TROPICS 3 (3/4): 249-276 Issued March, 1994

List of Plant Species Identified in the Northern Part of the Lope Reserve, Gabon*

Caroline E.G. TUTIN Centre International de Recherche Medicales de Franceville, Franceville, Gabon; Department of Biological and Molecular Sciences, University of Stirling, Scotland.

Lee J. T. WHITE NYZS-The Wildlife Conservation Society, U.S.A.; Institute of Cell,

Animal and Population Biology, University of Edinburgh, Scotland; Programme de Conservation et

Utilisation Rationelle des Ecosystemes Forestiers d'Afrique Centrale (ECOFAC), Composante Gabon (Projet FED, CCE DG VIII).

Elizabeth A. WILLIAMSON Psychology Department, University of Stirling, Scotland.

Michel FERNANDEZ Centre International de Recherche Medicales de Franceville, Franceville, Gabon; Department of Biological and Molecular Sciences, University of Stirling, Scotland;

Programme de Conservation et Utilisation Rationelle des Ecosystemes Forestiers d' Afrique Centrale

(ECOFAC), Composante Gabon (Projet FED, CCE DG VIII).

Gordon MCPHERSON Missouri Botanical Garden, St. Louis, Missouri, U.S.A.

ABSTRACT Research on lowland gorillas (Gorilla g. gorilla) and chimpanzees (Pan t. troglodytes)

began at the 'Station d'Etudes des Gorilles et Chimpanzes' in the Lope Reserve, central Gabon, in 1983

and is on-going. This paper lists 676 species of plants belonging to 91 families that occur in the 50 sq. km

study area. Data on trees with diameters of 10 cm or more were collected systematically along line

transects and opportunistic collections of fertile plants were made. For each plant species, the life-form,

habitat preference and density (for trees recorded on transects) are listed. For plants that provide food for

gorillas and chimpanzees, the part eaten is given. The plant species list is not complete but shows the flora of the SEGC study area to be diverse. The seven habitat types described range from Savanna to

Closed Canopy Forest but the study area is dominated by Marantaceae Forest. Gorillas and chimpanzees

at Lope have diverse diets and obtain food from plants in all of the habitat types. Some minor (in terms of

area) habitats provide large amounts of food in particular seasons. Comparison of ape diets in different

parts of Africa can only advance if vegetation inventories for each study site are compiled and published.

Key words: Gorilla g. gorilla / Pan t. troglodytes / plant species list / habitat-types / vegetation

description / conservation and reserve management

Gabon straddles the equator on the west coast of Africa and approximately 80% of the country's area of 267,000 km2 is covered by lowland tropical forest. Though still incompletely documented, the floristic diversity is great and Breteler (1990) estimated that at least 6,000 species of phanerogams occur. The patchy nature of botanical exploration within Gabon is highlighted by recent descriptions of new species (e.g. Halle, 1987; Halle & Louis, 1989; McPherson & Louis, 1991) and many new locality records (F. White, pers. comm.). The area of Gabon that has been most intensively studied is around Makokou, in the north-east of the

* Correspondence and Reprint Requests: Dr. C.E.G. Tutin, C.I.R.M.F., B.P. 769, Franceville, Gabon

C. E. G. Turnq, L. J. T. Wrure, E. A. Wu-unusoN, M. FEnNANDEZ & G. McRrnsoN

country. Six published lists cover a total of 1,233 plant species (Hall6, 1964;19651' Hall6 &I-e Thomas, 1967;1970; Hladik & Halld, 1973; Florence & Hladik, 1980).

Research at the Lopd Reserve, in central Gabon, began at the Station d'Etudes des

Gorilles et Chimpanzds (SEGC) in 1983 and is on-going. The focus of much of the research

has been the ecology of lowland gorillas and chimpanzees but this has involved collectingand identifying plant species eaten by apes and running transects to describe the vegetation ofthe apes' habitat. Among the important food species of apes at Lop6 (Tutin & Fernandez,

1993; Williamson et aI., 1990) were two new species of tree (Coln lizae [Sterculiaceae], N.Hall6, 1987 and Dialiurn lopense [Caesalpiniaceae], Breteler, in press). The discovery of the

former was particularly surprising as it is the commonest tree species within the main study

area and has distinctive, enonnous, leaves (Tirtin et al., t99la). Both of these new species

have restricted geographical ranges suggesting that many undescribed species remain in the

botanically unexplored parts of the country.

Recent work at Lopd has included more syst€matic botanical collecting and an extension ofthe enumeration of trees along ftansects. Analysis of these data has allowed the forest withinthe Reserve to be classed into 20 types that differ both in structure and species composition(Whire, 1992).

Similar ecological research, focussing on great apes, is underway at several other sites inthe ropical forests of cenfral Africa (e.9. Carroll, 1986; Fay, 1989; Kano & Mulavwa, 1984;

Kuroda, 1992; Malenky & Stiles, 1991; Mitani, 1992; Nishihara, 1992; Yamagiwa et al.,1992) but comparisons of diets between areas have been hindered by lack of background dataon the variety and density of plant species available as potential foods at each site.

Data on the vegetation ofthe Lopd Reserve are available from several sources, but publi-cations are limited to a single vegetation type and are not readily accessible (Descoings,

1974; Reitsma, 1988), or are in the form of unpublished Ph.D. theses (White, 1992;Williamson, 1988). Here we present a list of plant species, identified to at least family, thatoccur in the SEGC study area at Lop6. For each species, the life-form and typical habitat isgiven and the parts eaten by gorillas or chimpanzees are listed. Density data are included forall species of tree with diameters at breast height (dbh) > 10 cm recorded on vegetationtransects. The list is certainly incomplete and is biased in favour of habitats well represented

in the SEGC study area, but it provides the basis for inter-site comparisons both of apefeeding ecology and of botanical inventories.

STUDY AREA

The Lopd Reserve covers 5,000 km2 and extends in a tapering rectangular shape from itsnorthern boundary, the river Ogoou6 (0'3'S) to Mont Ibondji (l'10'S) between latitudes11"17'-11o50'8. Altitudes vary from 100-700 m. A chain of mountains runs almost north-south through the centre of the Reserve and elsewhere the terrain is rugged with numeroussmaller hills and steep valleys.

The Reserve includes a complex mosaic of vegetation types. In the north and east thereare areas of savanna interspersed with gallery forests and isolated forest patches (Figure l;see also aerial photograph in Harrison & Hladik, 1986). These savannas are thought to benatural in origin (Aubreville, 1967; White,1992), dating from the Pleistocene, and activerecolonisation by forest is underway except when arrested by annual burning. Most of the

List of Plant Species Identified in the Lope Reserve 25r

Fig. 1. The Lopd Reserve withsavanna areas (stippled), theSEGC study area (hatched box)and the loca-tions of the five 5 kmvegetation transects (Sites 1-5).

savannas are burnt each year for management purposes but some are protected bywatercourses or by their inaccessibility. Within the forest block, areas adjacent to thesavannas have discontinuous canopy cover and a dense unders0orey dominated by herbs ofthe Marantaceae and ZingSllrlraceae, classed as 'Marantaceae Forest' (Irtouzey, 1968). Thisforest type is thought to be of relatively recent origin, reflecting the re-colonisation ofPleistocene savannas (Aubreville, 1967; de Foresta, 1990; White,1992). Deeper into theforest block, canopy cover becomes continuous, plant species diversity increases and the

density of herbaceous plants in the understorey decreases dramatically: this forest type isclassed as 'Closed Canopy Forest'. Within these two major forest types there are smallerareas of permanent water, rocky outcrops, and areas where the vegetation has been affectedby selective logging in the past. In each of these cases, plant species occur that are rare, orabsent, elsewhere.

The SEGC study area covers about 50 km2(0o10'S, l1o 35'E) of mainly MarantaceaeForest to the south and west of the savanna zone but extends into Closed Canopy Forest !othe west of the major mountain range. Parts of the study area were selectively logged between1960-70. Asinglespecies, Aucoumeaklainearw wasextractedatanaveragedensityof 1.5

trees per hectare and logging occurred in both Marantaceae Forest and Closed Canopy Forest.

Mean annual rainfall is 1506 mm (1984-92). The climate is characterised by a long drysetson of about 3 months from mid-June to mid-September. Temperatures vary little ovgr the

year but are lowest during the dry season when constant cloud cover during the daylight

252 C. E. G. Turnr, L. J. T. Wrrre, E. A. Wu.unusoN, M. FEnNANDEz & G. McHunsoN

hours results in low evaporation rates and high relative humidity (Hladik, 1973).

METHODS

Vegetation data presented in this paper were collected during studies with different aims, and

hence methods vary somewhat. White (1992) conducted the most extensive vegetation

survey: five 5 km line-transects were established across major drainage features (c/. Norton-

Griffiths, 1978) in areas which had experienced different logging histories. Trees and lianes

)10 crn dbh were identified and measured in a strip 5m wide along each ffansect (providing a

sample of 2.5 ha in each area), and Eees 270 cm dbh were similarly enumerated in a stip50m wide (25 ha in each area). Figure 1 shows the location of the tansects. Sites 1, 4 and 5

were within the SEGC study area, while Sites 2 and 3 were 35 km south-west, in an active

logging concession and, as little information is available on ape diet in this area, these data

are not included here. Williamson (1988) enumerated a sample of 4 ha of ftees )10 cm dbh inlOm wide snips along a line-fiansect and several elephant paths, selected to sample various

gorilla habitats within the SEGC study area.

In addition to these vegetation samples, systematic collections were made of plant species

consumed by gorillas (Tutin & Fernandez, 1993; Williamson et al., 1990), chimpanzees

(Tutin & Fernandez,'/993), black colobus (Harrison & Hladik, 1986), grey-cheeked

mangabeys (Ham, in prep.), forest elephans (Whit€ et al., 1993) and other species of large

mammal (SEGC, unpublished data). Opportunistic botanical collections have been made ofplants encountered in flower. Voucher specimens are lodged at herbaria at Edinburgh (E),

Libreville (LBV), Kew (K), Missouri (MO), New York (NY), Oxford (FHO), SEGC and

Wageningen (WAG). Nomenclature follows the Flore du Gabon for published families(N=75) and the Flora of West Tropical Africa for other families, except where stated.

Plant life-forms were defined as follows:

Herb (H): Non woody monocotyledons and dicotyledons both free-standing and climbing, including grasses and

sedges.

Fern F): Members of the Pteridophyta rooted in the ground.

Eptphyticfern (Ep): Pt€ridophyta that re rooted on free trunks or branches.

Treefern (TD: Pteridophyta with rhizome rising like a nee runk.

Palm (P): Members of Palmae family.

Liue (L): Climbing woody plans that depend on other vegetation for support.

SIuzD (S): Woody plants, often with multiple stems that do not exceed 2m in height.

Climbing sftrrD (CS): Woody plants, often with multiple stems that do not exceed 2m height when free-

standing but" if supported by other vegetation, can grow much higher.

Tree (f): Woody plants, usually with single stems, growing to heights in excess of 2m.

Epiphyte (Ep): Woody plants that require sup'port of host early in life but may become free-standing once tlehost Fee dies.

Parusite (Pa): Plants wholly, or partly, dependent on other plants for nutrients.

The habitat-types used here are broad categories combining vegetation types defined by

White (1992) on the basis of species composition and structure and are defined as follows:Marantaceae Forasl (MF): The canopy is discontinuous particularly in the middle storey allowing light to

penefiate to the forest floor. The understorey is characterised by dense herbaceous growth of species ofMarantaceae and Zingiberaceae,

Closcd Canopy Forest (CCF): Characterised by more continuous canopy cover, especially in the middle layer

(10-20m). Herbaceous growth in the understorey is sprse,

Forest-savawu interface (F/S): This interface is clear-cut in areas where savannas are subject to annual buming

List of Plant Species Identified in the Lope Reserve 253

but less so if savannas are protected from fire, as,progressive colonisation by woody vegetation occurs.

Certain species of shrubs and trees are particularly common along edges although most also occur at lowerdensities in Marantaceae Forest.

Savanna (S): Areas dominated by Graminae with scattered fire-resistant shrubs.

Rocty (R): Vegetation growing in areas around rocky outcrops where the soil is thin. Trees rarely exceed 30 cmdbh and while the canopy can be close4 it is generally only 10-20 m high.

Water (W): Marshes and the banks of permanent sEeams.

Distarbed (D): Areas that have suffered disturbance from selective logging or road consfiuction in the recent past.

Foods of gorillas and chimpanzees have been recorded since 1984 from faecal analysisand observation, see Tutin & Fernandez (1993) and Williamson er al. (199O) for details ofmethods.

RESULTS

Species RecordedTable I lists 676 plant species, belonging to 9l families, that occur in the SEGC study area inthe lopd Reserve. Of these, 509 have been identified to species, 108 to genus and the remaining59 to family. Each species which is not, as yet, fully determined is given a collection number,which will be referred to in future publications and updates. For species recorded on fiansects(i.e., those that reach at least l0 cm dbh), density data are included, but many species wererare. Rubiaceae was the best represented family in numerical terms, with 90 species, while 30families were represented by a single species.

At least two species (Mangifura indica and Citrus sp?) are cultivars. Their precise historyis unknown but they are likely to be of ancient origin as from the 15th Century the RiverOgoou6, which is the northem limit of the Lopd Reserve, was a trading route and the savannazone has long been the site of human settlement. Young Citrus trees are not uncommon asgorillas and elephants disperse seeds but Mangifera does not appear to regenerate naturally:the majority of fruit are consumed by primates before they reach maturity and the remainingmature seeds are subject to intense predation by bushpigs and rodents.

Life-formsOf the 676 identifiedplants in Table 1,345 are trees, 113 herbs,55 fems,68 lianes and 63shrubs. Other life-fonns were represented by fewer than 12 species.

Habitat Preferences

Much of the SEGC study area is Marantaceae Forest (MF) and this forest-type was dominantin two 5 km transects (Sites I & 5 in Figure l) compared to a single transect where ClosedCanopy Forest (CCD predominated (Site 4). Thus, large tees (> 70 cm dbh) were enumeratedin 50 hectares of MF compared to 25 hectares of CCF and all trees > 10 cm dbh wereenumerated in 5 hectares of predominantly Marantaceae Forest compared to 2.5 hectares ofClosed Canopy Forest. The diversity of tree species recorded in the transect sample inMarantaceae Forest and in Closed Canopy Forest in the SEGC study area was similar, with129 and 146 species respectively. However, 130 additional species oftree were recorded ontwo 5 km transects in CCF (Sites 2 & 3 in Figure l) 35 km from the SEGC study area (Whire,1992). This clearly shows that CCF is much richer in tree species than is Marantaceae Forest.

The habitat preference of each species is listed in Table 1. [n cases of multiple habitats,

254 C. E. G. Turnq, L. J. T. Wmre, E. A. WurausoN, M. FEnNANDU & G. MCHNNSON

Thbte l. Plant species identified in the SEGC study area in the Lop6 Reserve

FAMILY / Species LIFE-FORMI PLANT PART HABITAT3EATEN2

Gorilla Chimp

DENS ITY (individualsTkmz)a> 10 cm dbh > 70 cm dbhMF CCF MF CCF

PTERIDOPHYTAAdiantum vogeliiArthropteris orientalisAspleniurn africanurnAsplenium emnrginatumAsplenium hemitomumAsplenium jaundeense

Asplenium variabileBlotiella curroriB olbitis acro stichoidesBolbitis auriculataBolbitis gabonensisBolbitis heudelotiiCeratopteris cornutaChristella dentataCyathea cameroonianaCyclosorus dentatusD icranopteris linearisDiplazium sammntiiDiplaziwn welwitschiiLastreopsis curroriLomariopsis hederaceaLomariopsis c ongoensisLomariopsis sp. SEGC 536Lycopodium cernuurnLygodium microphyllumMarattia fraxineaM ic r o gr ammt ow ar ie ns i s

Microlepia speluncae

Microsoriwn punctatum

Nephrolepis biserrataNephrolepis undulata

Oleandra distentaPellaea donianaPellaea holstiiP hynwto s o rus s c olopendria

P iryr o gr amm.e c alomelano s

Platycerium stemnriaPteris atrovirensPteris similisPteris acanthoneuxaPteris sp. SEGC 528

S e I a g ine lla c athe dr ifo liaSelaginella myosurus

Selaginella v ersic olorSelaginella vogeliiT ric ho nune s b al lardianum

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W (l'IF)w (ccF)wCCFCCFCCFMFF/SwR (NF)w(MF)R (NF)F/S

F/S

R(l!F)RR (l'F)F/S;MFsMFS

w(MF)MFMFw(Nlr')F/S

wMFw

I See Methods for definition of Life-forms;

2 L= leaf; F=fruit; p= pith; S= seed; FL = flower; B= bark; G- galls;

' See Methods for definitions of Habitat-types;a Density of trees calculated from ftansect sample: MF > l0 cm dbh, 5

dbh, 50 Ha; CCF > 70 dbh, 25 Ha (see text). * indicates present

transect sample.

Ha; CCF > 10 cm dbh,2.5 Ha: MF >70

as a tree of this size but not recorded in

List of Plant Species ldentified in the Lope Reserve 255

FAMILY / Species LIFE-FORM PLANT PART HABITATEATEN

Gorilla Chimp

DENS ITY (i ndividuals/km')> l0 cm dbh >70 cm dbhMF CCF MF CCF

T ric homane s c upre s s o ide s

Trichomanes guineense

Trichomnnes cf. manniiT rip I op hy ll urn b uc hho lz iiT r ipl o p hy ll wn p r o t e ns umT rip I op hy llwn s e c undifo rmeTriplophyllwn sp. SEGC 469Triplophyllwn vogeliiVittaria guineensis

ANGIOSPERMAE

DICOTYLEDONEAE

AcanthaceaeAnisotes mocrophyllaArystasia gangetica?DiclipteraUTW 0022Hypo estes vertic illarisJusticia tenellaP s e uder ant hemum tunic atumLJTW 0748LJTW 0910

AnacardiaceaeAntrocaryon klaineanwnLannea welwitschiiMangtfera indicaP seudospondias longifuliaP seudo spondias mic roc arpaSorindeia gilletiiTrichoscypha acuminataTrichoscypln abut

AnnonaceaeArtabotry s ? rhopalocarpwArtobotrys thomsoniiC leistopholis staudtiiErwntia chlorantlnHexalobw crlspiflorusM o runtho taxis c o n go ens isM onantho tatcis dec linaMonodora angolensisP ac hyp o dant hiwn s t audt iiP olyalthia sunveolens var. gabonicaP olyalthia sunv eolens v at . s runv e olensPopowia declinaUvaria sp. SEGC 253Uvaria sp. SEGC 256Uvaria sp. SEGC 262U v ar ias tr wn p i e r r e anurnUvariodendron sp. LJTW 0706Xylopia aethiopicaXylopia hypolampraXylopia parvifloraXylopia phloidoraXylopia quintasiiXylopia cf. quintasiiXylopia staudtiiLJTW 0608

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MF; DR(w)wccF (w)MF(w)

FMF; RDMFwMFFCCF

MFF/SMFFCCFR; F/S

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256 C. E. G. Turnr, L. J. T. Wrurn, E. A. Wn-uerrasoN, M. FEnNANDU & G. McHtnsoN

FAMILY / Species LIFE.FORM PLANT PART HABITATEATEN

Gorilla Chimp

DENS ITY (indi viduals/km2)> 10 cm dbh >70 cm dbhMF CCF MF CCF

ApocynaceaeAlstonia booneiHolarrhena floribundaLandolphia cf . heudelotti

I-,andolphia sp. LJTW 0079

Landolphia sp. SEGC 526

Picralima nitidaRauvolfia macrophyllaRauvoffia vomitoriaT ab e rnae mont ana c r as s a

LJTW 0306LJTW 479

AristolochiaceaeP araristolo c hia flo s - avis

BalanophoraceaeThoninngia sanguinea

BalsaminaceaeImpatiens sp. LJTW 0874

BegoniaceaeBegonia spp. (at least 2 species)

BignoniaceaeKigelia africanaNewbouldia laevis

Spathodea campanulataBixaceae

Bixa orellanaBombacaceae

Ceiba pentandra

BoraginaceaeCordia cf. millenii

BurseraceaeAucoumea klainearaC anar ium s c hw e infur t hiiDacryodes buettneriDacryodes edulisDacryodes klaineanaDacryodes normandiiDacryodes sp. LJTW 591

Santiria trimeraCaesalpiniaceaeAfzelia bellaAfzelia bipindensfsAfzelia sp. LJTW 450

Amphimas ferrugineusAnthonotha ferrugineaAnt hon o t ha mnc r op hy ll a

?Anthonotha sp. LJTW 0641

Ap lwn o c aly x emnr g i ne rv at us

Augounrdia letestuiBaikiaea insignisBerlinia auriculataBerlinia bracteosaCalpocalyx klaineiCassia manniiCassia mimosoides

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F/S *

MFCCF 40

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F/S *

MF I2OMF*

F/S

F/S 40

MF*

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Pa

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CCF 80 8

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MF 160 40

MF; F/S *

F/S

ccF 1600 *'!

F/S *

w (ccFl r20w (lv{F) 1S0 8

CCF 640F/S *

S

Cassia obtusifutin CS F/S

Cooaifera mildh-aedii T CCF * *

List of Plant Species ldentified in the Lope Reserve 257

FAMILY / Species LIT'E-FORM PLANT PART HABITATEATEN

DENSITY (individuals&m2)> 10 cm dbh > 70 cm dbhMF CCF MF CCF

Crudia gabonensisCryptosepalum staudtiiCynometra ?manniiDaniellia klaineiDetariurn macrocarpumDialium ?bipindensrsDialium dinklageiDialium lopenseDialiwn pachyphyllumDialium soyauxiiDialiwn sp. SEGC 525D is temonant hus benthamianusD uparquetia orc hidac eaErythrophleum ivorenseEurypetalwn batesiiG ilbertiodendron dew evreiGuibourtia demewiiGuibourtia ehieGuibourtia tessmanniiHylodendron gabunenseHymenostegia ?klaineiHymeno s te g ia p e ll e gr iniiJulberrurdia brieyiJulberrardia seretiiMezoneuron angolenseN e o c hev alier o de ndron st ep haniiP ar ab er I i nia b ifo li o lataP elle grini o de ndr o n dip hy I lwnS c orodop hlo eus ze nkeriSindoropsis le-testuiSwartzia fistuloidesT e s smannia dew i lde manni anaT e trab er I inia b ifo I io lat aLJTW 395

CelastraceaeErcnymus congolensisHippocratea myrianthaSalacia mayumbensisSalacia sp. LJTW 0308Salacia sp. SEGC 412

ChrysobalanaceaeAcioa sp. SEGC 499C hrysobalanw icaco atacorensisMagnistipula sp. LJTW 0701Maranthes aubrevilleiMaranthes gabunensisMaranthes glabraParinari excelsa

CombretaceaeCombretum pecoense

Combretum platypterumC ombretuml P teleopsis sp. LJTW 0369

CompositaeAgeratum sp. LJTW 0545Aspilia africarnBidens sp. LJTW 0215Bidens sp. LJTW 0544

Gorilla ChimT CCFT S;L;BS; L;BF(W)r F/S (W)T CCFT S; L MFT CCFTFFCCFT F; S; LF;S;LFTFFCCFT CCFT F/ST L L;FL FL F/ST CCFT CCFT CCFT CCFr w (ccF)TSFTSFr F/S (W)T CCFT S;L; BS; L; B F/S;MFr ccF (w)L F/STFT CCFr ccF (w)T ccFT CCFTLFr w (F/s)T CCFT CCF

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258 C. E. G. Turnq, L. J. T. WmrE, E. A. Wu-ueusoN, M. FENNANDU & G. MCHTENSON

FAMILY / Species LIFE.FORM PLANT PART HABITATEATEN

Gorilla Chimp

DENS ITY (individuals/km2)> l0 cm dbh > 70 cm dbhMF CCF MF CCF

Launea sp. LJTW 0533

LJTW 0423LJTW 0494

ConnaraceaeAgelaea paradoxaAgelaea pentagyna

Agelaea sp. 0092

Cnestis corniculataCnestis ferrugineaConnarus grffinianusJ olly dora dup arq uetianct

Manotes mncranthaRourea myrianthaRourea solanderiRourea tlnmsoniiLJTW 0344LJTW 0479

ConvolvulaceaeI p omo e a b lep lwr op hy ll a

Ipomoea involucrataIpomoea mauritianaHewittia scandens

CucurbitaceaeCogniauxia podolaena

LJTW 0408

DichapetalaceaeDichapetalum barteriDichapetalurn sp. LJTW 0048

Dichapetalum sp. LJTW 0140

Dichapetalwn sp. LJTW 0414

Tapura bouquetianaDilleniaceae

Tetracera podotrichaDipterocarpaceae

Marquesia excelsa

EbenaceaeDiospyros abyssinicaDiospyros boalaDiospyros cinnabarinaDiospyros dendoD io s py r o s kamerunens is

Diospyros manniiDiospyros melocarpa

Diospyros piscatoriaDiospyros polystemon

Diospyros soyauxiiDiospyros stumeolens

Diospyros viridicansDiospyros zenkeriDiospyros sp. LJTW 409

Diospyros sp. LJTW 1004

ErythroxylaceaeErythrorylum mannii

EuphorbiaceaeAlchornea cordifoliaAlclwrnea hirtella?Anthostemmn sP. LJTW 0677

S

MFS; MF

DF/S

CCFF/S; FF/S; FF/SccFDCCFMF

F F/S

MF(w)MF

sS

DS

F/S

MF

F;L;BF RFFMF

DF F lvtF (W)

MF

F/S

w

FFMFCCFCCF

F;S F;S F(R)F

F;S F FCCF

FFF;S F;S F

MFFFFLFF;S F R

RF/S

HHH

LLLCS

CSLsLLLLLS

LL

HHHH

TCS

LLT

TTTTTTTTTTTTTTT

F/S

F/SCCFw (lvtr)

320 40

340

:F

,F

*4040

1140 24020

180 {<

240160 80

1280 r20**s 40

60 40800 40

20tl.

60

20

CS

S

T

List of Plant Species Identified in the Lope Reserve 259

FAMILY / Species LIFE-FORM PLANT PART HABITATEATEN

DENS ITY (individual s/km2)> 10 cm dbh > 70 cm dbhMF CCF MF CCF

Antidesma laciniatwnAntidesnu vogelianumBridelia fercugineaBridelia sp. LJTW 0932?Cleistanthw sp. LJTW 546Croton sp. LJTW 0348D is co glypremne caloneuraD uv i gne audia inop i rnt aEuphorbia thymifoliaJ atrop hn g o s sypiifoliaKlaineanthw gaboniaeMacaranga barteriMacaranga gabunicaMacaranga monctndraMaesobotrya duseniiMaesobotrya cf . pynaertiiManniophyton fulvumMaprounea membranaceaMareya micranthaMareyopsis longifoliaPhyllanthw diandrus?Phyllanthw discoidewPlagiostyles africanaR ic ino de ndro n he ude I o tiiSapium ellipticutnThecacoris sp. nov.? GMcP 16015Uapaca guineensisUapaca heudelotiiUapaca paludosaUapaca aff. togoensisUapaca vanhoutteiUapaca sp. LITW 0993LJTW 526LJTW 567LJTW 0658LJTW 0661

FlacourtiaceaeCaloncoba glaucaCarnptostylus manniiCasearia barteriHomalium letestuiH omnlium ? aff . neur op hy llumHomalium sarcopetalwnHornaliwn sp. GMcP 16169Lindackeria dentataOncoba brachyantlrcraScottellia coriacea

GentianaceaeN eur othec a lo e se lioides

GuttiferaeAllanblackia sp. LITW 272Garcinia afzeliiGarcinia cf kolaG arc inia cf. ovalifoliaGarcinia sp. LJTW 152Garcinia sp. LJTW 0447Garcinia sp. LJTW 0767

FF

FFFFF

TTTTTTTTHS

TTLTTTLTTTTTTTTS

TTTTTTTTsL

Gorilla Chi

?F

FFF F/S; W; D

sF/S

CCFMF; D

F; L DCCFsF/S

CCFDDDCCFCCFFDCCFCCFFF/S

FFCCFMFMF

F F/S;MF(W)FFF CCF (W)F MF(W)F MF(W)

F/S (w)CCFMFCCFs (R)

MFFFRMFw (F/s)CCFD(r!F)F/S

MF

s

FCCFMFF/S (w)CCFCCFF/S (w)

140*40

40rF r&

40 280 2280

20* 240

40

500 1000

t20600

r20 720 280

202

20 l020*2

20*

8020

20

80 160r20 8060

tlc

,F

620

FLTTTTTTTsS

T

TTTTTTT

**rrk

280r|(

rF

40:1.

260 C. E. G. Tl-rrnr, L. J. T. Wrure, E. A. Wu.r-nusoN, M. FEnNANDEz & G. McHtnsoN

FAMILY / Species LIFE-FORM PLANT PART HABITATEATEN

Gorilla Chimp

DENSITY (individuals/km2)

> 10 cm dbh >70 cm dbhMF CCF MF CCF

Mammea africanaPentadesnwt buryraceaP entade smt grandifoliaSymphonia globuliftra

HumiriaceaeSac o glottis gabonensis

HypericaceaeHarungana sp. LJTW 0790

P s o r o sp ermum fe b rifu g um

P s o r o s p ermum tenuifo liumVismia guineensis

IcacinaceaeIcaciru manniiLasianthera africana

IrvingiaceaeD esborde sia glauc esc ens

Irvingia gabonensisIrttingia grardifoliaIrtingia roburKlainedoxa gabonensis

Klainedoxa trillesiiIxonanthaceae

O c ht lw c o smus c on g ol ensis

LabiataeSolenostemon sp. LJTW 0543

LauraceaeBeilschmiedia fulvaBeilschmiedia sp. LJTW 37

Beilschmiedia sp. LJTW 439

Beilschmiedia sp. LJTW 530

Beilschmiedia sp. LJTW 537

Hypodaphnis zenkeri

Ocotea gabonensis

LecythidaceaeN apoleonaea imperialisNapoleonaea cf . leonensis

P e t er s ianthus mac r o c arP us

LinaceaeHugonia planchonii

LoganiaceaeAnthocleista vogeliiAnt ho c le i st a ? s c hw einfur thiiMostuea hirsutaMostuea sp. LJTW 0769

Strychnos congolannStrychnos malacoclados

LoranthaceaeHelixanthera manniiLorantha.t sp. LJTW 0119

Tapinanthus sp. LJTW 0444LJTW 0802LJTW 0918

LJTW 0979Luxemburgiaceae

Testulea gabonensis

MalpighiaceaeAcr ido c arp us lo n g ifu lius

TFT F; STT

TF

S

S

TLS

LTL

TTFTFTFTFTF

T

H

TTTTTTLT

TTT

L

TLTS

S

LL

Pa

Pa

Pa

?PA

Pa

Pa

FFFCCF

F CCF

F F/S

sS

LDD; MF

MFF

FF; S FF; S FF CCFFFFF

F/S

S

FFFMF

ccF (w)CCFMFMFCCF

FCCFCCF

F/S

W; DF/S

ccF (D)F/S (w)CCFF/S

MFCCFCCFF/S

F/S

F/S

F

D

60 'F

40400

40

80

40

:N(

,F

4

40 160

160 480 l0220 240 20

60 40 12tl.

580 80 32

160 r20 6

,1.

40 4040

240240

20

560{.

40 r20

160

40

40 160

36

44**

20

t2

40:1.

1610

List of Plant Species ldentified in the Lope Reserve 26r

FAMILY / Species LIT'E-FORM PLANT PART HABITATEATEN

Gorilla Chimp

DENS ITY (individuals&m2)> 10 cm dbh >70 cm dbhMF CCF MF CCF

MalvaceaeHibiscw sp. LJTW 0920Sida rhombifoliaUrena ?lobata

MelastomataceaeCalvoa monticolaDissotis congolensrs

Tristemma leiocalyxLJTW 0023LJTW 0481

SEGC 319

MeliaceaeCarapa proceraE nt and o p hr a gma c ando I eiEntandophragma utileLovoa trichilioidesTrichilia monadelphaTrichilia prieureana

MenispermaceaeSEGC 43I

MimosaceaeAlbizia ferruginea?Calpocalyx sp. LJTW 0611

Cathormion altissimumCylic odis cus gabunensisDicrostachys cineraEntada gigasF illaeop sis disc ophoraNewtonia leucocarpaParkia bicolorParkia filicoideaP entac lethra eeweldeanaP entac lethra macr op hy llaP ip tadenias tr um afric anumSamnnea leptophyllaTetrapleura tetraptera

MoraceaeDorstenia aff. barteriFicw barteriFicw bubuFicw elasticoidesFicw ingens

Ficw kimuenzensisFicus macrospertnctFicw mucuso

Ficw ovataFicw politaFicw pseudomangiferaFicw recuryataFicw surFicw thonningiiFicw variifuliaFicns sp. SEGC 443M ilic ia (C hlorophora) excelsa

Musanga cecropioidesMyrianthus arboreusTreculia africarn

FCCFCCFF

SMFMF

F F/S

MFCCFF/S

FS (D)MFFF

FFFMFLFL; FL F

FCCFF

F/SMFS

DS

S

MFDMF

CS

HS

HHHHHLL

8*

,F ,F

100 600rF

:N<

**40

880

TTTTTT

L

TTTTS

LTTTTTTTTT

HEpEpEpTLEpTTEpEpEpTEpTEpTTTT

FFLLS

w (ccF)FFMFFFF/S

MFFMF

CCFFFMFF;L F MF

R (s)FFMF

F F/SF; L F MF

R (S)FFMFL;B L MFFFMFL; B; G F/S; MF

DF; B F MF(W)S S; L MF(w)

48

4:F

12

44r64*

:k

2

6

4t436

30

20 80

20*&0 240700 44040 :F

4040 'lc

80*64***

262

FAMILY / Species LIFE-FORM PLANT PART HABITATEATEN

Gorilla Chimp

DENSITY (individualsftm2)> 10 cm dbh >70 cm dbhMF CCF MF CCF

Treculia obovoideaMyristicaceae

Coelocaryon prewsiPycnanthus angolensisS cyp ho c ep haliwn o c hoc oaStaudtia gabonensis

MyrtaceaePsidium guineense

Syzygiwn sp. LJTW 3lSyzygiurn sp. LJTW 0147SyzygiwnlEugenia sp. LJTW 0381

NymphaeaceaeLJTW 0815

OchnaceaeC ampy I o s p e r murn el o n g atwnLophira alataOuratea flavaOuratea cf . myrioneura?Ouratea sp.LJTW 0432R lnb dop hy ll wn c al o p hy I I um

Sauvagesia erectaLJTW 0772LJTW 0882

OlacaceaeCoula edulisDiogoa zenkeriHeisteria pan ifuliaOngokea goreP ty c ho p etalum p e t io lat wnStrombosia pustulataStrombosia zenkeriStromb o siop sis tetrandra

OleaceaeLinociera aff . mannii

OxalidaceaeBiophytum petersianumBiophytum talbotii

Pandaceae

Centroplacw glaucinwPanda oleosa

PapilionaceaeAganope impressaAganope sp.

Crotularia pallidaDalbergia aff . rufaD e s mo diurn ramo s is s imumDesmodiurn scorpiurwEriosema glomeratumErythrina thollonianaI ndigofera c onj ugata v ar . occ identalisLep to derr is hyp ar gy re aMilleuia laurentiiMillenia sanaganaMilleuia conrauiMilleuia sp. LJTW 0038

Platysepalum sp. LJTW 0603

Pterocarpus soyawcii

TTTT

sTsT

FS;L

F

FF

F; L FL

FF

L; B; FL

L;B L

S;L;FL S;I

CCF

CCFFCCFCCF

sMFCCFCCF

w (s)

MFF/S; MFF/S; MFF/S; MFMFMFsCCFS

CCFCCFFFCCFCCFCCFCCF

F/S

S

w(R)

CCFF

CCFMFS

F/SS

S

sw (NrF)

sF/S

CCFF/SF/S

MF(w)CCFF

200

360140 r2020 920* 200

,1.

r2060

*

M286 r72**

2

TTTTTTHS

S

2740 'F 90 8*

:1.

TTTTTTTT

1760 60*r

2404480 14 12

160

4002720

r20 2000

100 272020'k**

40

20

24080 :rc 68 20

HH

TT

LLsLHHS

THLTTTTLT

List of Plant Species Identified in the Lope Reserve 263

FAMILY / Species LIFE-FORM PLANT PART HABITATEATEN

Gorilla Chimp

DENS ITY (i ndividual s/km2)> l0 cm dbh > 70 cm dbhMF CCF MF CCF

Tephrosia purpureaUraria pictaZornia latifuliaLJTW 0469LJTW 0591LJTW 0599LJTW 0,602LJTW 0609LJTW 0720LJTW 0950SEGC 395

PassifloraceaeBarteria fistulosaParopsia grewioidesLJTW 0360

RhizophoraceaeAnisophyllea sp. LJTW 0475Anopyxis klaineanaCassipourea congensisPoga oleosa

RubiaceaeAidia ochroleucaAtractogyne gaboniiBertiera batesiiBertiera mildbraediiBertiera sp. LJTW 0139Borreria pusillaBrenania brieyiCanthiwn sp. LJTW 0297

Cephaelr sp. LJTW 0202Corynanthc mayumbensisCrossopteryx febrtfugaD ic ty andra arboresc ens

Diodia scandensFeretia sp. LJTW 0646?Gaertnera sp. LJTW 0316Geophila afzeliiHeinsia crinitaLeptactirn cf . arnoldianaMassularia acuminataMitacarpus scaberMitragyru ciliataMwsaenda cf . debeawiiMussaenda cf . tenuitloraM ussaenda pp. LJTW 058 1

Nauclea didderrichiNauclea latifoliaNauclea vandergrchtiiOldenlandia corymbosaO lde nl and.i a I anc ifo I iaOtomeria guineezsisOtomeria micranthaOryanthus unilocularis?Oryanthus sp. LJTW 0903Pauridiantha aff . dewevreiPauridiantha efferataP aur idiant lrc fl o r i bunda

HHHS

LLLTTTL

S

S

S

F/S

CCFCCFccFCCFccF (w)F/S

MF

F/S; MF; DF/S; RCCF

CCFFRCCF

F/S

MF; Dw (ccF)MFCCFsMFCCFF/S

FS

MFMFF/S

CCFMFF(w)DFS

wDDMFFswS

sMFsF/S

MFF/SD; F/SD; F/S

80

4040

320

:F

520 600TTL

4,F

80:1.

TTTT

t6

60

FFF

TLS

TS

HTS

sTsTHsS

HTTTHTLLLTTTHsHHTTTTT

B; LF

FFF

*

180

rF

340 2240

40

80 r20

20 600

*

*220100 r20

2& C. E. G. Turw, L. J. T. Wru'rE, E. A. Wu.uausoN, M. FEnNANDEZ & G. McPrfiRSoN

FAMILY / Species LIFE-FORM PLANT PART HABITATEATEN

Gorilla Chimp

DENS ITY (individual s/km2)

> 10 cm dbh > 70 cm dbhMF CCF MF CCF

Pausinystalia johimbeP aus iny stal ia mac ro c er asPavetta hispidaPavetta aff . viridilobaPavetta sp. LJTW 0006Pavetta sp. LJTW 0205?Pavetta sp. LJTW 536Porterandia cladanthaPseudosabicea batesiiP s e u^do s ab i c e a fl or ib undaP seudo sabic ea mildbraediiP sy c ho tria p e dunc ulari sPsychotria vogelianaPsychotria sp. LJTW 0012Prychotria sp. LJTW 0l7lPsychotria sp. LJTW 0174Psychotria sp. LJTW 0209Psychotria sp. LJTW 0638

Psychotia sp. LJTW 0657

Rothmannia whitfteldiiRutidea dupuisiiSabicea efulenensisSabicea mollisS ac o sp errna p anic ulatumTarenna confertaTarenna eketensis

Tarenna sp. LJTW 0272Tricalysia anomalaT ric aly s ia mnc r o p hy ll aTricaly sia cf oligoneuraTricalysia cf . pallensTricalysia sp. LJTW 0081

Tricalysrc sp. LJTW 0082Trichostachys aureaUncaria africanaLJTW 0098LJTW 0372LJTW 0396LJTW 0417

LJTW 0433LJTW 0628LJTW 0638

LJTW 0642LJTW 0646LJTW 0662LJTW 140

LJTW I7O

LJTW 387LJTW 514LJTW 528

LJTW 566SEGC 288

SEGC 291

SEGC 508

RutaceaeAraliopsis sp. LJTW 353

Citrus sp. SEGC 557

F20MF 20

CCFMF 20

CCFF/SMF 40D; MF 120CCFDDMFF/S

F/S :F

CCFCCFMFMFCCFMFFMFMFF/S

DF/S

CCFDMF(W) ',rc

MFMFccF (w)MFMFF/SDMFCCFMFccFR; W (F/S)MFrrF (w)F/S

S

MF 40

DCCFCCFCCFMF 20

MFDMF

ccFF/S

404040

FFL

TTS

THS

TTHLHS

sTS

HS

S

S

TLLLLTLTTTS

sS

S

HLTsTsS

TsTS

TTTTTTTLLs

LL; FL

F; L F

200

80TT

List of Plant Species Identified in the Lope Reserve

FAMILY/Species LIFE-FORM PLANTPART HABITAT DENslTY(individuals/km2)EATIN ) l0 cm dbh > 70 cm dbh

Gorilla Chimp MF CCF MF CCFFagara?macrophylla T F/S; MF !8

Fagara tessnunnii T CCF 40Tecleaaff.verdooniana T R *Vepris cf. lonsii T R 20

SapindaceaeBlighiawelwitschii T MF r.

Chytranthus talbotii T MF(W)Eriocoelwnmatocarpwn T F/S; MF 540 280Eriocoelutn?paniculatwt T MF '* *Eriocoelwn sp. UTW 132 T MF nGanophyllwn gigannrmt T F F MF 80 Z0Lecaniodiscuscupanoides T F F MF 100Placodiscuscf.opacus T CCF g0

SapotaeaeBaillonella toxisperma T F F CCF ,& 40 Z 4Donellaogowewis T F F F/S {.

Donclla?prmiformis T CCF g0Gambeya africana T F F CCF * 40 !r g

GambeyasubnudaTFFF*40**?Lectomedoxn sp. LJTW 237 T MF * 2Letestua dwissima T MF * 4ManilkarafouilloyaruTFFCCFlFOmplalocarpwnprocerwnTF**:r*PachystelabrevipesTFFR**LITW 0228 T F F CCF 160 ,&

ScrophulariaceaeCycniwt carnporurn H S

ScytopetalaceaeScytopetalwt sp. IJTW 0017 T F F F l2O U0

SimaroubacraeOdyendyea gabonewis T CCF '*Qwssia gricana S F

SolanaceaeSchwenckia americana H S

SterculiaceaeCola lizae T F F; L; p MF 7460 g

Colanahoundensis T CCF (W) g0Colasp. SEGC 332 T MF *Lepnnychia echinocarpa S MFNesogordoniapapaverifera T F * * :r.

'*Pterygota?bequaertii T CCF :*

Scaplapenh,trnblackii T CCF 160Sterculia tagacantln T L MF 40 2

StyracaceaeAfrostyrax lepidopltyllus T CCF g0

ThymelaeaceaeDicranolepis buchlalzii S CCF?Diuanolepis sp. LJTW S MFOctolepis decalepis S MF

TiliaceaeClappertonialicifulia H S; DDuboscia ma$ocarpa T F; S F; S F 40 l2O 2 *Triwrfetu cordifolia S S

UlmaceaeCeltistessnannii T F;L F F * 200 g 2gCeltissp.LnW 444 T CCF 40

265

266 C. E. G. TUrNq, L. J. T. Wru.re, E. A. WnunusoN, M. FEnNANDEZ & G. MCHmnSoN

FAMILY / Species LIFE-FORM PLANT PART HABITATEATEN

Gorilla Chimp

DENS ITY (individuals/km2)> l0 cm dbh >70 cm dbhMF CCF MF CCF

VerbenaceaeVitex donianaVitex ?rivularis

ViolaceaeRinorea ilicifuliaRinorea sp. LJTW 0055

Rinorea sp. LJTW 0132Rinorea sp. LJTW 022IRinorea sp. LJTW 0596Rinorea sp. LJTW 0656Rinorea sp. LJTW 0899

VitaceaeCissru aff. barteriCisszs dinklageiCissns leonardi

VochysiaceaeErismadelphw exsul

MONOCOTYLEDONEAEAgavaceae

Dracaena fragansDracaena sp. LJTW 0916Sansevieria sp. SEGC 565

AmaryllidaceaeScadoxus cinnabarinus

AraceaeAnchomanes dffirmisNephytis sp. LJTW 0480

CommelinaceaeAneilema beninienseCommelina capitataPalisota ambigwPalisota sp. SEGC 551

CyperaceaeBulbostylis cf. densa

Bulbostylis lanicepsFimbristylis pilosaPycreus sp. LJTW 0526

Scleria boiviniiLJTW 0131

LJTW 0407

GramineaeAndr op o g o n fas t i g iatusAndr o p o g o n p s eudapr ic wAxonopus compressusBrachiaria jubata

Centotheca lappaceaHyparrhenia diplandraOlyra latifuliaOplismenw hirtellusPanicurn dregeanumPanicum grffiniiPanicurn cf . walense

Panicum sp. LJTW 0152Panicu.rn sp. LJTW 0208

Pospalum paniculatwnP ennis e twn po ly s tac hy on

TT

F F/S

CCF

S

TTTTsS

F;L FF

L;P

CCFw (ccF)w(MF)w (ccF)w (ccF)CCFCCF

DFMF

CCF

CCFFR

MF

Fw (ccF)

MFMFMFMF

S

S

S

S

F/S; Dw(MF)D(w)

S

S

S

S

MFsMFMFS

S

S

w (ccF)MFD (MF)S

80

*

40r20

40

LLL

TsH

HH

HHHH

HHHHHHH

HHHHHHHHHHHHHHH

List of Plant Species Identified in the Lope Reserve 267

FAMILY / Species LIFE-FORM PLANT PART HABITATEATEN

Gorilla Chimp

DENS ITY (indi vidual s/km2)

> 10 cm dbh > 70 cm dbhMF CCF MF CCF

Peratis indicaPobeguinea arrectaS c hiz ac hy rium p I aty p hy ll umSp or ob o lus py rarnidalisStreptogynn crinitaLJTW 0388LJTW 0404LJTW 0540

LiliaceaeCurculigo pilosa

MarantaceaeAtaenidia confertaHalopegia azureaH aunwnia liebr e c hs tianaHyps elodelphy s hirs utaHyp s el o de lp hy s p o g ge anaHyps elodelphy s vio lac eaM ar anto c hlo a c o rdifo li aMarantochloa filipesMarantochloa purpureaM e gap hry niwn mac r o s t ac hy umMegaphrynium velutinumT rac hy p hry niwn braunianurn

PalmaeCalamus sp. SEGC 554Elaeis guineensisEremospatha cabraePodococcus barteriRaphia sp. SEGC 555

PandanaceaePandanar sp. SEGC 556

PontederiaceaeH e t e rant he ra c all ifo lia

SmilacaceaeSmilax kraussiana

ZingiberaceaeAfr amomum I o n g ip e ti o I at umAfrarnomum sp. ?nov SEGC 152Aframomum sp. SEGC 239Afrarnomurn sp. SEGC 323Costw aferRenealmia cincinnataRenealmia macrocolea

S

ssS

MFS

MF(w)S

S

LLFL; P wS;L S;L FS;L S;L FS;L S;L CCFS;L S;L FL;P wPFL;P wF;L;PF;T FF;L;PF;L FPW

F/S; MFF; P F/S; MF

F; L; PF; P MFCCFw (s)

w (s)

W (I,F)

F/S

F; P F; P F/S

F; P F; P MFF;P F;P FF;P F;P CCF;DPPMFPFPPF

H

HHHHHHHH

HHHHHHHHHHHH

P

P

PP

P

180

HHHHHHH

C. E. G. TurN, L. J. T. Wrurg, E. A. Wu-ueusoN, M. FEnNANDU & G. MCHUNSON

the predominant one is marked first. Of the 676 plant species identified at least to family, 149

occur principally in MF and 152 in CCF. No obvious habitat preferences were found for 90

species (designated as 'Forest' (F) in Table 1) that occur both in MF and in CCF. One

hundred and thee species are found primarily close to the foresVsavanna interface, and 67

(including 23 Graminae) occur only in savannas. Smaller numbers of species were found to

be associated with water (53); with areas disturbed by past loggrng activities (34); or with

rocky outcrops within the forest (28). If these species occur only within a specific habitat-

type, this is indicated in Table l, e.g.R (MF) indicates a species found close to rocky

outcrops in Marantaceae Forest; and W (CCD, one associated with water in Closed Canopy

Forest.

Table 2 compares the l0 commonest species in Marantaceae Forest and Closed Canopy

Forest in terms of basal area and stem density. only two fiee species are arnongst the top ten

for basal area in both forest types and only one species figures among the ten commonest in

terms of stem density in both forest types. These data emphasise the differences between the

two major forest types in terms of species composition.

Foods of Gorillas and Chimpanzees

Of the 676 species recorded in the SEGC study area, 27Vo ptovide food for either

chimpanzees or gorillas, or both. Some families provided no, or few foods, e.g., Pteridophyte

(0 of 55 species) and Connaraceae (l of 13 species) while other families provide food

consistently e.g., Marantaceae (12 of 12 species) and Moraceae (15 of 21 species). Data on

Table 2. Top 10 species in Marantaceae Forest and Closed Canopy Forest atLop6 in terms of basal area and stem density

Species Family Rank (Basal Area) Rank (No. Stems)

MF CCF MF CCF

Aucoumea klaineanaCola lizaeLophira alataP entaclethra mac rop hy llaDacryodes buettneri

P entaclethra eetveldeanaDiospyros polystemon

Hylodendron gabunense

Xylopia quintasiiPycnanthus angolensisTrichilia cf . prieurearutS trombosiop s is teff andraDiospyros dendo

Diospyros zenkeri

Maprounea membranaceaS cyp hoceplnlium o c oc hoa

C entroplac us glaucinas

Xylopia aethiopicaSantiria trimeraCoula edulisAugounrdia letestuiSindoropsis le-testuiStrombosia zenkeriCy licodiscus gabonensisDialium soyawcii

Burseraceae

Sterculiaceae

OchnaceaeMimosaceaeBurseraceae

MimosaceaeEbenaceae

Caesalpiniaceae

AnnonaceaeMyristicaceaeMeliaceaeOlacaceae

Ebenaceae

Ebenaceae

EuphorbiaceaeMyristicaceaePandaceae

AnnonaceaeBurseraceae

OlacaceaeCaesalpiniaceae

Caesalpiniaceae

Olacaceae

Mimosaceae

2

I3

9

8

r05

1

2

3

45

6

7

8

9

10

l0

I7

3

5

6

9

45

6

8

10

Caesalpiniaceae

List of Plant Species Identified in the Lope Reserve 269

Table 3. Number of species of ape foods in different habitat-types at Lopd

Habitat-type No Species Gorilla Food Species Chimpanzee Food Species

Marantaceae ForestClosed Canopy ForestForestForest/savanna edgeSavannaWaterDisturbed areas

Rocky areas

Number Percen

42r753

223

Number Perce

36

2l50l92

r49r5290

103

67

53

34

28

2

4

6

9

6

6

28.2

rt.258.92r.3

4.5

r7.017.6

2r.4

24.213.8

55.618.4

3.0

3.7

I 1.8

21.4

TOTAL/AVERAGE 676 158 23.4 r40 20.7

ape feeding have been collected by faecal analysis and observation over ten consecutive yearsbut new foods are still being discovered. Some of the newly recorded foods come from rarespecies, or from species which fruit irregularly, but others are parts of common species thatare ealen only in unusual circumstances, e.g. when major fruit crops fail, or when apes rangein atypical habitats such as savannas.

Table 3 shows the vegetation types in which plants that provide foods for gorillas andchimpanzees primarily occur. Both gorillas and chimpanzees feed from plants in all of thehabitat-types. Foods from species found exclusively in Closed Canopy Forest are undoubtedlyunder-represented as areas of CCF are on the edges of the study area and less time was spentcollecting data on ape ecology in this vege0ation type. Few foods are provided by savannaspecies but a fifth of the species found at the forest/savanna interface provide food for apes.Both gorillas and chimpanzees eat parts from plants associated with rock outcrops and thosecolonising disturbed areas, but gorillas find more foods from the flora associated withpermanent water. Had this latter category been divided into 'marsh' and 'riverbank',chimpanzees would have emerged as having no food-plants in swamps while gorillas do feedon herbaceous and woody plants that grow in standing water. This reflects the generalavoidance ofwater by chimpanzees.

DISCUSSION

Species RecordedThe 676 species listed in Thble I show the flora of the SEGC study area to be diverse but thelist is far from complete. The transect samples were biased with respect to habitat-type asmost were located in Marantaceae Forest. This forest type is dominant in the north of theReserve, close to the savannas but the more botanically divene Closed Canopy Forest is thepredominant forest-type in the Reserve. This bias means that the species listed in Table Ishould not be considered as representative of the Lopd Reserve as a whole. To date, a total of864 species have been recorded for the Reserve but the southem half remains botanicallyunexplored.

Another source of bias is by life-form: Only tees and lianes with a dbh of l0 cm, or more,were included in the systematic transect enumerations, thus our data on herbaceous plants,shrubs and smaller lianes come from opportunistic collecting. The large number ofPteridophytes recorded (55 species) reflects two collecting rips by a specialist, but is far fromcomplete (N. Mundy, pers. comm.). Reitsma (1988), working in the Lop6 Reserve 30 km

270 C. E. G. Turnq, L. J. T. Wrure, E. A. Wu-unusoN, M. FEnNANDEz & G. McHmnsoN

south-west of the SEGC study area, recorded 69 species of tree of dbh < l0 cm in a one

hectare plot and 125 species in 0.02 hectares where all plants were enumerated. Of the

species recorded by Reitsma in the small plot,247o were Eees, 187o shrubs, l87o lianes and

4OVo lrerbs. This compares to 5lvo tees, 9Vo shrubs, 107o lianes and 25Vo herbs (including

fems) in our sample. An assumption of equivalent distribution of life-forms at the two sites at

Lop6 seems reasonable and this suggests that the species listed in Thble I represent about half

of the total divenity of the SEGC study area: If all tree species have been identified the 345

species represents 24Vo of the total plant diversity, equal to about 1,400 species. Tlvo decades

of botanical collecting in the Makokou area of Gabon resulted in identification of 1,233 plant

species (Florence & Hladik, 1980) giving support to the above extrapolation.

Of the 509 completely identified species, two were previously undescribed: Cola lizae,

Sterculiaceae (Hall6, 1987) and Dialiurn lopense, Caesalpiniaceae (Breteler, in press). At

least two other species of the 618 identified to genus are believed to be new: Aframomum

SEGC #152, Zingiberaceae (J.M. Lock, pers. comm) and Thecacoris GMcP #16015'

Euphorbiaceae. Among the fully identified species there are many new records for Gabon,

e.g. of 13 Ficus (Moraceae) species, two are new country records (Betg et al., 1984); and of

the 16 species of Diospyros (Ebenaceae) two af,e new records for Gabon (ktouzey & White,

1970), including D. polystemon wlichis the fifth commonest tree, in terms of stem density, in

Marantaceae Forest (Table 2).

Habitat-Preferences

Most plant species were found to occur only, or in much greater abundance, in one particular

habitat-type (see Table l). Dfferences between habitat-types are defined by many factors,

principally the amount of available light and water and the quality of the soil. Species that

grow only in, or beside, permanent water appear to need the water but species associated with

the poor soil of rock outcrops may not prefer this habitat but may be limited to them by an

inability to out-compete other species on richer soils. similarly, woody plants that grow in

savanna habitats are not only tolerant of exposure to sunlight but are also resistant to fire, as

annual burning by humans has been a recurring event for several thousand years (Oslisly &

Fontugne, 1993).

The major difference between MF and CCF, the two dominant forest types recognised in

this study, is that more light reaches the ground in the former because of the discontinuous

canopy. This undoubtedly affects the species composition of the understorey herbs and

shrubs, but the lower tree species diversity of MF may simply reflect its relatively recent

origins.In the savannas in the north of the SEGC study area, species of Graminae are ubiquitous

and only four species of shrubs occur in abundance (Nauclea latifulia, Crossopteryx

febrifuga, Psidium guineensis, Bridelia ferruginea). These have fire-resistant bark, or the

capacity for rapid re-glowth ftom root stock after the annual burning. In the savanna zone'

gallery forests gtow in thin ribbons along watercourses that are not swampy and there are a

few isolated forest blocks (1-5 hectares), where the forest limits are not obviously related to

water. The flora of the galleries and bosquets has not been studied systematically, but some

tree species occur only in these habitats (e.g. Cathormion altissitnum, Aphanocalyx

cynometroides, Pachystela brevipes) and some, generally rare MF species become common

(e.g. Uapaca guineensis, Pseudospondias microcarpa and Ficru spp')'

List of Plant Species Identified in the Lope Reserve 271

Savannas protected from annual buming at Lopd are being actively colonised and thisgives trees with seeds that can germinate at the forest/savanna interface an enormouspotential for reproduction. Tree species diversity at the interface is low but successionalchanges are clear with many young individuals of additional species appearing as soon as theedge colonisers provide shade. Large trees able to regenerate at, or close to, the forest edge atLopd include Aucoumea klaineana, Lophira alata and Sacoglottis gabonensis. All thesespecies can form almost pure stands, shading out Graminae and creating conditions thatappear favourable to invasion by other tree species, notably Cola lizae, Dialium lopense,Uvariastrum pierreanwn, Xylopia spp., Klainedoxn spp.

Of the tree species in Thble 1, three show an interesting variation in habitat-preference,depending on whether they grow in CCF or in MF. The three species, Santiria trimera(Burseraceae), Scyphoceplwliwn ochocoa and Staudtia gabonensis (Myristicaeae) are listedas CCF species and the two former are amongst the commonest ten species in this habitat (see

Table 2). All three of these species also occur in MF but there show a distinct micro-habitatpreference, growing almost exclusively on the banks of permanent streams. It seems likelythat this reflects the post-Pleistocene colonisation of savannas, with the flora growing alongstreams being old gallery forest now surrounded by the recently established MarantaceaeForest.

Ape foods, Habitat-types and Inter-site ComparisonsAt Lop6, gorillas and chimpanzees have diverse diets and obtain some foods from plants ineach of the habitat-types (Table 3). The sampling bias against CCF described above, appliesequally to our data on ape foods as far less time was spent observing and tracking apes in thisforest type than in MF. 'Forest' species are generalists that occur in both MF and in ClosedCanopy Forest and 55-6OVo of the 90 species in this category provide food for the apes. Thesespecies, being widespread, though not always common, provide much food as gorillas andchimpanzees both travel widely to exploit rare food resources. The mosaic of habitat-types inthe home ranges of our study groups at Lopd means that a wide range of food is available andperhaps explains why gorillas at Lop6 are the most frugivorous population of the genusstudied to date (Williamson et al., 1990). Foods from the minor (in terms of area) habitatsmake important contributions to ape diet and use of the savanna edge by both gorillas andchimpanzees is highly seasonal and clearly related to fruiting patterns of F/s species(unpublished data). Similarly, aquatic herbs particularly Marantochloa cordifulia(Marantaceae) provide important keystone foods for gorillas at Lort during the dry seasonwhen fruit is scarce (Rogers et al.1988; Tutin e, al.,l99lb).

Both gorillas and chimpanzees are known to show dietary flexibility and in different areasdo not always eat the same foods even when the plants are available. For example, Nishida etal. (1983) found that only 597o of 286 available plant foods were eaten by chimpanzees attwo sites, Gombe and Matrale in Tanzania, although they are only 150 krn apart. Four of the107 foods of lowland gorillas at Ndoki, Congo, listed by Nishihara (1992) occur ar Lop6 burare not eaten by gorillas. Of these, tfuee (Pennclethra seeds, Polyalthia fruit and Tetrapleuraseeds) are common within the SEGC study area Complete inter-site comparisons can only bemade once both lists of ape foods and plant species lists are published for each of the long-tenn study sites. What determines which plants apes eat, and which they do not, is stillunknown but comparisons between sites will clarify ecological differences (e.g. total plant

272 C. E. G. Turnr, L. J. T. Wru're, E. A. Wn r-nusoN, M. FEnNANDU & G. McHlEnsoN

species diversity at each site, the density of each species and the variety of habitat-types).

Information about the density of different plant species is important as the absence of arare species from the food list is less noteworthy than that of a common species. For example,

Musanga cercropioides (Moraceae) is eaten by apes elsewhere in Gabon (Hladik, 1973; Tutin

& Fernandez, 1985) but has never been recorded as an ape food at Lop6. However, while

Musanga is a common colonising tree in most of Gabon, it is extremely rare in the SEGC

study area with only two rees known in 50 sq. lan. In conEast, Pentaclethra spp. arc common

with an average of 1,300 trees per knf in Marantaceae Forest at Lop6, yet gorillas do not eat

the seeds. An importance rating for foods (Tutin & Fernandez, 1993) is also useful when

making comparisons between sites, as consumption of rare foods can easily be overlooked

while differences in important, or keystone, foods indicate real ecological or cultural differences

between populations. Inter-site comparisons will allow a deeper examination of nutrition and

food choice than has yet been possible. Analysis of the importance of factors such as inler-

specific competition and the determinants of population density will also be possible and

should advance our understanding of the geographical distribution of African apes.

Conservation and Management Implications

The value of tropical forest ecosystems is now appreciated, as is the need to protect their

extraordinary diversity of plants and animals. To achieve this, the first step is to compile

inventories, but this arduous task is too often overlooked. Obstacles to this logical first step

are legion but major ones are the expense, expertise and time that are required. Even ifinventory data are collected they are rarely published in an accessible form.

Systematic inventories of plants allow recognition and definition of habitat-types and it is

crucial that protected areas include representative portions of all habitat-types. Tropical rain

forests are rich in overall plant species but are rarely homogeneous: communities vary in

species composition forming a patchwork, or mosaic, that may seem deceivingly uniform to

the naive observer. All of the minor habitat-types at Lop6 provide important foods for gorillas

and chimpanzees and some, such as marshes, are crucial for gorillas during the dry season.

The savanna shrub Psidlzm guineensis produces fruit twice a year and gorillas, chimpanzees

and elephants arc attracted to the savannas to eat the fruit. Not only is it of interest to study

the behaviour of forest species in a very different habitat" but also visitors to the Reserve can

watch large mammals in ideal conditions. The Inpd savannas would be re-colonised by forest

if the gass was not burned annually. Thus, maintenance of part of the habitat (and species)

diversity at L,opE depends on active management to arrest a natural successional change.

RecommendationsThe value of systematic sampling of vegetation is obvious as it permits both qualitative

(presence versus absence) and quantitative (relative abundance) comparisons between sites.

Transects placed across the drainage give a representative sample of major habitat-types but

may include only small samples of some habitats (e.g. riverbanks or foresVsavanna interface).

These minor habilats may be of great interest and, if not sufficiently represented in transects'

should be sampled intensively using plots placed within each habitat-type. The enumeration

and identification of trees along transects and in sub-plots is time-consuming and difEcult but

is probably ttre best way to leam to recognise different plant species. Individual frees should

be labelled to allow long-term monitoring, checking of identification, and to act as a living

List of Plant Species Identified in the Lope Reserve 273

herbarium for new researchers. Non-specialists rarely have the taxonomic expertise to key out specimens with confidence, even if a local Flora is available; thus it is essential to collect good specimens that can be sent to specialists. We advise multiple collection and careful labelling of botanical specimens and the need for fertile material (preferably flowers). The Missouri Botanical Garden has produced a guide to plant collecting and specimen preparation that is useful. It is important to establish a good working relationship (good specimens in

exchange for rapid identifications) with National and International Herbaria as otherwise,

delays can be long and frustrating. It would be valuable if a list of taxomonic experts and

institutions interested in such collaboration with long-term field sites existed.

Finally, researchers interested in inter-site comparisons should continue their efforts to standardise methodology and to publish baseline data on ape foods and plant species lists

from their sites.

ACKNOWLEDGEMENTS We thank the L.S.B. Leakey Foundation, the World Wide

Fund for Nature, the Leverhulme Trust, NYZS-The Wildlife Conservation Society, the World Society for the Protection of Animals, The Royal Society, the Conder Conservation Trust, the Richard Brown Scholarship, the Conrad Zweig Trust, the Herbier de la Centre National de

Recherche Scientifique et Technique (Libreville) and especially the Centre International de Recherches Medicales de Franceville for financial and logistical support; also Alphonse Mackanga-Missandzou, Joseph Maroga-Mbina and the Direction de la Faune for assistance in

the Lope Reserve. This material is based, in part, upon work supported by: the ECOFAC

programme funded by the EEC (DGVIII) and managed by AGRECO-GEIE (LJTW & MF);

and the National Science Foundation and the Office of Forestry, Environment and Natural

Resources, Bureau of Science and Technology, of the U.S. Agency for International

Development under NSF Grant No. BSR-9024745 (GMcP). We are grateful to the following colleagues who contributed to data collection at the Lope: Stephanie Hall, Rebecca Ham,

Karen McDonald, Richard Parnell, Liz Rogers and Ben Voysey. We thank Patrick Blanc, EJ.

Breteler, David Harris, Mike Harrison, Philippe Hecketsweiler, Nicolas Halle, Annette Hladik, Carel Jongkind, Ard Louis, Nick Mundy, Frank White, and Chris Wilks for help with

botanical identifications.

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276

C. E. G. TUTIN. L. J. T. WHITE, E. A. WILLIAMSON, M. FERNANDEZ , G. MCPHERSON ガボン・口ぺ保護地域の同定された植物リスト

ガボン中央部に位置するロベ保護区 (the Lop� Reserve) の 'Station d'Etudes des Gorilles et

Chimpanz白 (SEGC)'におけるローランドゴリラ (Gorilla g. gorilla) とチンパンジー (Pan t.

troglodytes) に関する研究は、 1983 年に始まり現在も継続中である。本論文では、 50km2 の調査

地で確認できた 91 科 676 種の植物名をリストにして報告する。直径 10cm かそれ以上の木本の

データはライントランセクト法によりシステマティックに採集し、繁殖中の植物に関しては見つ

かれば吉己録するという方法をとった。各植物種について、生活形、ハピタツトの好み、そしてト

ランセクト上で記録した木本については密度を記載した。また、ゴリラやチンパンジーに食物を

提供する植物については、その探食部位を記載に入れた。植物種名リストは完全ではないが、

SEGC 調査地の植物の多様性は示している。記載した 7 つのハピタット・タイプはサバンナから

樹冠の閉じた森 (Closed Canopy Forest) までの幅があるが、調査地の主要な植生はクズウコン

科植物が林床に多い森林蜘1arantaceae Forest) である。ロベのゴリラとチンパンジーは幅の広い

食性を持ち、すべてのハピタット・タイプで植物から食物を得る。特定の季節には、いくつかの，

面積では狭い分布域しか有していない植物が多量の食物を供給する。アフリカの異なる調査地の

植生リストが明らかになって初めて、類人猿の採食対象の比較が可能になる。