Vegetatio 75: 81-86, 1988

Kluwer Academic Publishers, Dordrecht - Printed in the Netherlands 81

On the definition of ecological species groups in tropical rain forests
M. D. Swai ne a & T. C. Whi t mor e 2
IDepartment of Plant Science, The University, Aberdeen AB9 2UD, Scotland
ZOxford Forestry Institute, Department of Plant Sciences, South Parks Road, Oxford, 0)(1 3RB, England
Accepted 10.3.1988
Keywords: Ca nopy gap, Cl i max species, Ger mi nat i on, Pi oneer species, Seedling est abl i shment , Species guild
Abs t r a c t
The species ri chness of t r opi cal rai n forest s creat es di ffi cul t i es for ecol ogi cal anal ysi s. I t may useful l y be si mpl i -
fi ed by defi ni ng ecol ogi cal species gr oups whose member s share charact eri st i cs of i mpor t ance for det er mi ni ng
forest st r uct ur e and composi t i on. Many such cl assi fi cat i ons have been publ i shed, but few are pr oper l y ex-
pl ai ned. The t er mi nol ogy is conf used f r om l ack of precise defi ni t i ons. We pr opos e a si mpl e di vi si on of t ree
species i nt o t wo gr oups or guilds, pi oneer and non- pi oneer (or cl i max), based on seed ger mi nat i on and seedling
est abl i shment . Wi t hi n each guild t here is cont i nuous var i at i on and we r ecommend ar bi t r ar y subdi vi si on by
hei ght at mat ur i t y. We believe t hi s cl assi fi cat i on t o be appl i cabl e in all t r opi cal rai n forests.
Abbreviations: TRF = t r opi cal rai n forest(s); L HW = light har dwood species; HHW = heavy har dwood
species
Nomenclature." ei t her follows Hal l & Swai ne (1981), Whi t mor e (1972, 1973) and Ng (1978) or aut hor i t i es are cited.
I n t r o d u c t i o n
Ther e is now a general l y accept ed model of t r opi cal
rai n forest ( TRF) dynami cs. Gaps devel op in a cl osed
forest, new trees grow up in t hem, and a mat ur e
canopy is event ual l y at t ai ned. We may recogni se a
forest gr owt h cycle of gap, bui l di ng and mat ur e
phases (Watt 1947; Cousens 1974; Whi t mor e 1975,
1982). Gaps are a cruci al phase because what grows
up det er mi nes t he fl ori st i c compos i t i on of t he whol e
cycle ( Ha r t s hor n 1978; Ol de ma n 1974, 1978; Whi t -
mor e 1978). The mos t act i ve ar ea of TRF research
over t he past decade has been on ' gap- phas e repl ace-
ment ' t o el uci dat e processes of est abl i shment and
growt h, see for exampl e recent reviews by Br okaw
(1985) and Densl ow (1987). Ga p size is of f unda me n-
tal i mpor t ance. The bi gger t he gap t he great er t he so-
l ar r adi at i on at t he forest f l oor and t he great er t he
changes in ot her facet s of mi cr ocl i mat e above and
bel ow gr ound f r om condi t i ons beneat h cl osed cano-
py. Tree species di f f er in t hei r r esponse t o gaps. At
one extreme, s ome species can ger mi nat e bel ow a
canopy and t hei r seedlings est abl i sh and grow.
Ot her s need some i ncrease in sol ar r adi at i on f or
growt h. All t hese species have t he abi l i t y t o regener-
at e in situ bel ow a canopy. At t he ot her ext reme is
a gr oup or guild of species whose seedlings are not
f ound bel ow a canopy but whi ch a ppe a r af t er gap
creat i on. They do not regenerat e in situ. Thi s l ast
gr oup of species are of t en cal l ed ' pi oneer s' or ' secon-
dar y' species, t he ot hers ' cl i max' or ' pr i ma r y' spe-
cies. I t has l ong been known t o t emper at e zone for-
esters t hat t ree species di f f er in t he shade t ol er ance
of t hei r seedlings. I n Nor t h Amer i ca, for exampl e,
82
forest ers recogni se up t o five ' (shade) t ol er ance class-
es' (Baker 1950; Gr a ha m 1954). The st rong recent in-
terest in TRF dynami cs has led t o a si mi l ar recogni -
t i on and a very conf used nomencl at ur e has
devel oped wi t h a l ack of precise defi ni t i ons. No one
doubt s t he phe nome non, onl y how to name and clas-
si fy it. We believe t hat f ur t her progress in t he anal ysi s
and under st andi ng of TRF dynami cs, and especi al l y
t he cur r ent l y devel opi ng ar ea of ecophysi ol ogy (e.g.
Bazzaz & Pi cket t 1980) is hamper ed by this conf u-
sion.
Ecological species groups
The pur pos e of cl assi fi cat i on of species i nt o relative-
ly homogeneous gr oups is t o hel p peopl e c ommuni -
cat e and make general i zat i ons. I n t he st udy of TRF
such a cl assi fi cat i on is par t i cul ar l y necessary be-
cause many species are represent ed by onl y a few in-
di vi dual s in any st udy area. To cl assi fy species i nt o
gr oups or guilds i mposes a degree of si mpl i f i cat i on
whi ch reduces i nf or mat i on cont ent , but reveals
general pat t er ns and facilitates predi ct i ons about
forest processes. Taxonomi c gr oups by genus or fa-
mi l y are not sui t abl e because of t he poor cor r el at i on
in pl ant s bet ween t a xonomy and ecology.
The two main species groups
We believe t hat t here are t wo qual i t at i vel y di st i nct
gr oups of t ree species in TRF wi t h a si mpl e obser va-
ble di fference bet ween t hem. We call t hese pioneer
and climax or non-pioneer species respectively.
Pi oneer s are species whose seeds can onl y ger-
mi nat e in gaps in t he forest canopy open to t he sky
and in whi ch full sunl i ght i mpi nges at gr ound level
for at least par t of t he day.
Non- pi oneer or cl i max species are t hose whose
seeds can ger mi nat e under forest shade (very rarel y
in full sun as well). The seedlings can est abl i sh in for-
est shade and survive t here ( t hough in a few species
not for long). Young pl ant s are t hus c ommonl y
f ound bel ow a canopy, but may al so be seen in open
envi r onment s.
The t ri gger for ger mi nat i on for all TRF pi oneers
so far st udi ed is ei t her t he change in light qual i t y (an
i ncrease in red light fol l owi ng canopy removal ), or
t he st rongl y f l uct uat i ng t emper at ur e of soil exposed
for par t of t he day to full sunlight. Ot her t ri ggers
f ound in t he t emper at e zone, e.g. a fl ush of nut r i ent
ni t rogen, f l uct uat i ng soi l - moi st ur e cont ent , or a
wavel engt h - i ndependent i ncrease in r adi at i on (the
hi gh i r r adi ance react i on), have not yet been discov-
ered in TRE The i mpor t ant poi nt is t hat t hese are
all t ri ggers pr ovi ded by gap cr eat i on (see reviews by
Whi t mor e 1983; Vazquez-Yanes & Or ozco- Segovi a
1984). All pi oneers al so require full sunl i ght for seed-
ling est abl i shment and growt h. As a consequence
seedlings and young pl ant s are f ound in openi ngs in
t he forest (tree-fall gaps, roadsi des, landslips, felled
areas, etc.) and are never f ound under a closed forest
canopy, i ncl udi ng t hei r own.
These t wo gr oups have been wi del y recogni sed,
t hough rarel y preci sel y defi ned. Numer ous pai rs of
names have been appl i ed. Table 1 sets these out and
shows our obj ect i ons to mos t of t hem. We hope our
pr ef er r ed t er ms will now be adopt ed but realise this
may not happen. Mor e i mpor t ant t han t he names are
t he defi ni t i ons of t he groups. Loosel y appl i ed t ermi -
nol ogy has hel ped cause t he present conf usi on. All
t oo oft en, publ i shed wor k does not of f er cl ear defi -
ni t i ons, but it is evi dent t hat species are somet i mes
classed as pi oneers si mpl y because t hey are com-
monl y f ound in regrowt h af t er a ma j or di st urbance,
e.g. on a ba ndone d farms. Thi s is an i nadequat e test
as ma ny non- pi oneer s of t en col oni ze cleared l and
al ong wi t h t rue pi oneers by any def i ni t i on (e.g.
Swai ne & Hal l 1983) or may regrow f r om st umps or
root suckers.
Pi oneer species possess a whol e s yndr ome of
char act er s in addi t i on t o t he vital ones we have used
t o defi ne t hem. These are listed in Table 2. Collec-
tively t hese charact ers give a selective advant age f or
success in t he pi oneer ecol ogi cal niche. For example,
copi ous, wel l -di spersed seed i mproves t he chance of
reachi ng a new forest gap (van Steenis 1958). Rapi d
hei ght gr owt h enhanced t he chance to fill t he gap.
However not all pi oneers possess all these addi t i onal
charact ers.
83
Tabl e 1. Na me pai r s gi ven t o t he t wo ma j o r ecol ogi cal gr oups
of t ree speci es i n t r opi cal r ai n f or es t s . Pr e f e r r e d t e r ms ar e gi ven
in bol d t ype.
Pioneer/Non-pioneer: Al t h o u g h a wor d i n c o mmo n Engl i s h
usage, pi oneer is unl i kel y t o be mi s u n d e r s t o o d i n t hi s cont ext .
Co l o n i z i n g / Cl i ma x speci es: The us e o f ' cl i max' ma y be t aken t o
i mpl y accept ance o f Cl e me nt s i a n vi ews on s ucces s i on, a n d as s u-
mes a n accept abl e def i ni t i on o f cl i max veget at i on.
S e c o n d a r y / P r i ma r y speci es: Wi del y us ed, but easi l y c onf us e d
wi t h s e c onda r y a nd pr i ma r y s ucces s i on in veget at i on.
Sh a d e - b e a r e r s / Li g h t - d e ma n d e r s ( s hade t ol e r a nt / i nt ol e -
r ant ) : Appl i es t o seedl i ngs, not seeds, so t ha t not all ' i nt ol e r a nt s '
(l i ght de ma nde r s ) ar e st r i ct pi oneer s i n t he s ens e us e d her e.
No n - e q u i l i b r i u m/ Eq u i l i b r i u m speci es: Not mu c h us e d, a n d
r a t he r cr ypt i c; l i abl e t o c onf us i on wi t h appl i cat i on o f t hes e t e r ms
t o veget at i on.
r - s el ect ed/ K- s el ect ed speci es ( r - s t r at egi s t s / K- s t r at egi s t s ) : As s u -
mes a knowl e dge of how t he speci es evol ved. Of zool ogi cal or i gi n
( Ma c Ar t h u r & Wi l s on 1967).
We e ds / s pe c i e s o f cl osed veget at i on: We e ds ar e as s oci at ed wi t h
agr i cul t ur e a nd hor t i cul t ur e a nd t he t e r m is not mu c h us e d o f fo-
r est s.
Ep h e me r a l s / Pe r s i s t e n t s : Ephe me r a l is us ual l y appl i ed t o pl ant s
wi t h a l i f e- span of less t h a n one year ; l ar ge t ree pi oneer s coul d
r e a s ona bl y be cal l ed per s i s t ent .
No ma d / Dr y a d ( van St eeni s 1958): No ma d r ef er s t o c ont i nua l
mo v e me n t acr os s t he f or est at each gener at i on; dr ya ds ar e wood
n y mp h s , t r ue deni zens o f t he f or es t . Dr ya d ha s scar cel y been us e d
si nce i t s i nt r oduc t i on in t hi s sense.
Tabl e 2. Cha r a c t e r s y n d r o me o f pi oneer t ree speci es i n t r opi cal
r ai n f or es t . Not all pi oneer s pos s es s all t he char act er s iii-xvi.
i Seeds onl y ge r mi na t e i n c a nopy ga ps ope n t o t he s ky
a nd whi ch receive s o me full sunl i ght .
ii Pl a nt s c a nnot sur vi ve in s ha de - y o u n g pl ant s never
f o u n d u n d e r a cl osed f or es t canopy.
iii Seeds s mal l a n d pr oduc e d copi ous l y a n d mor e- or - l es s
c ont i nuous l y.
iv Seeds pr oduc e d f r o m ear l y in life.
v Seeds di s per s ed by a ni ma l s or wi nd.
vi Do r ma n t seeds us ual l y a b u n d a n t i n f or est soil (especi -
al l y f l es hl y- f r ui t ed speci es). Seeds o r t h o d o x ( no recal -
ci t r ant speci es known) (1).
vii Seedl i ng c a r bon- f i xa t i on r at e hi gh; c o mp e n s a t i o n
poi nt hi gh (2).
viii Hei ght gr owt h r api d.
ix Gr owt h i nde t e r mi na t e wi t h no r est i ng buds ( vi z syl l ep-
tic) (3).
x Br a nc hi ng rel at i vel y s par s e (4).
xi Leaves shor t - l i ved (2).
xii Root i ng super f i ci al (5).
xiii Wo o d us ual l y pal e, l ow dens i t y, not si l i ceous.
xi v Leaves suscept i bl e t o her bi vor y; s ome t i me s wi t h little
chemi cal def ence (6).
xv Wi de ecol ogi cal r a nge (7); phenot ypi cal l y pl ast i c (8).
xvi Of t e n shor t - l i ved.
(1) Te r mi nol ogy of Rober t s (1973), i ncl udes capaci t y f or
d o r ma n c y , Wh i t mo r e (1983).
(2) Ko y a ma (1978), Obe r ba ue r & St r ai n (1984).
(3) Booj h & Ra ma k r i s h n a n (1986).
(4) Wh i t n e y (1976).
(5) Shukl a & Ra ma k r i s h n a n (1986).
(6) Col ey (1983).
(7) Hal l & Swai ne (1980).
(8) Baker (1964).
Subdivision of the groups
Much confusion also arises from attempts at subdi-
vision of the two groups. We think that this is be-
cause within them variation is continuous. It is
quantitative rather than qualitative and there are not
sharp boundaries. Nevertheless, it is useful for many
purposes to recognise subgroups. These are accepta-
ble, so long as it is realised that they are arbitrary seg-
ments of a continuum.
We may plausibly advance many diverse factors as
criteria for the definition of subgroups, but we are
strongly constrained by what is known and practical.
Differences between species in their demands for
mineral nutrients, for water, or for photosynthetical-
ly active radiation may well be controlling influences
in forest ecology, but our knowledge is at present in-
adequate. Any classification must be based on
characters which we can reasonably expect to be
known for all species. We do not have a strong prefer-
ence for any particular scheme of subdivision. Some
measure of height achievable at maturity is one pos-
sibility (cf. Hall & Swaine 1981). For this the scheme
of Raunkiaer (1934) is suitable. In this scheme spe-
cies are allocated to nano-, micro-, meso- and mega-
phanerophyte classes, defined by the height above
84
the ground of their perennating buds, whi ch approx-
imates to their absol ute height at maturity. (The En-
glish words pygmy, small, medi um and large may
serve as alternatives). The cl assi fi cati on can be ex-
tended wi thout conf usi on to include shrubs, and
large and small herbs. Amongst pi oneer tree species,
longevity increases with stature, but we do not know
if large pioneers are generally shorter lived than large
non-pi oneers, so life span must onl y be used within
each group.
In Table 3 we present examples from all three TRF
regions of the pi oneer and cl i max species groups and
subgroups defined on height. The megaphanero-
phytes include 'emergents', whi ch is another term
difficult to define with precision. There is a similar
vagueness about terms such as 'canopy-top', 'mid-
Table 3. Examples of the pioneer and climax tree species groups
subdivided into height class subgroups.
canopy' , ' underst orey' which however are often
used, and for some purposes may be sufficiently pre-
cise. It may be noted t hat some species differ in-
fraspecifically at the subgroup level. For example
D&coglypremna coloneura, a West Afri can pioneer,
reaches only 20 m in Ghana but 50 m in Nigeria
(MDS pers. obs.).
Climax species response to solar radiation
Wi thi n the cl i max species groups there is variation
in the amount of solar radiation needed for seedling
growth ('release'). At one extreme there are species
whi ch need much solar radiation and then grow fast.
These tend to have seedlings with rapid mortal i ty be-
from Africa (AF), the Eastern Tropics (ET) and tropical America (AM)
Tree stature Pioneers (germinate in full
sun and require full sun for
survival and growth)
Climax (germinate in
shade, or rarely in full
sun, and seedlings can
survive and grow in shade)
Pygmy
(Nanophanerophytes)
<2 m tall
Small
(Microphanerophytes)
2- 7. 9 m tall
Medium
(Mesophanerophytes)
8- 29 m tall
Large
(Megaphanerophytes)
-> 30 m tall
probably none, class
occupied by shrubs eg
Solanum spp. (pantropical)
Rauvolfia vomitoria (AF)
most Trema (pantropical)
many Macaranga spp. (AF, ET)
Pipturus (ET)
Some Piper spp. (AM)
Musanga cecropioides (AF)
Anthocephalus (ET)
Macaranga hypoleuca (ET)
Cecropia spp. (AM)
most Sloanea spp. (AM)
Chlorophora excelsa (AF)
Terminalia ivorensis (AF)
Terminalia superba (AF)
Lophira alata (AF)
Pericopsis elata (AF)
Paraserianthes falcataria (ET)
Eucalyptus deglupta (ET)
Goupia glabra Aubl. (AM)
Laetia procera Eichl. (AM)
Cedrela odorata L. (AM)
Swietenia mahagoni (L.) Jacq. (AM)
Pycnocoma macrophylla (AF)
many arecoid palms
Coussarea spp. (AM)
Psychotria deflexa DC. (AM)
Microdesmis puberula (AF)
most Melastomataceae (ET, AM)
Drypetes ivorensis (AF)
Diospyros buxifolia (ET)
Turreanthus africanus (AF)
a few Dipterocarpaceae (ET)
Fagaceae (ET)
most Myristicaceae (ET, AM)
Minquartia guianensis Aubl. (AM)
Khaya ivorensis (AF)
Entandrophragma spp. (AF)
Funtumia elastica (AF)
Aningeria robusta (AF)
nearly all Dipterocarpaceae (ET)
Virola surinamensis Warb. (AM)
Pentaclethra macroloba (Willd.)
Kuntze (AM)
Couratari spp. (AM)
Vochysia maxima Ducke (AM)
Eschweilera spp. (AM)
low canopy shade, not persi st i ng f or long. I n Mal ay-
sia t hese are cal l ed Li ght Ha r dwoods ( LHW) and in-
cl ude t he Li ght Red Mer ant i Shorea spp.
( Di pt er ocar paceae) . Entandrophragma spp. (Mel i a-
ceae) are West Af r i can exampl es. These species are
i mpor t a nt t oday for t wo reasons. Silviculturally,
t hei r r egener at i on is f avour ed by t he massi ve canopy
di st ur bance caused by mode r n hi gh- vol ume t i mber
ext ract i on. Commer ci al l y, t hey have relatively pal e
and low densi t y t i mber. Apa r t f r om t he essent i al
char act er of ger mi nat i on and seedling est abl i shment
bel ow a l eafy canopy, L HW species resembl e pi-
oneers. At t he opposi t e extreme, vi a a c ont i nuum of
response, are ot her cl i max species whi ch requi re very
little or no i ncrease in sol ar r adi at i on f or release.
These grow mor e slowly. They have dar k, heavy,
of t en siliceous t i mber and in Mal aysi a are cal l ed
Heavy Ha r dwoods ( HHW) . They are less likely t o
regenerat e fol l owi ng c ont e mpor a r y hi gh-i nt ensi t y
l oggi ng. Thei r t i mber is less wi del y useful. Neo-
balanocarpus heimii, anot her di pt er ocar p, and
Cynometra alexandri ( Caesal pi ni aceae) are Mal ay-
si an and Af r i can exampl es.
Late secondary species
Some aut hor s have recogni sed a gr oup t hey call l at e
seral or l at e s econdar y species. These are i l l -defi ned
t er ms whi ch we believe shoul d be abandoned. Some-
t i mes t hey refer to species whi ch are domi nant in l at e
s econdar y forest, l at er t han pi oneers but bef or e cli-
max species become domi na nt (Budowski 1955). We
believe t hat in this usage l at e s econdar y species are
large pi oneers whi ch live l onger t han smal l ones (e.g.
Cedrela spp. and Swietenia spp. in t he neot ropi cs).
No one has demons t r at ed t he existence of a t hi rd ger-
mi nat i on/ es t abl i s hment ni che di fferent f r om t he pi-
oneer / cl i max niches as def i ned in this paper. We do
not believe t here is a class of species whi ch ger-
mi nat es and est abl i shes bel ow a pi oneer canopy but
not a cl i max one.
Anot he r usage of ' l at e seral ' is to refer t o a mor e
l i ght - demandi ng, fast er growi ng cl i max species, t he
L HW di scussed above. I n a forest whi ch is now ex-
peri enci ng smal l er gaps t han previously, HHW will
t end t o repl ace LHW. Thi s was t he si t uat i on for
85
whi ch Jones (1955/56) used t he t er m f or t he Okomu
forest, Ni geri a where Entandrophragma and ot her
Mel i aceae were regenerat i ng poor l y in situ, and at
Sungai Menyal a, Mal aysi a L HW megaphaner ophyt e
di pt er ocar ps have HHW di pt er ocar ps comi ng up
under neat h ( Whi t mor e 1975).
Conclusions
Uni versal adopt i on of t he def i ni t i ons we have made
here will be of great val ue in c ommuni c a t i on be-
t ween research workers in di fferent cont i nent s, and
shoul d hel p in compar at i ve studies of TRF dynami cs
and t he search f or general pri nci pl es. I f except i ons
exist, and we know of none, t hey will onl y be revealed
i f aut hor s (i ncl udi ng oursel ves) are qui t e explicit in
t he def i ni t i on o f t he t er ms t hey use.
Acknowledgements
Thi s paper has benef i t ed f r om t he cri t i ci sms of t hree
a nonymous referees.
References
Baker, E S. 1950. Pr i nci pl es o f si l vi cul t ure. McGr aw Hill, New
York.
Baker, H. B. 1964. Char act er i s t i cs a nd mo d e s o f or i gi n o f weeds.
In: Baker, H. G. & St ebbi ns, G. L. (eds), The genet i cs of
col oni zi ng speci es. Ac a de mi c Pr ess, London.
Bazzaz, F. A. & Pi cket t , S. T. A. 1980. Physi ol ogi cal ecol ogy o f
t r opi cal successi on: a c ompa r a t i ve review. An n . Rev. Ecol . Syst.
11: 2 8 7 - 3 1 0 .
Booj h, R. & Ra ma k r i s h n a n , P. S. 1982. Gr owt h st r at egy o f t rees
r el at ed t o s ucces s i onal s t at us I, II. For. Ecol . Manage. 4:
3 5 9 - 3 7 4 , 3 7 5 - 3 8 6 .
Brokaw, N. R. 1985. Treefalls, r egr owt h a n d c o mmu n i t y s t r uct ur e
i n t r opi cal f or est s. In: Pi cket t , S. T. A. & Whi t e, P. S. (eds), The
ecol ogy o f nat ur al di s t ur ba nc e a n d pat ch dynami cs . Academi c,
New York.
Budowski , G. 1955. Di s t r i but i on o f t r opi cal Ame r i c a n r ai n forest
speci es in t he l i ght of s ucces s i onal pr ocesses. Tur r i al ba 15:
40 - 42.
Coley, P. D. 1983. Her bi vor y a n d def ensi ve char act er i st i cs of tree
speci es in a l owl and t r opi cal forest . Ecol . Monogr . 53:
2O9- 233.
Cous e ns , J. 1974. An i nt r oduc t i on t o woodl a nd ecol ogy. Ol i ver
& Boyd, Edi nbur gh.
86
Densl ow, J. S. 1987. Tropi cal rai n f or est ga ps a nd t ree speci es
diversity. An n . Rev. Ecol . Syst. 18: 431- 451.
Gr a h a m, S. A. 1954. Scor i ng t ol er ance o f f or est trees. Mi c hi ga n
For est r y 4, Uni ver s i t y o f Mi chi gan, An n Ar bor .
Hal l , J. B. & Swai ne, M. D. 1980. Seed st ocks i n Gh a n a i a n forest
soils. Bi ot r opi ca 12: 2 5 6 - 2 6 3 .
Hal l , J. B. & Swai ne, M. D. 1981. Di s t r i but i on a n d ecol ogy o f vas-
cul ar pl ant s in a t r opi cal r ai n forest . J unk, The Hague.
Ha r t s h o r n , G. S. 1978. Tree fal l s a nd t r opi cal f or est dynami cs . In:
Toml i ns on, P. B. & Zi mme r ma n , M. H. (eds), Tropi cal t rees as
l i vi ng s ys t ems , Uni ver s i t y Press, Cambr i dge.
J ones , E. W. 1955- 1956. Ecol ogi cal st udi es on t he rai n forest of
s o u t h e r n Ni ger i a. IV. The pl at eau forest o f t he Ok o mu f or est
reserve. J. Ecol . 43: 5 6 4 - 5 9 4 ; 44: 83- 117.
Koyama, H. 1978. Phot os ynt he s i s st udi es i n Pa s oh f or est reserve.
Mal ay. Nat . J. 30: 2 5 3 - 2 7 8 .
Ma c Ar t hur , R. H. & Wi l s on, E. O. 1967. The t heor y of i sl and bi o-
geogr aphy. Pr i ncet on Uni ver s i t y Press, Pr i ncet on, New Jersey.
Ng, F. S. P. (ed.) 1978. Tree f l or a o f Mal aya, Vol. 3. Lo n g ma n , Kua-
la Lumpur .
Ober bauer , F. S. & St r ai n, B. R. 1984. Phot os ynt he s i s a nd succes-
si onal s t at us of Cos t a Ri can r ai n f or est trees. Pbot os yn. Res.
5: 2 2 7 - 2 3 2 .
Ol de ma n, R. A. A. 1974. Uar chi t ect ur e de la for~t guyanai s e.
Me moi r e s ORSTOM 73.
Ol de ma n, R. A. A. 1978. Ar chi t ect ur e a nd ener gy exchange of di-
c ot yl e donous t rees i n t he forest . In: Toml i ns on, P. B. & Zi m-
me r ma n , M. H. (eds), Tropi cal t rees as l i vi ng syst ems. Uni ver si -
t y Press, Cambr i dge.
Raunki aer , C. 1934. The l i f e- f or ms of pl ant s a n d st at i st i cal pl a nt
geogr aphy. Oxf or d Uni ver s i t y Press, London.
Rober t s, E. H. 1973. Pr edi ct i ng t he st or age life o f seeds. Seed
Sci ence a nd Technol ogy 1, 4 9 9 - 5 1 4 .
Shukl a, R. P. & Ra ma k r i s h n a n , P. S. 1986. Ar chi t ect ur e a nd
gr owt h st rat egi es o f t r opi cal t rees in r el at i on t o s ucces s i onal st a-
t us. J. Ecol. 74: 3 3 - 4 6 .
Swai ne, M. D. & Hal l , J. B. 1983. Ear l y s ucces s i on on cl eared for-
est l and i n Gh a n a . J. Ecol . 71: 6 0 1 - 6 2 7 .
Van St eeni s, C. G. G. J. 1958. Rej uvenat i on as a f act or for j udgi ng
t he s t at us o f veget at i on t ypes. The bi ol ogi cal n o ma d t heor y. In:
Pr oceedi ngs o f t he s y mp o s i u m on h u mi d t r opi cs veget at i on,
Kandy. UNESCO, Pari s.
Vazquez-Yanes, C. & Or ozco- Segovi a, A. 1984. Ecophys i ol ogy of
seed ge r mi na t i on. In: Medi na, E. e t a l . (eds), Physi ol ogi cal
ecol ogy of pl ant s o f t he wet t ropi cs. J unk, The Hague.
Wat t , A. S. 1947. Pat t er n a n d pr ocess in t he pl ant c ommuni t y. J.
Ecol . 35: 1 - 2 2 .
Whi t mor e , T. C. (ed.) 1972, 1973. Tree Fl or a of Mal aya, Vols 1,
2. Lo n g ma n , Kual a Lumpur .
Whi t mor e , T. C. 1975. Tropi cal rai n f or est s o f t he Far East .
Cl a r e ndon, Oxf or d.
Whi t mor e , T. C. 1978. Ga p s in t he f or est canopy. In: Toml i ns on,
P. B. & Zi mme r ma n , M. H. (eds), Tropi cal t rees as l i vi ng sys-
t ems. Uni ver s i t y Press, Cambr i dge.
Whi t mor e , T. C. 1982. On pat t er n a nd pr ocess in forest s. In: New-
ma n , E. I. (ed.), The pl ant c o mmu n i t y as a wor ki ng me c ha -
ni s m. Blackwell, Oxf or d.
Whi t mor e , T. C. 1983. Se c onda r y s ucces s i on f r om seed in t ropi cal
r ai n forest s. For. Abst r . 44: 7 6 7 - 7 7 9 .
Whi t ney, G. G. 1976. The bi f ur cat i on rat i o as a n i ndi cat or of
adapt i ve st r at egy in wo o d / p l a n t species. Bull. Torr. Bot . Cl ub
103: 6 7 - 7 2 .