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

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

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

List of Identified in the Northern Part of the Lope Reserve, *

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 belonging to 91 families that occur in the 50 sq. km study area. Data on 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 but the study area is dominated by 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 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 collecting and identifying plant species eaten by apes and running transects to describe the vegetation of the apes' habitat. Among the important food species of apes at Lop6 (Tutin & Fernandez, 1993; Williamson et aI., 1990) were two new species of (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, (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 of the enumeration of trees along ftansects. Analysis of these data has allowed the forest within the 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 in the ropical 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 data on 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, that occur in the SEGC study area at Lop6. For each species, the life-form and typical habitat is given and the parts eaten by gorillas or chimpanzees are listed. Density data are included for all species of tree with diameters at breast height (dbh) > 10 cm recorded on vegetation transects. 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 ape feeding ecology and of botanical inventories.

STUDY AREA

The Lopd Reserve covers 5,000 km2 and extends in a tapering rectangular shape from its northern boundary, the river Ogoou6 (0'3'S) to Mont Ibondji (l'10'S) between latitudes 11"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 numerous smaller hills and steep valleys. The Reserve includes a complex mosaic of vegetation types. In the north and east there are 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 be natural in origin (Aubreville, 1967; White,1992), dating from the Pleistocene, and active recolonisation 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 with savanna areas (stippled), the SEGC study area (hatched box) and the loca-tions of the five 5 km vegetation transects (Sites 1-5).

savannas are burnt each year for management purposes but some are protected by watercourses or by their inaccessibility. Within the forest block, areas adjacent to the savannas have discontinuous canopy cover and a dense unders0orey dominated by herbs of the Marantaceae and ZingSllrlraceae, classed as 'Marantaceae Forest' (Irtouzey, 1968). This forest type is thought to be of relatively recent origin, reflecting the re-colonisation of Pleistocene savannas (Aubreville, 1967; de Foresta, 1990; White,1992). Deeper into the forest block, canopy cover becomes continuous, plant species diversity increases and the density of herbaceous plants in the understorey decreases dramatically: this forest type is classed as 'Closed Canopy Forest'. Within these two major forest types there are smaller areas of permanent water, rocky outcrops, and areas where the vegetation has been affected by selective logging in the past. In each of these cases, plant species occur that are rare, or absent, elsewhere. The SEGC study area covers about 50 km2(0o10'S, l1o 35'E) of mainly Marantaceae Forest to the south and west of the savanna zone but extends into Closed Canopy Forest !o the west of the major mountain range. Parts of the study area were selectively logged between 1960-70. 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 dry setson 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 stip 50m 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 in lOm 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 of plants encountered in . 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 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 tle host 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 of Marantaceae 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 and trees are particularly common along edges although most also occur at lower densities 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 cm dbh 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 analysis and observation, see Tutin & Fernandez (1993) and Williamson er al. (199O) for details of methods.

RESULTS

Species Recorded Table I lists 676 plant species, belonging to 9l families, that occur in the SEGC study area in the lopd Reserve. Of these, 509 have been identified to species, 108 to and the remaining 59 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 were rare. was the best represented family in numerical terms, with 90 species, while 30 families were represented by a single species. At least two species (Mangifura indica and Citrus sp?) are cultivars. Their precise history is unknown but they are likely to be of ancient origin as from the 15th Century the River Ogoou6, which is the northem limit of the Lopd Reserve, was a trading route and the savanna zone has long been the site of human settlement. Young Citrus trees are not uncommon as gorillas 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 remaining mature seeds are subject to intense predation by bushpigs and rodents.

Life-forms

Of the 676 identifiedplants in Table 1,345 are trees, 113 herbs,55 fems,68 lianes and 63 shrubs. 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 dominant in two 5 km transects (Sites I & 5 in Figure l) compared to a single transect where Closed Canopy Forest (CCD predominated (Site 4). Thus, large tees (> 70 cm dbh) were enumerated in 50 hectares of MF compared to 25 hectares of CCF and all trees > 10 cm dbh were enumerated in 5 hectares of predominantly Marantaceae Forest compared to 2.5 hectares of Closed Canopy Forest. The diversity of tree species recorded in the transect sample in Marantaceae Forest and in Closed Canopy Forest in the SEGC study area was similar, with 129 and 146 species respectively. However, 130 additional species oftree were recorded on two 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 HABITAT3 DENS ITY (individualsTkmz)a EATEN2 > 10 cm dbh > 70 cm dbh Gorilla Chimp MF CCF MF CCF PTERIDOPHYTA Adiantum vogelii F MF(w) Arthropteris orientalis F R (ruF) Aspleniurn africanurn F R (}"tr) Asplenium emnrginatum F R(lr{F) Asplenium hemitomum F R(lvF) Asplenium jaundeense F R (MF) Asplenium variabile F w (ccF) Blotiella currori F w B olbitis acro stichoides F w(I!F) Bolbitis auriculata F w Bolbitis gabonensis F MF Bolbitis heudelotii F w Ceratopteris cornuta F W(lv[F) Christella dentata F w(I\F) Cyathea camerooniana Tf w Cyclosorus dentatus F w(lvF) D icranopteris linearis F S Diplazium sammntii F W (l'IF) Diplaziwn welwitschii F w (ccF) Lastreopsis currori F w Lomariopsis hederacea F (Ep) CCF Lomariopsis c ongoensis F CCF Lomariopsis sp. SEGC 536 F (Ep) CCF Lycopodium cernuurn F MF Lygodium microphyllum F F/S Marattia fraxinea F w M ic r o gr ammt ow ar ie ns i s F R (NF) Microlepia speluncae F w(MF) Microsoriwn punctatum F R (NF) Nephrolepis biserrata F F/S Nephrolepis undulata F F/S Oleandra distenta F R(l!F) Pellaea doniana F R Pellaea holstii F R (l'F)

P hynwto s o rus s c olopendria F (Ep) F/S;MF

P iryr o gr amm.e c alomelano s F s Platycerium stemnria F (Ep) MF Pteris atrovirens F S Pteris similis F w(MF) Pteris acanthoneuxa F MF Pteris sp. SEGC 528 F MF

S e I a g ine lla c athe dr ifo lia F w(Nlr') Selaginella myosurus F F/S Selaginella v ersic olor F w Selaginella vogelii F MF T ric ho nune s b al lardianum F w

I See Methods for definition of Life-forms; 2 L= ; 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 Ha; CCF > 10 cm dbh,2.5 Ha: MF >70 dbh, 50 Ha; CCF > 70 dbh, 25 Ha (see text). * indicates present as a tree of this size but not recorded in transect sample. List of Plant Species ldentified in the Lope Reserve 255

FAMILY / Species LIFE-FORM PLANT PART HABITAT DENS ITY (i ndividuals/km') EATEN > l0 cm dbh >70 cm dbh Gorilla Chimp MF CCF MF CCF

T ric homane s c upre s s o ide s F w Trichomanes guineense F R Trichomnnes cf. mannii F R T rip I op hy ll urn b uc hho lz ii F w T r ipl o p hy ll wn p r o t e ns um F MF T rip I op hy llwn s e c undifo rme F w Triplophyllwn sp. SEGC 469 F MF Triplophyllwn vogelii F CCF Vittaria guineensis F (Ep) CCF

ANGIOSPERMAE DICOTYLEDONEAE Acanthaceae Anisotes mocrophylla H F/S Arystasia gangetica H s ?DiclipteraUTW 0022 s MF; F/S Hypo estes vertic illaris H MF; D Justicia tenella H R(w) P s e uder ant hemum tunic atum H w LJTW 0748 s ccF (w) LJTW 0910 H MF(w) Anacardiaceae Antrocaryon klaineanwn T F F rF 40 Lannea welwitschii T B MF; R 2A 6 Mangtfera indica T F F D * P seudospondias longifulia T F F MF rF P seudo spondias mic roc arpa T F F w 60* Sorindeia gilletii T F F MF 40 Trichoscypha acuminata T F F F 460 r20 Trichoscypln abut T F F CCF 'F 200 Annonaceae Artabotry s ? rhopalocarpw L MF Artobotrys thomsonii L F/S C leistopholis staudtii T MF Erwntia chlorantln T F F F 500 200 Hexalobw crlspiflorus T F F CCF M o runtho taxis c o n go ens is CS F F R; F/S M onantho tatcis dec lina L CCF Monodora angolensis T R P ac hyp o dant hiwn s t audt ii T F/S :r P olyalthia sunveolens var. gabonica T F F 160 480 P olyalthia sunv eolens v at . s runv e olens T F F * Popowia declina L F MF Uvaria sp. SEGC 253 L F MF Uvaria sp. SEGC 256 CS F R Uvaria sp. SEGC 262 L F CCF U v ar ias tr wn p i e r r e anurn T F MF 260 Uvariodendron sp. LJTW 0706 S MF Xylopia aethiopica T F F MF; F/S; D 480 1440 Xylopia hypolampra T F F F M0 40 Xylopia parviflora T L F/S :1. Xylopia phloidora T MF 220 Xylopia quintasii T MF 2660 200 Xylopia cf. quintasii T MF * Xylopia staudtii T CCF 80 LJTW 0608 T MF 20 256 C. E. G. Turnr, L. J. T. Wrurn, E. A. Wn-uerrasoN, M. FEnNANDU & G. McHtnsoN

FAMILY / Species LIFE.FORM PLANT PART HABITAT DENS ITY (indi viduals/km2) EATEN > 10 cm dbh >70 cm dbh Gorilla Chimp MF CCF MF CCF

Apocynaceae Alstonia boonei T F Holarrhena floribunda T F/s :N( Landolphia cf . heudelotti L F/S; MF I-,andolphia sp. LJTW 0079 L D(w) Landolphia sp. SEGC 526 L F/S Picralima nitida T MF* Rauvolfia macrophylla T F/S * * Rauvoffia vomitoria T F/S * T ab e rnae mont ana c r as s a T F/S LJTW 0306 L MF LJTW 479 T CCF 40 Aristolochiaceae P araristolo c hia flo s - avis F Balanophoraceae Thoninngia sanguinea Pa F Balsaminaceae Impatiens sp. LJTW 0874 MF(w) Begoniaceae Begonia spp. (at least 2 species) CCF Bignoniaceae * Kigelia africana T F/S Newbouldia laevis T MF I2O Spathodea campanulata T MF* Bixaceae Bixa orellana F/S Bombacaceae Ceiba pentandra F/S 40 Cordia cf. millenii MF* Burseraceae T FL FL F; F/S 3140 640 570 3r2 Aucoumea klaineara **r1020 C anar ium s c hw e infur t hii T F F T F F 520 1160 72 272 Dacryodes buettneri :F Dacryodes edulis T F F 40 Dacryodes klaineana T F CCF * Dacryodes normandii T F F 190 Dacryodes sp. LJTW 591 T CCF 40 Santiria trimera T CCF 20 2160 Caesalpiniaceae * Afzelia bella T ccF (w) Afzelia bipindensfs T CCF :F Afzelia sp. LJTW 450 T CCF 4 Amphimas ferrugineus T CCF 80 8 Anthonotha ferruginea T CCF I2O 40 Ant hon o t ha mnc r op hy ll a T MF 160 * ?Anthonotha sp. LJTW 0641 T MF; F/S

Ap lwn o c aly x emnr g i ne rv at us T F/S *'! Augounrdia letestui T ccF 1600 * insignis T F/S Berlinia auriculata T w (ccFl r20 Berlinia bracteosa T w (lv{F) 1S0 8 Calpocalyx klainei T CCF 640 * Cassia mannii T F/S Cassia mimosoides H 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 HABITAT DENSITY (individuals&m2) EATEN > 10 cm dbh > 70 cm dbh Gorilla Chim MF CCF MF CCF Crudia gabonensis T CCF Cryptosepalum staudtii T S;L;BS; L;BF(W) Cynometra ?mannii r F/S (W) Daniellia klainei T CCF :F*2 4 Detariurn macrocarpum T S; L MF *10 Dialium ?bipindensrs T CCF * 8 Dialium dinklagei TFFCCF 160 rF Dialium lopense T F; S; LF;S;LF 280 160 2 r6 Dialiwn pachyphyllum TFFCCF 80 rF Dialium soyauxii T CCF I 160 4 Dialiwn sp. SEGC 525 T F/S D is temonant hus benthamianus T L L;FL F 40 40 12 4 D uparquetia orc hidac ea L F/S Erythrophleum ivorense T CCF 4 Eurypetalwn batesii T CCF 40 G ilbertiodendron dew evrei T CCF ,k* demewii T CCF 404 r w (ccF) 40 Guibourtia tessmannii TSF 208028 Hylodendron gabunense TSF 300 ,F 2g * Hymenostegia ?klainei r F/S (W) :r Hymeno s te g ia p e ll e gr inii T CCF 240 Julberrurdia brieyi T S;L; BS; L; B F/S;MF 206 Julberrardia seretii r ccF (w) rf Mezoneuron angolense L F/S N e o c hev alier o de ndron st ep hanii TF t20 40 P ar ab er I i nia b ifo li o lata T CCF *u P grini o rF elle de ndr o n dip hy I lwn r ccF (w) S c orodop hlo eus ze nkeri T ccF r20 r2 Sindoropsis le-testui T CCF 440 68 Swartzia fistuloides TLF 40* 10 ,F T e s smannia dew i lde manni ana r w (F/s) T e trab er I inia b ifo I io lat a T CCF rF LJTW 395 T CCF *g Celastraceae Ercnymus congolensis T R r20 Hippocratea myriantha L F/S; D Salacia mayumbensis L CCF Salacia sp. LJTW 0308 CS CCF Salacia sp. SEGC 412 CS L F/S Chrysobalanaceae Acioa sp. SEGC 499 T F/S C hrysobalanw icaco atacorensis S F/S (w) Magnistipula sp. LJTW 0701 T W(l,tr) Maranthes aubrevillei T F/S; F :F40*g Maranthes gabunensis T F 402002* Maranthes glabra T CCF 'F I20 'F 68 Parinari excelsa T CCF *g Combretaceae Combretum pecoense L CCF 40 Combretum platypterum L L F/S; MF C ombretuml P teleopsis sp. LJTW 0369 T MF Compositae

Ageratum sp. LJTW 0545 H S Aspilia africarn H s Bidens sp. LJTW 0215 H MF Bidens sp. LJTW 0544 H S 258 C. E. G. Turnq, L. J. T. WmrE, E. A. Wu-ueusoN, M. FENNANDU & G. MCHTENSON

FAMILY / Species LIFE.FORM PLANT PART HABITAT DENS ITY (individuals/km2) EATEN > l0 cm dbh > 70 cm dbh Gorilla Chimp MF CCF MF CCF

Launea sp. LJTW 0533 H S LJTW 0423 H MF LJTW 0494 H S; MF Connaraceae Agelaea paradoxa L D Agelaea pentagyna L F/S Agelaea sp. 0092 L CCF Cnestis corniculata CS F/S; F Cnestis ferruginea CS F/S; F Connarus grffinianus L F/S J olly dora dup arq uetianct s ccF Manotes mncrantha L D Rourea myriantha L CCF Rourea solanderi L MF Rourea tlnmsonii L F F/S LJTW 0344 L MF(w) LJTW 0479 S MF Convolvulaceae

I p omo e a b lep lwr op hy ll a H s Ipomoea involucrata H S Ipomoea mauritiana H D Hewittia scandens H S Cucurbitaceae Cogniauxia podolaena L F/S LJTW 0408 L MF Dichapetalaceae Dichapetalum barteri T F;L;BF R 320 40 Dichapetalurn sp. LJTW 0048 CS FFMF Dichapetalum sp. LJTW 0140 L D Dichapetalwn sp. LJTW 0414 L F F lvtF (W) Tapura bouquetiana T MF 340 Dilleniaceae :F Tetracera podotricha F/S Dipterocarpaceae ,F Marquesia excelsa w Ebenaceae * Diospyros abyssinica T FFMF Diospyros boala T CCF 40 Diospyros cinnabarina T CCF 40 Diospyros dendo T F;S F;S F(R) 1140 240 20 D io s py r o s kamerunens is T F {< Diospyros mannii T F;S F F 180 Diospyros melocarpa T CCF 240 Diospyros piscatoria T FF 160 80 Diospyros polystemon T F;S F;S F 1280 r20 T MF * Diospyros soyauxii *s Diospyros stumeolens T FFF 40 Diospyros viridicans T LF 60 40 Diospyros zenkeri T F;S F R 800 40 Diospyros sp. LJTW 409 T R 20 tl. Diospyros sp. LJTW 1004 T F/S Erythroxylaceae Erythrorylum mannii F/S 60 Alchornea cordifolia CS F/S Alclwrnea hirtella S CCF ?Anthostemmn sP. LJTW 0677 T w (lvtr) 20 List of Plant Species Identified in the Lope Reserve 259

FAMILY / Species LIFE-FORM PLANT PART HABITAT DENS ITY (individual s/km2) EATEN > 10 cm dbh > 70 cm dbh Gorilla Chi MF CCF MF CCF Antidesma laciniatwn T F FF 140 Antidesnu vogelianum T F F F/S; W; D *40 Bridelia fercuginea T s Bridelia sp. LJTW 0932 T F/S ?Cleistanthw sp. LJTW 546 T CCF 40 Croton sp. LJTW 0348 T MF; D rF r& D is co glypremne caloneura T F; L D 40 280 2 D uv i gne audia inop i rnt a T CCF 280 thymifolia H ?F s J atrop hn g o s sypiifolia S F/S Klaineanthw gaboniae T CCF 20 Macaranga barteri T D * 240 Macaranga gabunica L D Macaranga monctndra T D dusenii T CCF Maesobotrya cf . pynaertii T CCF 40 Manniophyton fulvum L F Maprounea membranacea T D 500 1000 micrantha T CCF t20 Mareyopsis longifolia T CCF 600 Phyllanthw diandrus T F ?Phyllanthw discoidew T F/S Plagiostyles africana T FF r20 720 2 R ic ino de ndro n he ude I o tii T CCF 80 Sapium ellipticutn T MF 202 Thecacoris sp. nov.? GMcP 16015 S MF Uapaca guineensis T F F F/S;MF(W) 20 l0 T F FF 20*2 Uapaca paludosa T F F CCF (W) Uapaca aff. togoensis T F F MF(W) 20 Uapaca vanhouttei T F F MF(W) * Uapaca sp. LITW 0993 T F/S (w) LJTW 526 T CCF 80 LJTW 567 T MF 20 LJTW 0658 s CCF LJTW 0661 L s (R) Caloncoba glauca T FL MF 20 Carnptostylus mannii T F 80 160 barteri T F r20 80 Homalium letestui T R 60 H omnlium ? aff . neur op hy llum T MF tlc Homalium sarcopetalwn T w (F/s) Hornaliwn sp. GMcP 16169 T CCF ,F Lindackeria dentata s D(r!F) Oncoba brachyantlrcra S F/S Scottellia coriacea T MF 620 Gentianaceae N eur othec a lo e se lioides s Guttiferae Allanblackia sp. LITW 272 T F **rrk afzelii T CCF 280 Garcinia cf kola T MF r|( G arc inia cf. ovalifolia T F/S (w) rF Garcinia sp. LJTW 152 T CCF 40 Garcinia sp. LJTW 0447 T CCF :1. Garcinia sp. LJTW 0767 T F/S (w) 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 HABITAT DENSITY (individuals/km2) EATEN > 10 cm dbh >70 cm dbh Gorilla Chimp MF CCF MF CCF Mammea africana TF FF :N( ,F Pentadesnwt buryracea T F; S F 60 'F P entade smt grandifolia T CCF 40 4 Symphonia globuliftra T F CCF 400 Humiriaceae Sac o glottis gabonensis TF F F/S 40 Hypericaceae Harungana sp. LJTW 0790 S s S P s o r o sp ermum fe b rifu g um S 80 P s o r o s p ermum tenuifo lium TL LD Vismia guineensis S D; MF 40 Icaciru mannii L MF Lasianthera africana TL F 40 160 D esborde sia glauc esc ens T F 160 480 l0 36 TF F; S F 220 240 20 4 Irttingia grardifolia TF F; S F 60 40 12 4 tl. * Irtingia robur TF F CCF * gabonensis TF FF 580 80 32 Klainedoxa trillesii TF FF 160 r20 6 20 Ixonanthaceae ,1. O c ht lw c o smus c on g ol ensis T F/S Labiatae Solenostemon sp. LJTW 0543 H S Lauraceae Beilschmiedia fulva T FF 40 40 Beilschmiedia sp. LJTW 37 T FMF 40 Beilschmiedia sp. LJTW 439 T ccF (w) 240 Beilschmiedia sp. LJTW 530 T CCF 240 Beilschmiedia sp. LJTW 537 T MF 20 Hypodaphnis zenkeri TL MF 560 {. Ocotea gabonensis T CCF N apoleonaea imperialis T F 40 r20 Napoleonaea cf . leonensis T CCF 160 t2 P e t er s ianthus mac r o c arP us T CCF Linaceae Hugonia planchonii L F/S Anthocleista vogelii TL W; D 40 40 :1. Ant ho c le i st a ? s c hw einfur thii T F/S Mostuea hirsuta S ccF (D) Mostuea sp. LJTW 0769 S F/S (w) Strychnos congolann L CCF Strychnos malacoclados L F/S Loranthaceae Helixanthera mannii Pa MF Lorantha.t sp. LJTW 0119 Pa CCF Tapinanthus sp. LJTW 0444 Pa CCF LJTW 0802 ?PA F/S LJTW 0918 Pa F/S LJTW 0979 Pa F/S Luxemburgiaceae Testulea gabonensis F 40 160 10 16 Malpighiaceae Acr ido c arp us lo n g ifu lius D List of Plant Species ldentified in the Lope Reserve 26r

FAMILY / Species LIT'E-FORM PLANT PART HABITAT DENS ITY (individuals&m2) EATEN > 10 cm dbh >70 cm dbh Gorilla Chimp MF CCF MF CCF Malvaceae Hibiscw sp. LJTW 0920 CS F/S Sida rhombifolia H MF Urena ?lobata S S Melastomataceae Calvoa monticola H D Dissotis congolensrs H S Tristemma leiocalyx H S LJTW 0023 H MF LJTW 0481 H D SEGC 319 LL MF Carapa procera T F 100 600 rF E nt and o p hr a gma c ando I ei T CCF 8 Entandophragma utile T CCF :N< * Lovoa trichilioides T F ** ,F ,F Trichilia monadelpha T SMF 40 Trichilia prieureana T MF 880 Menispermaceae SEGC 43I L F F/S Mimosaceae Albizia ferruginea T MF ?Calpocalyx sp. LJTW 0611 T CCF Cathormion altissimum T F/S Cylic odis cus gabunensis T F Dicrostachys cinera S S (D) Entada gigas L MF F illaeop sis disc ophora T F :k 4 Newtonia leucocarpa T F 2 8 Parkia bicolor T F FF 20 80 6 4 Parkia filicoidea T F FMF 20* 4 :F P entac lethra eeweldeana T L LF &0 240 t4 12 P entac lethra macr op hy lla T L L; FL F 700 440 36 44 P ip tadenias tr um afric anum T S F 40 :F 30 r6 Samnnea leptophylla T CCF 40 4 * Tetrapleura tetraptera T F 40 'lc Moraceae Dorstenia aff. barteri H w (ccF) Ficw barteri Ep FFMF Ficw bubu Ep FFF/S Ficw elasticoides Ep MF Ficw ingens T FMF Ficw kimuenzensis L CCF Ficus macrospertnct Ep FFMF Ficw mucuso T F;L F MF Ficw ovata T R (s) Ficw polita Ep FFMF Ficw pseudomangifera Ep F F/S Ficw recuryata Ep F; L F MF Ficw sur T R (S) Ficw thonningii Ep FFMF Ficw variifulia T L;B L MF Ficns sp. SEGC 443 Ep FFMF M ilic ia (C hlorophora) excelsa T L; B; G F/S; MF 80*64 Musanga cecropioides T D * Myrianthus arboreus T F; B F MF(W) * Treculia africarn T S S; L MF(w) * 262

FAMILY / Species LIFE-FORM PLANT PART HABITAT DENSITY (individualsftm2) EATEN > 10 cm dbh >70 cm dbh Gorilla Chimp MF CCF MF CCF Treculia obovoidea CCF 200 Myristicaceae Coelocaryon prewsi T CCF 360 * Pycnanthus angolensis T F F 140 r20 M28 S cyp ho c ep haliwn o c hoc oa T S;L CCF 20 920 6 r72 Staudtia gabonensis T F CCF * 200 ** Myrtaceae Psidium guineense s FF s Syzygiwn sp. LJTW 3l T MF ,1. 2 Syzygiurn sp. LJTW 0147 s CCF SyzygiwnlEugenia sp. LJTW 0381 T CCF Nymphaeaceae LJTW 0815 w (s) Ochnaceae

C ampy I o s p e r murn el o n g atwn T MF Lophira alata T F/S; MF 2740 'F 90 8 Ouratea flava T F/S; MF * Ouratea cf . myrioneura T F/S; MF ?Ouratea sp.LJTW 0432 T MF

:1. R lnb dop hy ll wn c al o p hy I I um T MF Sauvagesia erecta H s LJTW 0772 S CCF LJTW 0882 S S edulis T CCF 1760 60 Diogoa zenkeri T CCF *r Heisteria pan ifulia T F; L F F r20 24044 Ongokea gore T L F 60 80 14 12 P ty c ho p etalum p e t io lat wn T FF CCF 160 Strombosia pustulata T CCF 400 Strombosia zenkeri T CCF 2720 Stromb o siop sis tetrandra T CCF r20 2000 Oleaceae Linociera aff . mannii F/S Oxalidaceae Biophytum petersianum H S Biophytum talbotii H w(R) Pandaceae Centroplacw glaucinw T CCF 100 2720 T F 20'k** Papilionaceae Aganope impressa L CCF Aganope sp. L MF Crotularia pallida s S Dalbergia aff . rufa L L; B; FL F/S

D e s mo diurn ramo s is s imum H S Desmodiurn scorpiurw H S Eriosema glomeratum S s Erythrina tholloniana T w (NrF)

I ndigofera c onj ugata v ar . occ identalis H s Lep to derr is hyp ar gy re a L F/S Milleuia laurentii T CCF Millenia sanagana T L;B L F/S 40 Milleuia conraui T F/S Milleuia sp. LJTW 0038 T MF(w) 20 Platysepalum sp. LJTW 0603 L CCF 240 Pterocarpus soyawcii T S;L;FL S;I F 80 :rc 68 20 List of Plant Species Identified in the Lope Reserve 263

FAMILY / Species LIFE-FORM PLANT PART HABITAT DENS ITY (i ndividual s/km2) EATEN > l0 cm dbh > 70 cm dbh Gorilla Chimp MF CCF MF CCF Tephrosia purpurea H S Uraria picta H S Zornia latifulia H S LJTW 0469 S F/S LJTW 0591 L CCF 80 LJTW 0599 L CCF 40 LJTW 0,602 L ccF 40 LJTW 0609 T CCF 320 LJTW 0720 T ccF (w) LJTW 0950 T F/S :F SEGC 395 L MF fistulosa T F/S; MF; D 520 600 Paropsia grewioides T F/S; R LJTW 0360 L CCF Rhizophoraceae Anisophyllea sp. LJTW 0475 T CCF 80 4 Anopyxis klaineana T F * :1. ,F Cassipourea congensis T R 180 Poga oleosa T CCF Rubiaceae Aidia ochroleuca T F/S rF Atractogyne gabonii L MF; D Bertiera batesii S w (ccF) Bertiera mildbraedii T MF Bertiera sp. LJTW 0139 S CCF Borreria pusilla H s Brenania brieyi T MF Canthiwn sp. LJTW 0297 S CCF Cephaelr sp. LJTW 0202 s F/S Corynanthc mayumbensis T F 340 2240 Crossopteryx febrtfuga s S D ic ty andra arboresc ens T MF Diodia scandens H MF Feretia sp. LJTW 0646 s F/S ?Gaertnera sp. LJTW 0316 S CCF Geophila afzelii H MF Heinsia crinita T F(w) Leptactirn cf . arnoldiana T D Massularia acuminata T F 40 Mitacarpus scaber H S Mitragyru ciliata T B; L w 80 r20 t6 Mwsaenda cf . debeawii L F D Mussaenda cf . tenuitlora L D M ussaenda pp. LJTW 058 1 L MF Nauclea didderrichi T F F F 20 600 Nauclea latifolia T F F s Nauclea vandergrchtii T F F w 60 Oldenlandia corymbosa H S O lde nl and.i a I anc ifo I ia s s Otomeria guineezsis H MF Otomeria micrantha H s Oryanthus unilocularis T F/S * ?Oryanthus sp. LJTW 0903 T MF Pauridiantha aff . dewevrei T F/S * Pauridiantha efferata T D; F/S 220 P aur idiant lrc fl o r i bunda T D; F/S 100 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 HABITAT DENS ITY (individual s/km2) EATEN > 10 cm dbh > 70 cm dbh Gorilla Chimp MF CCF MF CCF Pausinystalia johimbe T F20 200 P aus iny stal ia mac ro c er as T MF 20 Pavetta hispida S L CCF Pavetta aff . viridiloba T L; FL MF 20 Pavetta sp. LJTW 0006 H CCF Pavetta sp. LJTW 0205 S F/S ?Pavetta sp. LJTW 536 T MF 40 Porterandia cladantha T F; L F D; MF 120 Pseudosabicea batesii H CCF P s e u^do s ab i c e a fl or ib unda L D P seudo sabic ea mildbraedii H D

P sy c ho tria p e dunc ulari s S MF Psychotria vogeliana s F/S Psychotria sp. LJTW 0012 T F/S :F Prychotria sp. LJTW 0l7l S CCF Psychotria sp. LJTW 0174 H CCF Psychotria sp. LJTW 0209 S MF Psychotria sp. LJTW 0638 S MF Psychotia sp. LJTW 0657 S CCF Rothmannia whitfteldii T MF Rutidea dupuisii L F Sabicea efulenensis L MF Sabicea mollis L MF

S ac o sp errna p anic ulatum L F/S Tarenna conferta T D Tarenna eketensis L F/S Tarenna sp. LJTW 0272 T CCF Tricalysia anomala T D

T ric aly s ia mnc r o p hy ll a T MF(W) ',rc Tricaly sia cf oligoneura S MF Tricalysia cf . pallens s MF Tricalysia sp. LJTW 0081 S ccF (w) Tricalysrc sp. LJTW 0082 S MF Trichostachys aurea H MF Uncaria africana L F/S LJTW 0098 T D LJTW 0372 s MF LJTW 0396 T CCF LJTW 0417 s MF LJTW 0433 S ccF LJTW 0628 T R; W (F/S) LJTW 0638 s MF LJTW 0642 T rrF (w) LJTW 0646 S F/S LJTW 0662 T S LJTW 140 T MF 40 LJTW I7O T D LJTW 387 T CCF 40 LJTW 514 T CCF 40 LJTW 528 T CCF 40 LJTW 566 T MF 20 SEGC 288 L F MF SEGC 291 L F D SEGC 508 s L MF Rutaceae Araliopsis sp. LJTW 353 T ccF 80 Citrus sp. SEGC 557 T F/S List of Plant Species Identified in the Lope Reserve 265

FAMILY/Species LIFE-FORM PLANTPART HABITAT DENslTY(individuals/km2) EATIN ) l0 cm dbh > 70 cm dbh Gorilla Chimp MF CCF MF CCF Fagara?macrophylla T F/S; MF !8 Fagara tessnunnii T CCF 40 Tecleaaff.verdooniana T R * Vepris cf. lonsii T R 20 Sapindaceae Blighiawelwitschii T MF r. Chytranthus talbotii T MF(W) Eriocoelwnmatocarpwn T F/S; MF 540 280 Eriocoelutn?paniculatwt T MF '* * Eriocoelwn sp. UTW 132 T MF n Ganophyllwn gigannrmt T F F MF 80 Z0 Lecaniodiscuscupanoides T F F MF 100 Placodiscuscf.opacus T CCF g0 Sapotaeae Baillonella toxisperma T F F CCF ,& 40 Z 4 Donellaogowewis T F F F/S {. Donclla?prmiformis T CCF g0 Gambeya africana T F F CCF * 40 !r g GambeyasubnudaTFFF*40** ?Lectomedoxn sp. LJTW 237 T MF * 2 Letestua dwissima T MF * 4 ManilkarafouilloyaruTFFCCFlF OmplalocarpwnprocerwnTF**:r* PachystelabrevipesTFFR** LITW 0228 T F F CCF 160 ,& Scrophulariaceae Cycniwt carnporurn H S Scytopetalaceae Scytopetalwt sp. IJTW 0017 T F F F l2O U0 Simaroubacrae Odyendyea gabonewis T CCF '* Qwssia gricana S F Solanaceae Schwenckia americana H S Sterculiaceae lizae T F F; L; p MF 7460 g Colanahoundensis T CCF (W) g0 Colasp. SEGC 332 T MF * Lepnnychia echinocarpa S MF Nesogordoniapapaverifera T F * * :r. '* Pterygota?bequaertii T CCF :* Scaplapenh,trnblackii T CCF 160 Sterculia tagacantln T L MF 40 2 Styracaceae Afrostyrax lepidopltyllus T CCF g0 Thymelaeaceae Dicranolepis buchlalzii S CCF ?Diuanolepis sp. LJTW S MF Octolepis decalepis S MF Tiliaceae Clappertonialicifulia H S; D Duboscia ma$ocarpa T F; S F; S F 40 l2O 2 * Triwrfetu cordifolia S S Ulmaceae Celtistessnannii T F;L F F * 200 g 2g Celtissp.LnW 444 T CCF 40 266 C. E. G. TUrNq, L. J. T. Wru.re, E. A. WnunusoN, M. FEnNANDEZ & G. MCHmnSoN

FAMILY / Species LIFE-FORM PLANT PART HABITAT DENS ITY (individuals/km2) EATEN > l0 cm dbh >70 cm dbh Gorilla Chimp MF CCF MF CCF Verbenaceae Vitex doniana T F F/S Vitex ?rivularis T CCF 80 Violaceae Rinorea ilicifulia S CCF Rinorea sp. LJTW 0055 T w (ccF) * Rinorea sp. LJTW 0132 T w(MF) Rinorea sp. LJTW 022I T w (ccF) 40 Rinorea sp. LJTW 0596 T w (ccF) r20 Rinorea sp. LJTW 0656 s CCF Rinorea sp. LJTW 0899 S CCF Vitaceae Cissru aff. barteri L D Cisszs dinklagei L F;L F F Cissns leonardi L F MF Vochysiaceae Erismadelphw exsul CCF 40

MONOCOTYLEDONEAE Agavaceae Dracaena fragans T CCF Dracaena sp. LJTW 0916 s F Sansevieria sp. SEGC 565 H R Amaryllidaceae Scadoxus cinnabarinus MF Araceae Anchomanes dffirmis H L;P F Nephytis sp. LJTW 0480 H w (ccF) Commelinaceae Aneilema beniniense H MF Commelina capitata H MF Palisota ambigw H MF Palisota sp. SEGC 551 H MF Cyperaceae Bulbostylis cf. densa H S Bulbostylis laniceps H S Fimbristylis pilosa H S Pycreus sp. LJTW 0526 H S Scleria boivinii H F/S; D LJTW 0131 H w(MF) LJTW 0407 H D(w) Gramineae Andr op o g o n fas t i g iatus H S Andr o p o g o n p s eudapr ic w H S Axonopus compressus H S Brachiaria jubata H S Centotheca lappacea H MF Hyparrhenia diplandra H s Olyra latifulia H MF Oplismenw hirtellus H MF Panicurn dregeanum H S Panicum grffinii H S Panicurn cf . walense H S Panicum sp. LJTW 0152 H w (ccF) Panicu.rn sp. LJTW 0208 H MF Pospalum paniculatwn H D (MF)

P ennis e twn po ly s tac hy on H S List of Plant Species Identified in the Lope Reserve 267

FAMILY / Species LIFE-FORM PLANT PART HABITAT DENS ITY (indi vidual s/km2) EATEN > 10 cm dbh > 70 cm dbh Gorilla Chimp MF CCF MF CCF

Peratis indica H S Pobeguinea arrecta H s S c hiz ac hy rium p I aty p hy ll um H s Sp or ob o lus py rarnidalis H S Streptogynn crinita H MF LJTW 0388 H S LJTW 0404 H MF(w) LJTW 0540 H S Liliaceae

Curculigo pilosa H S Marantaceae Ataenidia conferta H LLF Halopegia azurea H L; P w H aunwnia liebr e c hs tiana H S;L S;L F Hyps elodelphy s hirs uta H S;L S;L F Hyp s el o de lp hy s p o g ge ana H S;L S;L CCF Hyps elodelphy s vio lac ea H S;L S;L F M ar anto c hlo a c o rdifo li a H L;P w filipes H PF H L;P w M e gap hry niwn mac r o s t ac hy um H F;L;PF;T F Megaphrynium velutinum H F;L;PF;L F T rac hy p hry niwn braunianurn H PW Palmae Calamus sp. SEGC 554 P F/S; MF P F; P F/S; MF 180 Eremospatha cabrae P F; L; PF; P MF Podococcus barteri P CCF Raphia sp. SEGC 555 P w (s) Pandanaceae Pandanar sp. SEGC 556 w (s) Pontederiaceae H e t e rant he ra c all ifo lia W (I,F) Smilacaceae Smilax kraussiana F/S Zingiberaceae Afr amomum I o n g ip e ti o I at um H F; P F; P F/S Afrarnomum sp. ?nov SEGC 152 H F; P F; P MF Aframomum sp. SEGC 239 H F;P F;P F Afrarnomurn sp. SEGC 323 H F;P F;P CCF;D Costw afer H PPMF Renealmia cincinnata H PF Renealmia macrocolea H PPF 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 at Lop6 in terms of basal area and stem density

Species Family Rank (Basal Area) Rank (No. Stems) MF CCF MF CCF Aucoumea klaineana Burseraceae 1 2 Cola lizae Sterculiaceae 2 I Lophira alata Ochnaceae 3 3 P entaclethra mac rop hy lla Mimosaceae 4 9 Dacryodes buettneri Burseraceae 5 8 P entaclethra eetveldeana Mimosaceae 6 r0 Diospyros polystemon Ebenaceae 7 5 Hylodendron gabunense Caesalpiniaceae 8 Xylopia quintasii Annonaceae 9 Pycnanthus angolensis Myristicaceae 10 Trichilia cf . prieurearut Meliaceae

S trombosiop s is teff andra Olacaceae Diospyros dendo Ebenaceae Diospyros zenkeri Ebenaceae Maprounea membranacea Euphorbiaceae l0

S cyp hoceplnlium o c oc hoa Myristicaceae C entroplac us glaucinas Pandaceae I Xylopia aethiopica Annonaceae 7 Santiria trimera Burseraceae 9 3 Coula edulis Olacaceae 4 5 Augounrdia letestui Caesalpiniaceae 5 6 Sindoropsis le-testui Caesalpiniaceae 6 Strombosia zenkeri Olacaceae 8 Cy licodiscus gabonensis Mimosaceae 10 Dialium soyawcii 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 Number Percen Number Perce Marantaceae Forest r49 42 28.2 36 24.2 Closed Canopy Forest r52 r7 rt.2 2l 13.8 Forest 90 53 58.9 50 55.6 Forest/savanna edge 103 22 2r.3 l9 18.4 Savanna 67 3 4.5 2 3.0 Water 53 9 r7.0 2 3.7 Disturbed areas 34 6 17.6 4 I 1.8 Rocky areas 28 6 2r.4 6 21.4

TOTAL/AVERAGE 676 158 23.4 r40 20.7

ape feeding have been collected by faecal analysis and observation over ten consecutive years but new foods are still being discovered. Some of the newly recorded foods come from rare species, or from species which fruit irregularly, but others are parts of common species that are ealen only in unusual circumstances, e.g. when major fruit crops fail, or when apes range in atypical habitats such as savannas. Table 3 shows the vegetation types in which plants that provide foods for gorillas and chimpanzees primarily occur. Both gorillas and chimpanzees feed from plants in all of the habitat-types. Foods from species found exclusively in Closed Canopy Forest are undoubtedly under-represented as areas of CCF are on the edges of the study area and less time was spent collecting data on ape ecology in this vege0ation type. Few foods are provided by savanna species 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 those colonising disturbed areas, but gorillas find more foods from the flora associated with permanent 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 feed on herbaceous and woody plants that grow in standing water. This reflects the general avoidance ofwater by chimpanzees.

DISCUSSION

Species Recorded

The 676 species listed in Thble I show the flora of the SEGC study area to be diverse but the list is far from complete. The transect samples were biased with respect to habitat-type as most were located in Marantaceae Forest. This forest type is dominant in the north of the Reserve, close to the savannas but the more botanically divene Closed Canopy Forest is the predominant forest-type in the Reserve. This bias means that the species listed in Table I should not be considered as representative of the Lopd Reserve as a whole. To date, a total of 864 species have been recorded for the Reserve but the southem half remains botanically unexplored.

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 of Pteridophytes recorded (55 species) reflects two collecting rips by a specialist, but is far from complete (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 bark, or the febrifuga, Psidium guineensis, Bridelia ferruginea). These have fire-resistant 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 this gives trees with seeds that can germinate at the forest/savanna interface an enormous potential for reproduction. Tree species diversity at the interface is low but successional changes are clear with many young individuals of additional species appearing as soon as the edge colonisers provide shade. Large trees able to regenerate at, or close to, the forest edge at Lopd include Aucoumea klaineana, Lophira alata and Sacoglottis gabonensis. All these species can form almost pure stands, shading out Graminae and creating conditions that appear 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 listed as 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-habitat preference, growing almost exclusively on the banks of permanent streams. It seems likely that this reflects the post-Pleistocene colonisation of savannas, with the flora growing along streams being old gallery forest now surrounded by the recently established Marantaceae Forest.

Ape foods, Habitat-types and Inter-site Comparisons At Lop6, gorillas and chimpanzees have diverse diets and obtain some foods from plants in each of the habitat-types (Table 3). The sampling bias against CCF described above, applies equally to our data on ape foods as far less time was spent observing and tracking apes in this forest type than in MF. 'Forest' species are generalists that occur in both MF and in Closed Canopy Forest and 55-6OVo of the 90 species in this category provide food for the apes. These species, being widespread, though not always common, provide much food as gorillas and chimpanzees both travel widely to exploit rare food resources. The mosaic of habitat-types in the home ranges of our study groups at Lopd means that a wide range of food is available and perhaps explains why gorillas at Lop6 are the most frugivorous population of the genus studied to date (Williamson et al., 1990). Foods from the minor (in terms of area) habitats make important contributions to ape diet and use of the savanna edge by both gorillas and chimpanzees 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 season when fruit is scarce (Rogers et al.1988; Tutin e, al.,l99lb). Both gorillas and chimpanzees are known to show dietary flexibility and in different areas do not always eat the same foods even when the plants are available. For example, Nishida et al. (1983) found that only 597o of 286 available plant foods were eaten by chimpanzees at two sites, Gombe and Matrale in , although they are only 150 krn apart. Four of the 107 foods of lowland gorillas at Ndoki, Congo, listed by Nishihara (1992) occur ar Lop6 bur are not eaten by gorillas. Of these, tfuee (Pennclethra seeds, Polyalthia fruit and Tetrapleura seeds) are common within the SEGC study area Complete inter-site comparisons can only be made 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 still unknown 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 a rare 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 if inventory 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 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.

Recommendations The 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 ). 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.

REFERENCES

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Y., Sackett, G.P. & Thompson, K.R. (eds.), Topics in Prima to logy Volume 2: Behavior, Ecology and Conservation, 215-224. University of Tokyo Press, Tokyo. Nishida, T., Wrangham, RW., Goodall, J. & Uehara, S. 1983. Local differences in plant­ feeding habits of chimpanzees between the Mahale Mountains and Gombe National Park, Tanzania. Journal of Human Evolution 12: 467-480. Nishihara, T. 1992. A preliminary report on the feeding habits of western lowland gorillas (Gorilla g. gorilla) in the Ndoki Forest, northern Congo. In: Itoigawa, N., Sugiyama, Y., Sackett, G.P. & Thompson, K.R. (eds.), Topics in Primatology Volume 2: Behavior, Ecology and Conservation, 225-240. University of Tokyo Press, Tokyo. Norton-Griffiths, M. 1978. Counting Animals. African Wildlife Foundation, Nairobi, . Oslisly, R & Fontugne, M. 1993. La fin du stade neolithique et Ie debut de l'age du fer dans la moyenne vallee de l'Ogooue au Gabon. Problemes chronologiques et changements culturels. C. R. Acad. Sci. Paris Serie II 316: 997-1003. Reitsma, J. M. 1988. Vegetation Forestiere du Gabon. Forest Vegetation of Gabon. Tropenbos Technical Series 1, Ede, The Netherlands. Rogers, M. E., Williamson, E. A., Tutin, e. E. G. & Fernandez, M. 1988. Effects of the dry season on gorilla diet in Gabon. Primate Report 22: 25-33. Tutin, C. E. G. & Fernandez, M. 1985. Foods consumed by sympatric populations of Gorilla g. gorilla and Pan t. troglodytes in Gabon: Some preliminary data. International Journal of Primatology 6: 27-43. --& --1993. Composition of the diet of chimpanzees and comparisons with that of sympatric lowland gorillas in the Lope Reserve, Gabon. American Journal of Prima to logy 30: 195-211. --, --, Rogers, M. E., Williamson, E.A., McGrew, W.e. 1991b. Foraging profiles of sympatric lowland gorillas and chimpanzees in the Lope Reserve, Gabon. Phil. Trans. R. Soc. Lond. B 334: 179-186. --, Williamson, E. A., Rogers, M. E. & Fernandez, M. 1991a. A case study of a plant­ animal relationship:Cola lizae and lowland gorillas in the Lope Reserve, Gabon. Journal of Tropical Ecology 7: 181-199. White, L. J. T. 1992. Vegetation History and Logging Disturbance: Effects on Rain Forest Mammals in the Lope Reserve, Gabon (with Special Emphasis on Elephants and Apes). Ph.D. Thesis, University of Edinburgh. --, Tutin, C. E. G. & Fernandez, M. 1993. Group composition and diet of forest elephants, Loxodonta africana cyclotis , Matschie 1900, in the Lope Reserve, Gabon. African Journal of Ecology 31: 181-199. Williamson, E. A. 1988. Behavioural Ecology of Western Lowland Gorillas in Gabon. Ph.D. Thesis, University of Stirling. --, Tutin, e. E. G., Rogers, M. E. & Fernandez, M. 1990. Composition of the diet of lowland gorillas at Lope in Gabon. American Journal of Primatology 21: 265-277. Yamagiwa, J., Mwanza, N., Yumoto, T. & Maruhashi, T. 1992. Travel distances and food habits of eastern lowland gorillas: A comparative analysis. In: Itoigawa, N., Sugiyama, Y., Sackett, G.P. & Thompson, K.R (eds.), Topics in Primatology Volume 2: Behavior, Ecology and Conservation, 267-281. University of Tokyo Press, Tokyo. 276 276

C. 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 (SEG C)'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、 50km 2 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。

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