-) . I S B N 9 1 -7 2 1 0-003-6. 9 1 - 72 1 0-020-6.
4 . B. Lindquist, Om den vildvaxande skogs almens raser 2 1 . H. Sjdrs, Myrvegetation i B ergslagen. (Summary :
och deras utbredning i Nordvasteuropa. (Summary : Mire vegetation in Bergslagen, Sweden.) 1 94 8 . 3 6 :
The races of spontaneous Ulmus glabra Huds. and (24 :-). I S B N 9 1 - 72 1 0-02 1 -4 .
their d istribution in N W . Europe.) 1 9 3 2. 1 2 :- ( 8 : -). 22. S . A hlner, Utbredningstyper bland nordiska barr
ISBN 9 1 - 7 2 1 0-004-4. trad slavar. (Zusammenf. : Verbreitungstypen unter
5. H. Osvald, Vegetation of the Pacific coast bogs of fennos kandi schen N adelbaumflechten.) 1 94 8 . 3 0 :
N orth America. 1 9 3 3 . 1 2 :- ( 8 :-). I S B N 9 1 - 7 2 1 0- (20: -). I S B N 9 1 - 72 1 0-022-2.
005 -2. 23. E. Julin, Vessers· udde. M ark och vegetation i en igen
6. G. Samuelsson, Die Verbreitung der hoheren Was ser vaxande lovang vid Bj arka-Saby. (Zusammenf. : Ves
pflanzen in N ordeuropa. 1 9 3 4 . 2 1 :- ( 1 4 :-). I S B N 9 1 - sers udde. Boden und Vegetation in einer verwachsen
72 1 0-006-0. den Laubwiese bei Bj arka-S aby in Gstergotland, Siid
7 . G. Degelius, Das ozeanische Element der Strauch schweden.) 1 94 8 . 2 1 :- ( 1 4 :-). I S B N 9 1 - 7 2 1 0-023 -0.
und Laubflechtenflora von Skandinavien. 1 9 3 5 . 3 6 : 24. M. Fries, Den nordi ska utbredningen av Lactuc a
(24 :-). I S B N 9 1 - 7 2 1 0-007-9. alpina, A conitum septentrionale, R anunculus platani
8 . R . Serna nder, Granskar och F i b y urskog. En studie folius och Polygonatum verticillatum. (Z usammenf. :
over stormluckornas och marbuskarnas betydelse i Die nordis che Verbreitung von Lactuca alpina. . . )
den svenska granskogens regeneration. (Summary : 1 94 9 . 1 5 : - ( 1 0 : -) . I S B N 9 1 - 7 2 1 0-024-9.
The primitive forests of Granskar and Fiby. A study 25. 0. Gjcerevoll, Sn0leievegetasjonen i Oviksfjellene.
of the part played by storm-gaps and dwarf trees in the ,(Summary : The snow-bed vegetation of Mts Oviks
regeneration of the Swedish spruce forest.) 1 9 3 6 . 2 1 : f] allen, J amtland, Sweden.) 1 9 49. 1 8 : - ( 1 2 :-). I S B N
( 1 4 :-). I S B N 9 1 - 7 2 1 0-008- 7 . 9 1 - 7 2 1 0-02 5 - 7 .
9 . R . Sterner, Flora der I n sel Gland. D i e Areale d e r Ge 26. H . Osvald, Notes on the vegetation o f British and Irish
fasspflanzen Glands nebst Bemerkungen zu ihrer mosses. 1 94 9 . 1 2 :- ( 8 :-). ISBN 9 1 - 7 2 1 0-026-5 .
Oekologie und Soziologie. 1 9 3 8 . 2 1 :- ( 1 4 :-). I S B N 27. S. Selander, Floristic phytogeography of South
9 1 - 72 1 0-009- 5 . Western Lule Lappmark (Swedish Lapland) I. 1 9 50.
1 0. B . L indquist, D al by Soderskog. E n sklmsk lovskog i 2 1 : - ( 1 4 :-). ISBN 9 1 - 7 2 1 0-02 7 - 3 .
forntid och nutid. (Zusammenf. : Ein Laubwald in 28. S . Selander, Floristic phytogeography o f South
S chonen in der Vergangenheit und Gegenwart.) 1 9 3 8 . Western Lule Lappmark (Swedish Lapland) 11. Karl
2 7 :- ( 1 8 :-). I S B N 9 1 - 7 2 1 0-0 1 0-9. vaxtfloran i sydvastra Lule Lappmark. (Summary :
1 1 . N. Stalberg, Lake Vattern. Outlines of its natural Vascular flora.) 1 9 05. 1 8 : - ( 1 2 :-). I S B N 9 1 - 7 2 1 0-
history, especially its vegetation. 1 9 3 9 . 1 2 :- ( 8 :-). 028- 1 .
I S B N 9 1 - 7 2 1 0-0 1 1 - 7. 29. M. Fries, Pollen analytiska vittnesbord o m senkvartar
1 2 . G.E. Du Rietz, A . G. Hannerz, G. Lohammar, R. vegetationsutveckling, sarskilt skogshi storia, i nord
. Santesson & M. Wcern, Zur Kenntnis der Vegetation vastra Gotal and. (Z usamm enf. : Pollenanalytische
des Sees Takern. 1 9 3 9 . 1 2 :- ( 8 :-). I S B N 9 1 - 7 2 1 0- Zeugnisse der spatquartaren Vegetationsentwicklung,
0 1 2-5 .. . hauptsachlich der Waldgeschichte, im nordwestlichen
1 3 . Viixtgeografiska studier tilliignade Car! Skoltsberg Gotaland, Sii d schweden.) 1 9 5 1 . 2 1 :- ( 1 4 :-). I S B N
pa sextioarsdagen 1 / 1 2 1 940. ( Geobotanical studies 9 1 - 7 2 1 0-02 9-X.
dedicated to C. Skottsberg.) 1 9 40. 3 6 : - (24 :-). I S B N 30. M. Wcern, Rocky-shore algae in the Gregrund Archi
9 1 - 7 2 1 0-0 1 3 - 3 . pelago. 1 9 5 2 . 3 3 : - (22 :-). I S B N 9 1 - 72 1 0-030- 3 .
1 4. N . Hylander, De svenska formerna a v Mentha gentilis 3 1. 0. R une, Plant life on serpentines a n d rel �ted rocks in
L. coli. (Zusammenf. : D ie schwedischen Formen der the North of Sweden . 1 9 5 3 . 1 8 :- ( 1 2 :-). I S B N 9 1 -
Mentha gentilis L . sensu cot!.) 1 9 4 1 . 1 2 : - ( 8 :-). I SB N 7 2 I 0-0 3 1 - I .
9 1 - 7 2 1 0-0 1 4- 1 . 3 2. P . Kaaret, Was servegetation d e r Seen Orlangen und
1 5 . T. E. Hasselrot, Till kannedomen om nagra nordiska Trehorni ngen . 1 9 5 3 . 1 2 :- ( 8 :-). I S B N 9 1 - 7 2 1 0-03 2-
umbilicariaceers utbredning. (Zusammenf. : Zur X.
Kenntnis der Verbreitung einiger U mbilicariaceen in 3 3 . T. E. Hasselrot, Nordliga lavar i Syd- och Mellan
Fennoscandia.) 1 9 4 1 . 1 2 :- ( 8 :-). I S B N 9 1 - 72 1 0- sverige. (Nordliche Flechten in Sii d - und M ittelschwe
0 1 5-X. den . ) 1 9 5 3 . 2 2 :- ( 1 4 : -). I S B N 9 1 - 7 2 1 0-03 3-8.
· 1 6. G . Samuelsson, D i e Verbreitung d e r Alchemilla-Arten 34. H. Sjdrs, Slatterangar i Grangiirde fi nnmark. (Sum
aus der Vulgaris-Gruppe in N ordeuropa. 1 9 4 3 . 1 8 : mary : Meadows in Grangarde Finnmark, S W . D alar
( 1 2 :-). ISBN 9 1 - 7 2 1 0-0 1 6-8. n a, Sweden . ) 1 9 5 4 . 1 8 :- ( 1 2 : -). I S B N 9 1 - 7 2 1 0-034-
I 7 . Th. A rwidsson, Studien iiber d i e Gefasspflanzen in 6.
den Hochgebirgen der Pite Lappmark. 1 9 4 3 . 2 7 : 35. S. Ki/ander, Karlvaxternas ovre gran ser pa fj all i syd
( 1 8 :-). I S B N 9 1 - 7 2 1 0-0 1 7-6. vastra J amtl and samt an grans an de delar av H arjeda
1 8 . N. Dahlbeck, Strandwiesen am siidostlichen Gre len och Norge. [ S u m m ary : Upper limits of vascular
sund. (Summary : Salt marshes on the S . E. coast of plants on mountains in Southwestern J a mtland and
Gresund.) 1 9 4 5 . 1 8 : - ( 1 2 :-). I S B N 9 1 - 7 2 1 0-0 1 8-4 . . adja cent parts of Harjed alen (Sweden) and N orway . ]
1 9 . E . von Krusenstjerna, Bladmossvegetation och blad- 1 9 5 5 . 2 2 : - ( 1 4 :-). I S B N 9 1 - 7 2 1 0-03 5 -4.
Distributor: Svenska Viixtgeografis k a Siillskapet, Box 5 5 9, S- 7 5 1 22 Uppsala, S weden
36. N. Quennerstedt, D i atomeern a i Limgans sjovegeta 48. E. Sjogren, Epilithische und eJ?igiiische Moosvegetati
tion. ( S u m m ary : Diatoms i n the lake vegetation of the on in Laubwiildern der Insel Oland, Schweden. (Sum
Langan drainage area, J iimtland, Sweden.) 1 9 5 5 . 2 2 :- · m ary : Epilithic and epigeic m o s s vegetation i n
( 1 4 :-). I S B N 9 1 - 7 2 1 0-03 6-2. deciduo us woods on t h e island of Gland, Sweden.)
3 7 . M.-B. Florin, Plankton of fresh and brackish waters in 1 9 64. 40:- (27 :-). I S B N 9 1 - 7 2 1 0-048-6 ( I S B N 9 1 -
the Sodertiilje area. 1 9 5 7 . 20:- ( 1 4 :-). ISBN 9 1 - 72 1 0-448- 1 ).
7 2 1 0-03 7-0: 49. 0. Hedberg, Features of afroalpine plant ecology.
3 8 . M.-B. Flo rin, l n sjostudier i Mellansverige. M ik ro Resume franc;ais. 1 9 64. 3 6 :- (24 : - ). I S B N 9 1 -7 2 1 0-
vegetation och pollenregn i vikar av Gstersjobiickenet 04 9 -4 (IS B N 9 1 - 7 2. 1 0-449-X).
och insjoar fran preboreal tid till nutid. (Summ ary : 50. The Pla n t Cover of S weden. A study dedicated to G.
Lake studies in Central Sweden. Microvegetation and Einar D u Rietz on his 70th birthday by his pupils.
pollen rain in inlets of the B altic basin and in lakes 1 9 6 5 . 7 2 : - (4 8 :- ). I S B N 9 1 - 7 2 1 0-050-8.
from Preboreal time to the present d ay.) 1 9 5 7 . 1 0 : 5 I. T. Flensburg, Desmids and other benthic algae of L ake
(6 :-). I S B N 9 1 - 7 2 1 0-03 8-9. Kiivsjon and Store Mo sse, SW S weden. 1 9 6 7 . 3 9 :
3 9 . M. Fries, Vegetationsutveckling och odlingshistoria i (26 :-). I S B N 9 1 - 7 2 1 0-05 1 -6 ( I S B N 9 1 - 7 2 1 0-45 1 - 1 ).
V arnhem strakten. En pollenan alytisk undersokning i
5 2 . E. Skye, Lichens and air pollution. A study of crypto
gamic epiphytes and environment i n the Stockholm
V iistergotl and. [Z usammenf. : V egetationsentwick
lung und Siedlungsgeschichte im Gebiet von V arnhem . region. 1 9 68. 4 2 : - ( 2 8 : -). I S B N 9 1 - 72 1 0-05 2-4
Eine pollen analytische Untersuchung aus V ii stergot ( I S B N 9 1 - 72 1 0-45 2-X).
land (Slidschweden).] 1 9 5 8 . 1 6 :- ( 1 0 : -). I S B N 9 1 - 5 3 . Jim Lundqvist, Plant cover and environment of steep
7 2 1 0-03 9 - 7 . hill- sides in Pite Lappm ark. (Resume : La couverture
4 0 . Bengt Pettersson, Dynamik o c h konstans i Gotl ands vegeta!e et !'habitat des fl ancs escarpes des collines de
flora och vegetation. (Resume : Dynamik und Kon Pite Lappm ark .) 1 9 6 8 . 4 8 :- ( 3 2 : -). I S B N 9 1 - 72 1 0-
stanz in der Flora und Vegetation von Gotland, 0 5 3 - 2 ( I S B N 9 1 - 7 2 1 0-45 3 -8).
S chweden .) 1 9 5 8 . 40:- ( 2 7 :-). I S B N 9 1 - 7 2 1 0-040-0. 5 4. Conservation of Vegetatioil in Africa South of the
4 1 . E. Uggla, Skogsbrandfiilt i M uddus n ationalpark. Sahara. Proc. of sixth plen. meeting of AETF AT. Ed.
( S u m m ary : Forest fire areas i n M uddus N ation al by Inga and Olov Hedberg. 1 96 8 . 60:- ( 40:-) , cloth
Park, Northern Sweden.) 1 9 5 8 . 2 1 :- ( 1 4 :-). I S B N 7 0 : - (47 :-). I S B N 9 1 - 7 2 1 0-05 4-0 ( I S B N 9 1 - 7 2 1 0-
9 1 - 72 1 0-04 1 - 9. 454-6).
42. K. Thomasson, N ahuel H u api. Plankton of some lakes 55. L . -K. Ko1�(gsson, The Holocene hi story of the Great
in an Argentine N ational P ark, with notes on terrestri Alvar of Ol and. 1 9 6 8 . 5 4 :- ( 3 6 :-). I S B N 9 1 - 7 2 1 0-
al vegetation. 1 9 5 9 . 2 1 :- ( 1 4 : -). I SB N 9 1 - 7 2 1 0-042- 05 5 - 9 ( I S B N 9 1 - 7 2 1 0-45 5 -4).
7. 56. H.P. Hallberg, Vegetation auf den Schalenablage
rungen in Bohusliin, Schweden . ( S u m m ary : Vegetati
4 3 . V. Gillner, Vegetations- und Standortsuntersuchungen
on on shell deposits in B ohusliin, Sweden.) 1 9 7 1 . 4 8 :
in den Strandwiesen der schwedischen Westkliste.
(3 2 : - ). I S B N 9 1 - 7 2 1 0-05 6 - 7 (ISBN 9 1 - 7 2 1 0-45 6-2).
1 9 60. 3 3 :- (22 :-). I S B N 9 1 - 7 2 1 0-04 3 - 5 .
5 7 . S. Fransson, Myrvegetation i sydviistra V iir mland.
4 4 . E . Sjogren, Epiphytische Moosvegetation in Laub
(Sum m ary : Mire vegetation i n south-western V iirm
wiildern der I n sel Gl and, Schweden. (Summ ary :
land, Sweden .) 1 9 72. 4 2 :- ( 2 8 :-). ISBN 9 1 - 7 2 1 0-
Epiphytic moss communities in deciduous woods on
05 7-5 ( I S B N 9 1 - 72 1 0-45 7-0).
the i sland of Gland, S weden. ) 1 9 6 1 . 24 : - ( 1 6 : -).
I S B N 9 1 - 72 1 0-044-3 (ISB N 9 1 - 7 2 1 0-444-9). 58. G. Wa!lin, Lovskogsvegetation i Sjuhiiradsbygden.
45. G. Wistrand, Studier i Pite Lappmarks kiirlviixtflora, ( S u m m ary : Deciduous woodlands in Sjuhiiradsbyg
den, V iistergotland, south-western Sweden.) 1 9 7 3 .
med siirskild hiinsyn till skogsl andet och de isolerade
42 :--(2 8 : -) I S B N 9 1 - 72 1 0-05 8 - 3 ( I S B N 9 1 - 7 2 1 0-.
fj iillen. (Z usammenf. : Studien U ber die Gefiisspflan
zenflora der Pite Lappmark mit besonderer Berlick 45 8-9).
sichti gung d e s Waldl andes u n d der i solierten niederen 5 9. D . Johansson, Ecology of vascular epiphytes in West
Fjel de.) 1 9 6 2 . 3 6 : - (24 :- ). I S B N 9 1 - 7 2 1 0-04 5 - 1 African rain forest. (Resume : Ecologie des epiphytes
( I S B N 9 1 - 72 1 0-4 4 5 - 7). vascul aires dans la foret dense h um ide d'Afrique oc
46. R. Ivarsson, Lovvegetationen i Mollosunds sock en. cidentale.) 1 9 74. 42 :-. I S B N 9 1 - 7 2 1 0-05 9 - 1 ( IS B N
(Zusammenf. : Die Laubvegetation i m K irch spiel 9 1 - 7 2 1 0-4 5 9-7).
Mollosund, Bo husliin, Schweden.) 1 9 6 2. 3 0 : - (20:- ). Limited n umbers of cloth-bound copies of Acta 44, 45,
I S B N 9 1 - 7 2 1 0-046-X (ISBN 9 1 - 72 1 0-446-5).
46 , 4 8 , 49, 5 1 , 52, 5 3 , 5 5 , 5 6 , 5 7, 58, 59 are available
4 7. K. Thomasson, Araucanian Lakes. Plankton studtes in
through the Society at an additional cost of 1 0 :- per
North Patagonia, with notes on terrestrial vegetation.
1 9 6 3 . 3 6 : - ( 24 : - ) . I S B N 9 1 - 7 2 1 0-04 7-8. copy. I S B N nos. in brackets refer to cloth-bound copies.
V A XT E K O LO G I S K A STUDI ER
Sallskapet bar till andamru att vacka och underhrula The object of the Society is to promote investigation in
intresse for vaxtgeografien i vidstracktaste mening, att flora and vegetation, . their history and their ecological
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ISBN 9 1 -7210-059- 1