Part of the Lopé Reserve, Gabon - l'Observatoire des forêts d ...

IList of Plant Species Identified in the NorthernI Part of the Lopé Reserve, Gabon

ICaroline E.G. Tutin ^ 2, Lee J.T. White 3 * 4- 5, Elizabeth A. Williamson 6,

Michel Fernandez 2 > 5 and Gordon McPherson 7.

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Av1 Centre International de Recherche Médicales de Franceville, F

2 Department of Biological and Molecular Sciences, University of Stirling, Scotland.3 NYZS-The Wildlife Conservation Society, U.S.A.4 Institute of Cell, Animal and Population Biology, University of Edinburgh, Scotland.5 Programme de Conservation et Utilisation Rationelle des Ecosystèmes Forestièrs d’Afrique

Centrale (ECOFAC), Composante Gabon, (Projet FED, CCE DG VIII).6 Psychology Department, University of Stirling, Scotland.

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

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

Abstract

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

at the ‘Station d’Etudes des Gorilles et Chimpanzés' in the Lopé 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 tobe

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 Lopé 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.Introduction

Gabon straddles the equator on the west coast of Africa and approximately 80% of the

country’s area of 267,000 km^ 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. Hallé, 1987; Hallé & 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

country. Six published lists cover a total of 1,233 plant species (Hallé, 1964; 1965; Hallé &

Le Thomas, 1967; 1970; Hladik & Hallé, 1973; Florence & Hladik, 1980).Research at the Lopé Reserve, in central Gabon, began at the Station d’Etudes des Gorilles et

Chimpanzés (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 Lopé (Tutin & Fernandez, 1993; Williamson et al.,1990) were two new species of tree (Cola lizae [Sterculiaceae], N. Hallé, 1987 and Dialiumlopense fCaesalpiniaceael. 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,

enormous, leaves (Tutin et al., 1991a). 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 Lopé has included more systematic botanical collecting and an extension of the

enumeration of trees along transects. Analysis of these data has allowed the forest within theReserve to be classed into 20 types that differ both in structure and species composition(White, 1992).

Similar ecological research, focussing on great apes, is underway at several other sites inthe tropical forests of central Africa (e.g. 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 of the Lopé Reserve are available from several sources, but

publications 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 Lopé. 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 SEGCstudy area, but it provides the basis for inter-site comparisons both of ape feeding ecology and

of botanical inventories.Study Area

The Lopé Reserve covers 5,000 km^ and extends in a tapering rectangular shape from its

northern boundary, the river Ogooué (0°3’S) to Mont Ibondji (1°10’S) between latitudes11°17’-11°50’E. 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 1; 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

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 understorey dominated by herbs of the Marantaceae and

Zingiberaceae, classed as ‘Marantaceae Forest’ (Letouzey, 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 affectedby selective logging in the past.In

each of these cases, plant species occur that are rare, or absent, elsewhere.(Figure 1 about here)

The SEGC study area covers about 50 km^ (0°10’S, 11° 35’E) of mainly Marantaceae Forest

to the south andwest of the savanna zonebut extendsinto ClosedCanopy Forest to the west of the

major mountain range. Parts of the study area were selectively logged between 1960-70. A

single species, Aucoumea klaineana was extracted at an average density of 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 season of about 3 months

from mid-June to mid-September. Temperatures vary little over the year but are lowest

during the dry season when constant cloud cover during the daylight 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 (cf. Norton-Griffiths, 1978) in areas which had experienced different logging histories. Trees and lianes

>10 cm dbh were identified and measured in a strip 5m wide along each transect (providing a

sample of 2.5 ha in each area), and trees >70 cm dbh were similarly enumerated in a strip

50m wide (25 ha in each area). Figure 1 shows the location of the transects. 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 trees >10 cm dbh in 10m

wide strips along a line-transect 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, 1993), black colobus (Harrison & Hladik, 1986), grey-cheeked

mangabeys (Ham, in prep.), forest elephants (White et aL, 1993) and other species of largemammal (SEGC, unpublished data). Opportunistic botanical collections have been made of plantsencountered 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 WestTropical 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.Epiphytic fern (Ep): Pteridophyta that are rooted on tree trunks or branches.Tree fern (Tf): Pteridophyta with rhizome rising like a tree trunk.Palm (P): Members of Palmae family.Liane (L): Climbing woody plants that depend on other vegetation for support.Shrub (S): Woody plants, often with multiple stems that do not exceed 2m in height.Climbing shrub (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 (T): Woody plants, usually with single stems, growing to heights in excess of 2m.Epiphyte (Ep): Woody plants that require support of host early in life but may become free-standing

once the host tree dies.Parasite (Pa): Plants wholly, or partly, dependent on other plants for nutrients.

The habitat-types used here are broad categories combining vegetation types defined byWhite (1992) on the basis of species composition and structure and are defined as follows:Marantaceae Forest (MF): The canopy is discontinuous particularly in the middle storey allowing light

to penetrate to the forest floor. The understorey is characterised by dense herbaceous growth of

species of Marantaceae and Zingiberaceae.Closed Canopy Forest (CCF): Characterised by more continuous canopy cover, especially in the middle

layer (10-20m). Herbaceous growth in the understorey is sparse.Forest-savanna interface (F/S): This interface is clear-cut in areas where savannas are subject to

annua! burning 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 lower densities in Marantaceae Forest.Savanna (S): Areas dominated by Graminae with scattered fire-resistant shrubs.Rocky (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 closed, it is generally only 10-20 m high.Water (W): Marshes and the banks of permanent streams.Disturbed (D): Areas that have suffered disturbance from selective logging or road construction 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 et al. (1990) for details ofmethods.

ResultsSpecies Recorded

Table 1 lists 676 plant species, belonging to 91 families, that occur in the SEGC study areain the Lopé Reserve. Of these, 509 have been identified to species, 108 to genus and theremaining 59 to family. Each species which is not, as yet, fully determined is given a collectionnumber, which will be referred to in future publications and updates. For species recorded ontransects (i.e., those that reach at least 10 cm dbh), density data are included, but many specieswere rare. Rubiaceae was the best represented family in numerical terms, with 90 species,while 30 families were represented by a single species.

(Table 1 about here)At least two species (Manqifera 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 RiverOgooué, which is the northern limit of the Lopé 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 Manqifera does not appear to regenerate naturally: themajority of fruit are consumed by primates before they reach maturity and the remainingmature seeds are subject to intense predation by bushpigs and rodents.Life-forms

Of the 676 identified plants in Table 1, 345 are trees, 113 herbs, 55 ferns, 68 lianes and63 shrubs. Other life-forms 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 1 & 5 in Figure 1) compared to a single transect where ClosedCanopy Forest (CCF) predominated (Site 4). Thus, large trees (> 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, with 129and 146 species respectively. However, 130 additional species of tree were recorded on two 5km transects in CCF (Sites 2 & 3 in Figure 1) 35 km from the SEGC study area (White,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. In cases of multiple habitats, the

predominant one is marked first. Of the 676 plant species identified at least to family, 149occur principally in MF and 152 in CCF. No obvious habitat preferences were found for 90species (designated as ‘Forest’ (F) in Table 1) that occur both in MF and in CCF. One hundredand three species are found primarily close to the forest/savanna interface, and 67 (including23 Graminae) occur only in savannas. Smaller numbers of species were found to be associatedwith water (53); with areas disturbed by past logging activities (34); or with rocky outcropswithin the forest (28). If these species occur only within a specific habitat-type, this isindicated in Table 1, e.g. R (MF) indicates a species found close to rocky outcrops inMarantaceae Forest; and W (CCF), one associated with water in Closed Canopy Forest.

Table 2 compares the 10 commonest species in Marantaceae Forest and Closed Canopy Forestin terms of basal area and stem density. Only two tree species are amongst the top ten for basalarea 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 foresttypes in terms of species composition.

(Table 2 about here)Foods of Gorillas and Chimpanzees

Of the 676 species recorded in the SEGC study area, 27% provide food for eitherchimpanzees or gorillas, or both. Some families provided no, or few foods, e.g., Pteridophyte (0of 55 species) and Connaraceae ( 1 of 13 species) while other families provide food consistentlye.g., Marantaceae (12 of 12 species) and Moraceae (15 of 21 species). Data on ape feeding havebeen collected by faecal analysis and observation over ten consecutive years but new foods arestill being discovered. Some of the newly recorded foods come from rare species, or fromspecies which fruit irregularly, but others are parts of common species that are eaten only inunusual circumstances, e.g. when major fruit crops fail, or when apes range in atypicalhabitats such as savannas.

(Table 3 about here)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 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 vegetation type. Few foods are provided by savanna speciesbut a fifth of the species found at the forest/savanna interface provide food for apes. Bothgorillas 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 feed on

herbaceous and woody plants that grow in standing water. This reflects the general avoidance ofwater by chimpanzees.

DiscussionSpeciesRecorded

The 676 species listed in Table 1 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 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 diverse Closed Canopy Forest is the predominantforest-type in the Reserve. This bias means that the species listed in Table 1 should not beconsidered as representative of the Lopé Reserve as a whole. To date, a total of 864 species havebeen recorded for the Reserve but the southern half remains botanically unexplored.

Another source of bias is by life-form: Only trees and lianes with a dbh of 10 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 trips by a specialist, but is farfrom complete (N. Mundy, pers. comm.). Reitsma (1988), working in the Lopé Reserve 30 kmsouth-west of the SEGC study area, recorded 69 species of tree of dbh < 10 cm in a one hectareplot and 125 species in 0.02 hectares where all plants were enumerated. Of the speciesrecorded by Reitsma in the small plot, 24% were trees, 18% shrubs, 18% lianes and 40%herbs. This compares to 51% trees, 9% shrubs, 10% lianes and 25% herbs (including ferns)in our sample. An assumption of equivalent distribution of life-forms at the two sites at Lopéseems reasonable and this suggests that the species listed in Table 1 represent about half of thetotal diversity of the SEGC study area: If aM tree species have been identified the 345 speciesrepresents 24% of the total plant diversity, equal to about 1,400 species. Two decades ofbotanical collecting in the Makokou area of Gabon resulted in identification of 1 ,233 plantspecies (Florence & Hladik, 1980) giving support to the above extrapolation.

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

Sterculiaceae (Hallé, 1987) and Dialium lopense. Caesalpiniaceae (Breteler, in press). Atleast 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 (Berg et al., 1984); and of the 16 species ofDiospyros (Ebenaceae) two are new records for Gabon (Letouzey & White, 1970), including ITpolystemon which is the fifth commonest tree, in terms of stem density, in Marantaceae Forest(Table 2).

Habitat-PreferencesMost plant species were found to occur only, or in much greater abundance, in one particular

habitat-type (see Table 1). Differences 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 latifolia, Crossopteryx febrifuqa.Psidium quineensis. Bridelia ferruqinea). These have fire-resistant bark, or the capacity for

rapid re-growth from root stock after the annual burning. In the savanna zone, gallery forests

grow 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 altissimum. Aphanocalyx cvnometroides. Pachystela

brevipes) and some, generally rare MF species become common (e.g. Uapaca quineensis.Pseudospondias microcarpa andFicus spp.).

Savannas protected from annual burning at Lopé 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 colonisersprovide shade. Large trees able to regenerate at, or close to, the forest edge at Lopé include

Aucoumea klaineana. Lophira alata and Sacoqlottis qabonensis. 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 pierreanum. Xvlopia spp..Klainedoxa spp.

Of the tree species in Table 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). Scyphocephalium ochocoa andStaudtia qabonensis (Myristicaeae) are listed as

CCF species and the two former are amongst the commonest ten species in this habitat (see Table2). 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 likely thatthis 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 Lopé, 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 55-60% 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 Lopé means that a wide range of food is available andperhaps explains why gorillas at Lopé are the most frugivorous population of the genus studiedto date (Williamson et al., 1990). Foods from the minor (in terms of area) habitats makeimportant contributions to ape diet and use of the savanna edge by both gorillas and chimpanzeesis highly seasonal and clearly related to fruiting patterns of F/S species (unpublished data).Similarly, aquatic herbs particularly Marantochloa cordifolia (Marantaceae) provideimportant keystone foods for gorillas at Lopé during the dry season when fruit is scarce (Rogerset al. 1988; Tutin et al., 1991b).

Both gorillas and chimpanzees are known to show dietary flexibility and in different areas donot always eat the same foods even when the plants are available. For example, Nishida et al.(1983) found that only 59% of 286 available plant foods were eaten by chimpanzees at two

sites, Gombe and Mahale in Tanzania, although they are only 150 km apart. Four of the 107foods of lowland gorillas at Ndoki, Congo, listed by Nishihara (1992) occur at Lopé but are not

eaten by gqrillas. Of these, three (Pentaclethra seeds,Polvalthia fruit and Tetrapleura seeds)are common within the SEGC study area. Complete inter-site comparisons can only be made onceboth lists of ape foods and plant species lists are published for each of the long-term study sites.What determines which plants apes eat, and which they do not, is still unknown but comparisonsbetween sites will clarify ecological differences (e.g. total plant 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 rarespecies from the food list is less noteworthy than that of a common species. For example,Musanqa cercropioides (Moraceae) is eaten by apes elsewhere in Gabon (Hladik, 1973; Tutin &Fernandez, 1985) but has never been recorded as an ape food at Lopé. However, while Musanqa

is a common colonising tree in most of Gabon, it is extremely rare in the SEGC study area withonly two trees known in 50 sq. km. In contrast, Pentaclethra spp. are common with an average

of 1,300 trees per km^ in Marantaceae Forest at Lopé,yet gorillas do not eat the seeds. An

importance rating for foods (Tutin & Fernandez, 1993) is also useful when makingcomparisons between sites, as consumption of rare foods can easily be overlooked whiledifferences in important, or keystone, foods indicate real ecological or cultural differencesbetween populations. Inter-site comparisons will allow a deeper examination of nutrition andfood choice than has yet been possible. Analysis of the importance of factors such as inter-specific competition and the determinants of population density will also be possible and shouldadvance 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 theirextraordinary 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 arelegion 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 thenaive observer. All of the minor habitat-types at Lopé provide important foods for gorillas andchimpanzees and some, such as marshes, are crucial for gorillas during the dry season. Thesavanna shrub Psidium quineensis produces fruit twice a year and gorillas, chimpanzees andelephants are attracted to the savannas to eat the fruit. Not only is it of interest to study thebehaviour of forest species in a very different habitat, but also visitors to the Reserve can

watch large mammals in ideal conditions. The Lopé savannas would be re-colonised by forest ifthe grass was not burned annually. Thus, maintenance of part of the habitat (and species)

diversity at Lopé 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 butmay include only small samples of some habitats (e.g. riverbanks or forest/savanna interface).These minor habitats 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

2

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

probably the best way to learn to recognise different plant species. Individual trees should belabelled to allow long-term monitoring, checking of identification, and to act as a livingherbarium 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 goodspecimens that can be sent to specialists. We advise multiple collection and careful labelling ofbotanical specimens and the need for fertile material (preferably flowers). The MissouriBotanical Garden has produced a guide to plant collecting and specimen preparation that isuseful. It is important to establish a good working relationship (good specimens in exchange forrapid identifications) with National and International Herbaria as otherwise, delays can be longand frustrating. It would be valuable if a list of taxomonic experts and institutions interested insuch 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 fromtheir sites.

Acknowledgements

We thank the L.S.B. Leakey Foundation, the World Wide Fund for Nature, the LeverhulmeTrust, NYZS-The Wildlife Conservation Society, the World Society for the Protection ofAnimals, The Royal Society, the Conder Conservation Trust, the Richard Brown Scholarship, theConrad Zweig Trust, the Herbier de la Centre National de Recherche Scientifique et Technique(Libreville) and especially the Centre International de Recherches Médicales de Franceville forfinancial and logistical support; also Alphonse Mackanga-Misandzou, Joseph Maroga-Mbina andthe Direction de la Faune for assistance in the Lopé Reserve. This material is based, in part,upon work supported by: the ECOFAC programme funded by the EEC (DGVIII) and managed byAGRECO-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 forInternational Development under NSF Grant No. BSR-9024745 (GMcP). We are grateful to thefollowing colleagues who contributed to data collection at the Lopé: Stephanie Hall, Rebecca Ham,Karen McDonald, Richard Parnell, Liz Rogers and Ben Voysey. We thank Patrick Blanc, F.J.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 botanicalidentifications.

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r •• •

m «•«• • «

« •«• • •• • •• 9«« • • • « «•«

River Ogoouô•«« • • 9 9

« ••« V

4NSite 4te 1v

Site S

tt\ Site 3 / Site 2\\

\

\t\

\\\\

vv\i\

T\12°v

V\ CAMEROUN

10 km

Figure 1. The Lopé Reserve with savanna areas (stippled), the SEGC study area (hatched box)and the locations of the five 5 km vegetation transects (Sites 1-5).

Table 1. Plant species identified in the SEGC study area in the Lopé Reserve

HABITAT 3 DENSITY (individuals/km2) 4> 10 cm dbh > 70 cm dbh

MF CCF

LIFE-FORM 1 PU\NT PART EATEN 2Gorilla Chimp

FAMILY / Species

MF CCF

ACANTHACEAEAnisotes macrophvllaAsystasia qanqeticaTDicliptera UTW 0022Hvooestes verticillarisJusticia tenellaPseuderanthemum tunicatumLJTW 0748UTW 0910

AGAVAŒAEDracaena fraqansDracaena sp. LJTW 0916Sansevieria sp. SEGC 565

AMARYLLIDACEAEScadoxus cinnabarinus

ANACARDIACEAEAntrocarvon klaineanumLannea welwitschiiManqifera indicaPseudospondias lonqifoliaPseudospondias microcarpaSorindeia qilletiiTrichoscypha acuminataTrichoscypha abut

ANNONACEAEArtabotrvs ? rhopalocarpusArtobotrvs thomsoniiCleistopholis staudtiiEnantia chloranthaHexalobus crispiflorus

F/SLLHSHMF; F/SMF; DR (W)

SHH

WHCCF (W)MF (W)

SH

*CCFTFSRH

MFH

* 40F F 4FT20 6MF; RBT* *DF FT+F MFFT60 •k *w 2FFT40MFFFT460 120F FFT* 200CCFF FT

MFLF/SLMFT

500 200FF FTCCFF FT

HABITAT 3 DENSITY (individuals/km2) 4

> 10 cm dbh > 70 cm dbhMF CCF MF CCF

LIFE-FORM 1 PLANT PART EATEN 2

Gorilla ChimpFAMILY / Species

R; F/SCS F FMonanthotaxis conqoensisMonanthotaxis déclinaMonodora anqolensisPachypodanthium staudtiiPolvalthia suaveolens var. qabonicaPolvalthia suaveolens var. suaveolensPopowia déclinaUvaria sp. SEGC 253Uvaria sp. SEGC 256Uvaria sp. SEGC 262Uvariastrum pierreanumUvariodendron sp. LJTW 0706Xvlopia aethiopicaXvlopia hypolampraXvlopia parvifloraXvlopia phloidoraXvlopia quintasiiXvlopia cf. quintasiiXvlopia staudtiiLJTW 0608

APOCYNACEAEAlstonia booneiHolarrhena floribundaLandolphia cf. heudelottiLandolphia sp. LJTW 0079Landolphia sp. SEGC 526Picralima nitidaRauvolfia macrophvllaRauvolfia vomitoriaTabemaemontana crassa

CCFLRF FTF/S *T

160 480FFT*FFT

F MFLLMFFL

CS F RFCCFFLMF 260F FTMFSMF; F/S; D 480 1440

440 40FFT

2F FT FF/S *T LMF 220T

2660 200F MFT F*MFT

CCF 80T20MFT

* *FT*F/ST

F/S; MFD (W)

FL FL

F/SFL F*T MF

F/S *TF/STF/ST

F MFFLJTW 0306LJTW 479

L40CCFT

HABrTAT 3 DENSITY (individuals/km2) 4> 10 cm dbh > 70 cm dbh

MF CCF MF CCF


FAMILY / Species

ARACEAEAnchomanes difformisNephvtis sp. LJTW 0480

ARISTOLOCHIACEAEPararistolochia flos-avis

BALANOPHORACEAEThoninnqia sanquinea

BALSAMINACEAEImpatiens sp. LJTW 0874

BEGONIACEAEBegonia spp. (at least 2 species)

BIGNONIACEAEKiqelia africanaNewbouldia laevisSpathodea campanulata

BIXACACEAEBixa orellana

BOMBACACEAECeiba pentandra

BORAGINACEAECordia cf. millenii

BURSERACEAEAucoumea klaineanaCanarium schweinfurthiiDacrvodes buettneriDacrvodes edulisDacrvodes klaineanaDacrvodes normandiiDacrvodes sp. LJTW 591Santiria trimera

L; P FHW (CCF)H

FL

FPa

MF (W)H

CCFH

F/S *T120MFT*MFT

S F/S

F/S 40 6T

MF +T

F; F/S 3140 640 570 31210 20

520 1160 72 272

FLT FL**F FT

F FT* 40F FT

F CCFT•k180 *F FFT

40CCFT20 2160F CCFT F

CAESALPINIACEAE




FAMILY / Species

Afzelia bellaAfzelia bipindensisAfzelia sp. UTW 450Amphimas ferruqineusAnthonotha ferruaineaAnthonotha macrophvlla? Anthonotha sp. LJTW 0641Aphanocalvx emarqinervatusAuqouardia letestuiBaikiaea insiqnisBerlinia auriculataBerlinia bracteosaCalpocalvx klaineiCassia manniiCassia mimosoidesCassia obtusifoliaCopaifera mildbraediiCrudia qabonensisCryptosepalum staudtiiCvnometra ?manniiDaniellia klaineiDetarium macrocarpumDialium ?bipindensisDialium dinklaqeiDialium lopenseDialium pachyphvllumDialium sovauxiiDialium sp. SEGC 525Distemonanthus benthamianusDuparquetia orchidaceaErvthrophleum ivorenseEurypetalum batesii

CCF (W)TCCFT *CCF *T 4CCFT 80 8CCF 120T

T MF 160 40MF; F/S *TF/STCCF 1600T *F/STW (CCF)W (MF)

T 120T L 180 8

CCFT 640T F/S

SHCS F/S

CCFT **T CCF 4

F (W)F/S (W)

S; L; BT S; L; B * *TT CCF ** 2 4T S; L MF ic 10T CCF 8T F CCFF 160 *

F; S; L F; S; L 280 160T F 2 16CCF 80T F FCCF 1160T 4F/ST

L; FL 40 40T L F 12 4F/SL

T CCF 4CCFT 40



LIFE-FORM 1 PLANT PART EATEN 2Gorilla Chimp

FAMILY / Species

CCF •k *Gilbertiodendron dewevreiGuibourtia demeusiiGuibourtia ehieGuibourtia tessmanniiHvlodendron gabunenseHvmenosteqia ?klaineiHvmenosteqia pelleqriniiJulbernardia brieviJulbernardia seretiiMezoneuron anqolenseNeochevalierodendron stephaniiParaberlinia bifoliolataPelleqriniodendron diphvllumScorodophloeus zenkeriSindoropsis le-testuiSwartzia fistuloidesTessmannia dewildemannianaTetraberlinia bifoliolataUTW 395

CELASTRACEAEEuonvmus conqolensisHippocratea mvrianthaSalacia mayumbensisSalacia sp. UTW 0308Salacia sp. SEGC 412

CHRYSOBALANACEAEAcioa sp. SEGC 499Chrvsobalanus icaco atacorensisMaqnistipula sp. UTW 0701Maranthes aubrevilleiMaranthes qabunensisMaranthes glabra

T40CCFT 4

W (CCF)T 40S 20 80

300 *T F 2 8T S 28F

F/S (W)T *240CCFT

F/S; MFCCF (W)

20T S; L; B S; L; B 6T *

F/SL120 40T F

CCF *T 24CCF (W) *TCCF 120T 1 2CCF 440T 68

40T L F 10W (F/S)TCCFT *CCFT 8

120RTF/S; DLCCFLCCFCSF/SCS L

F/STF/S (W)W (MF)F/S; F

ST

* 40 k 8TF 40 200 2 *T

* 120 *CCF 68T

DENSITY (individuals/km2) 4


LIFE-FORM 1 PUNT PART EATEN 2 HABITAT 3Gorilla Chimp

FAMILY / Species

* 8CCFParinari excelsaCOMBRETACEAE

Combretum pecoenseCombretum platypterumCombretum/Pteleopsis sp. LJTW 0369

COMMELINACEAEAneilema beninienseCommelina capitataPalisota ambiguaPalisota sp. SEGC 551

COMPOSITAEAgératum sp. LJTW 0545Aspilia africanaBidens sp. LJTW 0215Bidens sp. LJTW 0544Launea sp. LJTW 0533LJTW 0423LJTW 0494

CONNARACEAEAqelaea paradoxaAqelaea pentaqynaAqelaea sp. 0092Cnestis corniculataCnestis ferruqineaConnarus qriffonianusJollvdora duparquetianaManotes macranthaRourea mvrianthaRourea solanderiRourea thomsoniiLJTW 0344LJTW 0479

T

40CCFLF/S; MFL L

* *MFT

MFHMFHMFPHMFH

SHSHMFHSHSHMFHS; MFH

DLF/SLCCFLF/S; FF/S; F

CSCS

F/SLCCFSDLCCFLMFLF/SFLMF (W)LMFS

HABrTAT 3 DENSITY (individuals/km2) 4> 10 cm dbh > 70 cm dbh

MF CCF MF CCF


FAMILY / Species

CONVOLVULACEAEIpomoea blepharophvllaIpomoea involucrataIpomoea mauritianaHewittia scandens

CUCURBITACEAECoqniauxia podolaenaLJTW 0408

CYPERACEAEBulbostvlis cf. densaBulbostvlis lanicepsFimbristvlis pilosaPvcreus SP. LJTW 0526Scleria boiviniiLJTW 0131LJTW 0407

DICHAPETALACEAEDichapetalum barteriDichapetalum sp. LJTW 0048Dichapetalum sp. LJTW 0140Dichapetalum sp. LJTW 0414Tapura bouquetiana

DILLENIACEAETetracera podotricha

DIPTEROCARPACEAEMarquesia excelsa

EBENACEAEDiospyros abvssinicaDiospvros boalaDiospyros cinnabarinaDiospvros dendoDiospvros kamerunensis

SHH SH D

SH

F/SLMFL

SHSHSHSHF/S; DW (MF)D (W)

HHH

320 40F; L; B F RTCS F F MFL D

MF (W)L F F340T MF

F/S *L

W * *T

F F MF *TCCF 40TCCF 40TF (R) 1140 240F; S F; sT

20FT

HABrrAT 3 DENSITY (individuals/km2) 4> 10 cm dbh > 70 cm dbh

MF CCF MF CCF


FAMILY / Species

Diospyros manniiDiospyros melocarpaDiospyros piscatoriaDiospyros polvstemonDiospyros sovauxiiDiospyros suaveolensDiospyros viridicansDiospyros zertkeriDiospyros sp. LJTW 409Diospyros sp. LJTW 1004

ERYTHROXYLACEAEErvthroxylum mannii

EUPHORBIACEAEAlchomea cordifoliaAlchomea hirtella? Anthostemma sp. LJTW 0677Antidesma laciniatumAntidesma voqelianumBridelia ferrugineaBrideliasp. UTW 0932TCIeistanthus sp. LJTW 546Croton sp. LJTW 0348Discoqlypremna caloneuraDuviqneaudia inopinataEuphorbia thvmifoliaJatropha gossvpiifoliaKlaineanthus qaboniaeMacaranqa barteriMacaranqa qabunicaMacaranqa monandraMaesobotrya duseniiMaesobotrya cf. pynaertii

F; S F F 180 *TCCF 240TF 160 80

1280 120T F

F; ST F; S F 2MF *T

F F * 40T FF 60 40T L

800 40F; S RT F20RT

F/S •kT

F/S 60T

CS F/SS CCF

W (MF) 20TF FT F 140

*F F/S; W; D 40T FT S

F/STT CCF 40T MF; D * *T 40 280 2B F; L D

CCF 280T? F SH

S F/SCCF 20T

T * 240DL DT D *

CCF *TCCF 40T

PLANT PART EATEN 2 HABITAT 3 DENSITY (individuals/km2) 4


1FAMILY / Species LIFE-FORMGorilla Chimp

Manniophvton fulvumMaprounea membranaceaMareva micranthaMarevopsis lonqifoliaPhvllanthus diandrus?Phvllanthus discoideusPlaqiostvles africanaRicinodendron heudelotiiSapium ellipticumThecacoris sp. nov.? GMcP 16015Uapaca quineensisUapaca heudelotiiUapaca paludosaUapaca aff. toqoensisUapaca vanhoutteiUapaca sp. LJTW 0993LJTW 526LJTW 567LJTW 0658LJTW 0661

FLACOURTIACEAECaloncoba qlaucaCamptostvlus manniiCasearia barteriHomalium letestuiHomalium ?aff. neurophvllumHomalium sarcopetalumHomalium sp. GMcP 16169Lindackeria dentataOncoba brachvantheraScottellia coriacea

GENTIANACEAE

L S F500 1000T D

120T CCFCCF 600T

T FF/ST

120 720T F F F 2CCF 80TMF 20T 2

S MFF/S;MF(W) 20 10T F F

20T F * 2 4F FCCF (W)MF (W)MF (W)F/S (W)

T F F20T F F*T FF

TCCF 80T

20MFTCCFSS (R)L

MF 20T FL80F 160T120 80FT60RT*MFT

W (F/S)TCCFTD (MF)SF/SS L

620MF 4T

DENSITY (individuals/km^) 4


LIFE-FORM 1 PLANT PART EATEN 2 HABITAT 3Gorilla Chimp

FAMILY / Species

SHNeurotheca loeselioidesGRAMINAE

Andropoqon fastiqiatusAndroooqon pseudapricusAxonopus compressusBrachiaria iubataCentotheca lappaceaHvparrhenia diplandraOlvra latifoliaQplismenus hirtellusPanicum dreqeanumPanicum ariffaniiPanicum cf. walensePanicum sp. LJTW 0152Panicum sp. LJTW 0208Paspalum paniculatumPennisetum polvstachvonPeratis indicaPobequinea arrectaSchizachvrium platyphyllumSporobolus pyramidalisStreptoqyna crinitaLJTW 0388LJTW 0404LJTW 0540

GUTTIFERAEAllanblackia SP. LJTW 272Garcinia afzeliiGarcinia cf kolaGarcinia cf. ovalifoliaGarcinia sp. LJTW 152Garcinia sp. LJTW 0447

SHSHSHSHMFHSHMFHMFHSHSHSHW (CCF)HMFHD (MF)HSHSHSHSHSHMFHSHMF (W)HSH

•k* * *FTCCF 280FT

*MF icTF/S (W) •kTCCF 40TCCF *T

r

LIFE-FORM 1 PUNT PART EATEN 2Gorilla Chimp


> 10 cm dbh > 70 cm dbhFAMILY / Species

MF CCF MF CCF

Garcinia sp. LJTW 0767Mammea africanaPentadesma butvraceaPentadesma qrandifoliaSvmphonia qlobulifera

HUMIRIACEAESacoalottis qabonensis

HYPERICACEAEHarunqana sp. LJTW 0790Psorospermum febrifuqumPsorospermum tenuifoliumVismia quineensis

ICACINACUEIcacina manniiLasianthera africana

IRVINGIACEAEDesbordesia qlaucescensIrvinqia qabonensisIrvinqia qrandifoliaIrvinqia roburKlainedoxa qabonensisKlainedoxatrillesii

IXONANTHACEAEOchthocosmus conqolensis

UBIATAESolenostemon sp. LJTW 0543

UURACEAEBeilschmiedia fulvaBeilschmiedia SP. LJTW 37Beilschmiedia sp. LJTW 439Beilschmiedia sp. LJTW 530Beilschmiedia sp. LJTW 537

T F/S (W)T F F F 'k *T F; S 60F 'k *T CCF 40 4T F CCF 400

T F F F/S 40

S SS ST L L D 80S D; MF 40

L MF160T L F 40

T F 160 480220 240

10 36T F; SF F 20 4

12 4T F; SF F 60 40CCFT F F *

80T F F F 580 32 *120T F F F 160 6 20

T F/S

H S

T F F 40 40T F MF 40

CCF (W)T 240CCFT 240

T MF 20

DENSITY (individuals/km^) 410 cm dbh > 70 cm dbh


FAMILY / Species

MF CCF MF CCF

560MFT LHvpodaphnis zenkeriQcotea qabonensis

LECYTHIDACEAENapoleonaea imperialisNapoleonaea cf. leonensisPetersianthus macrocarpus

LILIACEAECurculiqo pilosa

LINACEAEHuqonia planchonii

LOGANIACEAEAnthocleista voqeliiAnthocleista TschweinfurthiiMostuea hirsutaMostuea sp. UTW 0769Strvchnos conqolanaStrvchnos malacoclados

LORANTHACEAEHelixanthera manniiLoranthus sp. UTW 0119Tapinanthus sp. UTW 0444UTW 0802UTW 0918UTW 0979

LUXEMBURGIACEAETestulea qabonensis

MALPIGHIACEAEAcridocarpus lonqifolius

MALVACEAEHibiscus sp. UTW 0920Sida rhombifoliaUrena ?lobata

*CCFT

40 120T FCCFTCCF 160 12T

SH

F/SL

W; D 40 40LTF/S *TCCF (D)F/S (W)

SS

CCFLF/SL

MFPaCCFPaCCFPaF/S?PaF/SPaF/SPa

40 160L F 10 16T

DT

CS F/SMFH

S S

LIFE-FORM 1 PLANT PART EATEN 2 HABTAT 3 DENSITY (individuals/km2) 4


10 cm dbh > 70 cm dbhMF CCF MF CCF

MARANTACEAEAtaenidia confertaHalopeqia azureaHaumania liebrechstianaHvpselodelphvs hirsutaHypselodelphvs poqqeanaHvpselodelphvs violaceaMarantochloa cordifoliaMarantochloa filipesMarantochloa purpureaMeqaphrvnium macrostachvsMeqaphrvnium velutinumTrachvphrvnium braunianum

MELASTOMATACEAECalvoa monticolaDissotis conqolensisTristemma leiocalvxLJTW 0023UTW 0481SEGC 319

MELIACEAECarapa proceraEntandophraqma candoleiEntandophraqma utileLovoa trichilioidesTrichiIia monadelphaTrichilia prieureana

MENISPERMACEAESEGC 431

MIMOSACEAEAlbizia ferruqinea?Calpocalvx SP. LJTW 0611

L L FHWL; PH

S; L S; L FHS; L S; L FHS; L S; L CCFHS; L S; L FH

WL; PHP FH

WH L; PF; L; PF; L; P

F; L FHF; L FH

WH P

DHSHSHMFH

H DMFL L

100 600FTCCF * 8TCCF *TF * *T

S MF 40T880MFT

F/SFL

MF *TCCF 640T

LIFE-FORM 1 PLANT PART EATEN 2 HABITAT 3 DENSITY (individuals/km2) 4


> 10 cm dbhMF CCF

> 70 cm dbhMF CCF

F/STCathormion altissimumCvlicodiscus qabunensisDicrostachys cineraEntada qiqasFillaeopsis discophoraNewtonia leucocarpaParkia bicolorParkia filicoideaPentaclethra eetveldeanaPentaclethra macrophvllaPiptadeniastrum africanumSamanea leptophvllaTetrapleura tetraptera

MORACEAEDorstenia aff. barteriFicus barteriFicus bubuFicus elasticoidesFicus inqensFicus kimuenzensisFicus macrospermaFicus mucusoFicus ovataFicus politaFicus pseudomanqiferaFicus recurvataFicus surFicus thonninqiiFicus variifoliaFicus sp. SEGC 443Milicia (Chlorophora) excelsaMusanqa cecropioides

* 80 * 8T FS (D)SMFL

*F 4T2 86 4

T F20 80F FT F20 *F F MF *T 4640 240700 440

L F 14 1236 4430 16

T LT L L; FL F

40T S FCCF 40T 4

40 *T F 2 *W (CCF)HMFEp F FF/SEp FFMFEp

T MFFCCFL

F F MFEpF; L F MFT

R (S)TMFEp F FF/SFEp

F; L F MF * 2EpR (S)T

F F MFEpL; B L MF *T 2

Ep F F MFL; B; G F/S; MF 80 *T 6 4

*T D

1

PLANT PART EATEN 2 HABITAT 3 DENSITY (individuals/km2) 4> 10 cm dbh

MF CCF

1FAMILY / Species LIFE-FORMGorilla Chimp > 70 cm dbh

MF CCF

MF (W)MF (W)

Mvrianthus arboreusTreculia africanaTreculia obovoidea

MYRISTICACEAECoelocarvon preussiPvcnanthus angolensisScyphocephalium ochocoaStaudtia qabonensis

MYRTACEAEPsidium quineenseSvzyqium sp. LJTW 31Svzyqium sp. LJTW 0147Svzyqium/Euqenia sp. LJTW 0381

NYMPHAEACEAELJTW 0815

OCHNACEAECampvlospermum elonqatumLophira alataOuratea flavaOuratea cf. mvrioneura? Ouratea sp. UTW 0432Rhabdophvllum calophvllumSauvaqesia erectaLJTW 0772UTW 0882

OLACAŒAECoula edulisDioqoa zenkeriHeisteria parvifoliaOnqokea gorePtvchopetalum petiolatumStrombosia pustulata

T F; B FT S S; FLT S S CCF 200

CCFT 360 •k

T F F 140 12020 920

44 28T S; L CCF 1726

CCF * 200T F *S F F ST *MF 2S CCF

CCF kT *W (S)H

T MFT G F/S; MF

F/S; MFF/S; MF

2740 *L 90 8T *TT MF

MF *TSH

S CCFS S

CCFT 1760 60CCFT

F; L 120 24060 80

T F F 4 414 12T FL

CCFT F 160FCCF 400T

HABITAT 3 DENSITY (individuals/km2) 4> 10 cm dbh > 70 cm dbh

MF CCF MF CCF

LIFE-FORM 1 PLANT PART EATEN 2


2720120 2000

CCFStrombosia zenkeriStrombosiopsis tetrandra

OLEACEAELinociera aff. mannii

OXAUDACEAEBiophvtum petersianumBiophvtum talbotii

PALMAECalamus sp. SEGC 554Elaeis quineensisEremospatha cabraePodococcus barteriRaphia sp. SEGC 555

PANDACEAECentroplacus glaucinusPanda oleosa

PANDANACEAEPandanus sp. SEGC 556

PAPILIONACEAEAaanope impressaAganope sp.Crotularia pallidaDalberqia aff. rufaDesmodium ramosissimumDesmodium scorpiurusEriosema glomeratumErvthrina thollonianaIndiqofera coniuqata var. occidentalisLeptoderris hyparqyreaMillettia laurentiiMillettia sanaqana

Millettia conraui

TCCFT

F/ST

SHW (R)H

F/S; MFF/S; MF

P180F; PP

F; L; P MFP F; PCCFPW (S)P

CCF 100 2720T20 *F *T

W (S)T

CCFLL MF

SSL; B; FL F/SL

SHSHSsW (MF) •kTSHF/SLCCFT 4F/S 40L; B LTF/ST

LIFE-FORM 1 PLANT PART EATEN 2 HABITAT 3 DENSITY (individuals/km2) 4> 10 cm dbh > 70 cm dbh

MF CCF MF CCF

FAMILY / SpeciesGorilla Chimp

MF (W)Millettia sp. UTW 0038Platvsepalum sp. LJTW 0603Pterocarpus soyauxiiTephrosia purpureaUraria pictaZornia latifoliaUTW 0469UTW 0591UTW 0599LJTW 0602LJTW 0609LJTW 0720LJTW 0950SEGC 395

PASSIFLORACEAEBarteria fistulosaParopsia qrewioidesLJTW 0360

PONTEDERIACACEAEHeteranthera callaefolia

PTERIDOPHYTEAdiantum voqeliiArthropteris orientalisAsplénium africanumAsplénium emarainatumAsplénium hemitomumAsplénium iaundeenseAsplénium variabileBlotiella curroriBolbitis acrostichoidesBolbitis auriculataBolbitis qabonensis

20TCCF 240L

S; FLT S; L; FL F 80 68 20SHSHSH

S F/SCCF 80LCCFL 40CCF 40LCCF 320TCCF (W)T

T F/S *L B MF

T F F/S; MF; DF/S; R

520 600FT

CCFL F

W (MF)H

MF (W)R (MF)R (MF)R (MF)R (MF)R (MF)W (CCF)

FFFFFFFF W

W (MF)FWFMFF



LIFE-FORM 1 PU\NT PART EATEN 2 HABÎTAT 3Gorilla Chimp

FAMILY / Species

Bolbitis heudelotiiCeratopteris cornutaChristella dentataCvathea cameroonianaCvclosorus dentatusDicranopteris linearisDiplazium sammatiiDiplazium welwitschiiLastreopsis curroriLomariopsis hederaceaLomariopsis conqoensisLomariopsis sp. SEGC 536Lycopodium cernuumLycopodium microphvllumMarattia fraxineaMicroaramma owariensisMicrolepia speluncaeMicrosorium punctatumNephrolepis biserrataNephrolepis undulataOleandra distentaPellaea donianaPellaea holstiiPhvmatosons scolopendriaPityroqramma calomelanosPlatvcerium stemariaPteris atrovirensPteris similisPteris acanthoneuraPteris sp. SEGC 528Selaqinella cathedrifoliaSelaqinella mvosurus

WFW (MF)W (MF)

FF

WTfW (MF)FSFW (MF)W (CCF)

FFF WF (Ep) CCF

CCFFF (Ep) CCF

MFFF/SFWFR (MF)W (MF)R (MF)

FFF

F/SFF/SFR (MF)F

F RR (MF)F/S;MF

FF (Ep)

SFF (Ep) MF

SFW (MF)F

F MFF MF

W (MF)FF/SF

LIFE-FORM 1 PLANT PART EATEN 2 HABrTAT 3 DENSITY (individuals/km2) 4> 70 cm dbh

MF CCF

FAMILY / Species> 10 cm dbh

MF CCFGorilla Chimp

WFSelaqinella versicolorSelaqinella voaeliiTrichomanes ballardianumTrichomanes cupressoidesTrichomanes quineenseTrichomanes cf.manniiTriplophvllum buchholziiTriplophvllum protensumTriplophvllum secundiformeTriplophvllum sp. SEGC 469Triplophvllum voqeliiVittaria quineensis

RHIZOPHORACEAEAnisophyllea sp. LJTW 0475Anopyxis klaineanaCassipourea conqensisPoqa oleosa

RUBIACEAEAidia ochroleucaAtractoqyne qaboniiBertiera batesiiBertiera mildbraediiBertiera sp. LJTW 0139Borreria pusillaBrenania brieyiCanthium sp. LJTW 0297Cephaelis sp. LJTW 0202Corvnanthe mavumbensisCrossoptervx febrifuqaDictvandra arborescensDiodia scandensFeretia sp. LJTW 0646

MFFWFWFRFRFWFMFFWFMFFCCFF

F (Ep) CCF

80CCF 4T* ieF 2T

180RTCCF * 4T

F/S *TF MF; D

W (CCF)L FS

MFTCCFSSHMFTCCFSF/SS

340 2240L FTSSMFTMFHF/SS

HABrrAT 3 DENSITY (individuals/km2) 4


LIFE-FORM 1 PLANTT PART EATEN 2Gorilla Chimp

FAMILY / Species

CCFS?Gaertnera sp. UTW 0316Geophila afzeliiHeinsia crinitaLeptactina cf. arnoldianaMassularia acuminataMitacarpus scaberMitraqyna ciliataMussaenda cf. debeauxiiMussaenda cf. tenuifloraMussaenda sp. UTW 0581Nauclea didderrichiNauclea latifoliaNauclea vanderauchtiiOldenlandia corvmbosaOldenlandia lancifoliaOtomeria quineensisOtomeria micranthaOxvanthus unilocularis?Oxvanthus sp. UTW 0903Pauridiantha aff. dewevreiPauridiantha efferataPauridiantha floribundaPausinvstalia iohimbePausinvstalia macrocerasPavetta hispidaPavetta aff. viridilobaPavetta SP. LJTW 0006Pavetta sp. UTW 0205?Pavetta sp. UTW 536Porterandia cladanthaPseudosabicea batesiiPseudosabicea floribunda

MFHF (W)TDT

40*FTSH

80 120 2 16WB; LTDFLDLMFL

20 600 2 4FFFTSF FT

60 *F W 4FTSHSSMFHSHF/STMFTF/STD; F/SD; F/S

220T100 12020 200

TFT

20MFTCCFS L

20MFL; FLTCCFHF/SS

40MFT120 *D; MFFF; LT

CCFHDL

DENSITY (individuals/km^) 4> 10 cm dbh > 70 cm dbh


FAMILY / Species

MF CCFMF CCF

DPseudosabicea mildbraediiPsvchotria peduncularisPsvchotria voqelianaPsvchotria sp. LJTW 0012Psvchotria sp. LJTW 0171Psvchotria sp. LJTW 0174Psvchotria sp. LJTW 0209Psvchotria sp. LJTW 0638Psvchotria sp. LJTW 0657Rothmannia whitfieldiiRutidea dupuisiiSabicea efulenensisSabicea mollisSacosperma paniculatumTarenna confertaTarenna eketensisTarenna sp. LJTW 0272Tricalvsia anomalaTricalvsia macrophvllaTricalvsia cf oliqoneuraTricalvsia cf pallensTricalvsia sp. LJTW 0081Tricalvsia sp. LJTW 0082Trichostachvs aurea

HMFS FF

F F/SS FF/STCCFSCCFH

S MFS MF

CCFSMFT

F FLMFLMFLF/SLDTF/SLCCFTDTMF (W) *TMFSMFSCCF (W)SMFSMFHF/SUncaria africana

LJTW 0098LJTW 0372LJTW 0396LJTW 0417LJTW 0433LJTW 0628LJTW 0638

LDTMFSCCFTMFSCCFSR; W (F/S)TMFS


> 10 cm dbh > 70 cm dbhLIFE-FORM 1 PU\NT PART EATEN 2 HABITAT 3


MF CCF MF CCF

MF (W)LJTW 0642UTW 0646LJTW 0662UTW 140UTW 170LJTW 387LJTW 514UTW 528LJTW 566SEGC 288SEGC 291SEGC 508

RUTACEAEAraliopsis sp. UTW 353Citrus sp. SEGC 557Faqara TmacrophvllaFaqara tessmanniiTeclea aff. verdoonianaVepris cf. lousii

SAPINDACEAEBliqhia welwitschiiChvtranthus talbotiiEriocoelum macrocarpumEriocoelum ?paniculatumEriocoelum sp. UTW 132Ganophvllum qiqanteumLecaniodiscus cupanoidesPlacodiscus cf. opacus

SAPOTACEAEBaillonella toxispermaDonella oqowensisDonella ?pruniformis

TF/SSST

40MFTT D

40CCFT40CCFT40CCFT

20MFTF MFLF DL

S L MF

CCF 80TF/ST FF/S; MF *TCCF 40T

-kT R20RT

+MFTMF (W)F/S; MF

T540 280T

MFT20MFT80MF 20T F F

MF 100F FTCCF 80T

CCF 40 2 4T F FF/S *F FTCCF 80T



LIFE-FORM 1 PUNT PART EATEN 2


8CCF 40T FGambeva africanaGambeva subnuda? Lectomedoxa sp. LJTW 237Letestua durissimaManilkara fouillovanaOmphalocarpum procerumPachvstela brevipesLJTW 0228

SCROPHUURIACEAECvcnium camporum

SCYTOPETAUCEAEScvtopetalum sp. LJTW 0017

SIMAROUBACEAEOdvendvea qabonensisQuassia africana

SMIUCACEAESmilax kraussiana

SOUNACEAESchwenckia americana

STERCULIACEAECola lizaeCola mahoundensisCola sp. SEGC 332Leptonvchia echinocarpaNesoqordonia papaveriferaPteryqota ?bequaertiiScaphopetalum blackiiSterculia traqacantha

STYRACACUEAfrostvrax lepidophvllus

F*40F FT F2MFT

ieMF 4TCCFFT F

* * *FT*F RT F

160CCFFT F

SH

120 240FF FT

*CCFTFS

F/SL

SH

7460F; L; P MF 8T FCCF (W) 80TMF *T

S MF* •kFT

CCFTCCF 160T

40MF 2T L

80CCFT

THYMEUEACEAE

PU\NT PART EATEN 2 HABITAT 3 DENSITY (individuals/km2) 4


1FAMILY / Species LIFE-FORMGorilla Chimp

CCFSDicranolepis buchholzii?Dicranolepis sp. LJTWOctolepis decalepis

TILIACEAEClappertonia ? ficifoliaDuboscia macrocarpaTriumfetta cordifolia

ULMACEAECeltis tessmanniiCeltis sp. LJTW 444

VERBENACEAEVitex donianaVitex Trivularis

VIOLACEAERinorea ilicifoliaRinorea sp. LJTW 0055Rinorea sp. LJTW 0132Rinorea sp. LJTW 0221Rinorea sp. LJTW 0596Rinorea sp. LJTW 0656Rinorea sp. LJTW 0899

VITACEAECissus aff. barteriCissus dinklaqeiCissus leonardi

VOCHYSIACEAEErismadelphus exsul

ZINGIBERACEAEAframomum lonqipetiolatumAframomum sp. ?nov SEGC 152Aframomum sp. SEGC 239Aframomum sp. SEGC 323

MFSMFS

S; DHF; S 40 120F; S F 2T

Ss200 8 28F FT F; L

CCF 40T

F/S •kFT FCCF 80T 4

S CCFW (CCF)W (MF)W (CCF)W (CCF)

*T*T

40T120T

S CCFS CCF

L DF; L F + *L F

L F MF

CCF 40T 8

F/SH F; P F; PH F; P F; P MFH F; P F; P F

CCF; DH F; P F; P

LIFE-FORM 1 PLANT PART EATEN 2 HABITAT 3 DENSITY (individuals/km2) 4



Costus aferRenealmia cincinnataRenealmia macrocolea

H P P MFH P FH P P F

1 See Methods for definition of Life-forms;2 L= leaf; F=fruit; P= pith; S= seed; FL = flower; B= bark; G= galls;3 See Methods for definitions of Habitat-types;4 Density of trees calculated from transect sample: MF > 10 cm dbh, 5 Ha; CCF > 10 cm dbh, 2.5 Ha: MF >70 dbh, 50 Ha; CCF > 70dbh, 25 Ha (see text). * indicates present as a tree of this size but not recorded in transect sample.

Table 2. Top 10 species in Marantaceae Forest and Closed Canopy Forest at Lopé in terms ofbasal area and stem density

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

Aucoumea klaineana BURSERACEAESTERCULIACEAEOCHNACEAEMIMOSACEAEBURSERACEAEMIMOSACEAEEBENACEAECAESALPINIACEAEANNONACEAEMYRISTICACEAEMELIACEAEOLACAŒAEEBENACEAEEBENACEAEEUPHORBIACEAEMYRISTICACEAEPANDACEAEANNONACEAEBURSERACEAEOLACAŒAECAESALPINIACEAECAESALPINIACEAEOLACAŒAEMIMOSACEAECAESALPINIACEAE

1 22Cola lizae 2 1Lophira alata 3 3Pentaclethra macrophvlla 4 9Dacrvodes buettneri 5 81Pentaclethra eetveldeana 6 10Diospyros polystemon 7 5Flvlodendron qabunense 8Xvlopia quintasii 9 4Pycnanthus anqolensis 10Trichilia cf. prieureana 7Strombosiopsis tetrandraDiospyros dendoDiospyros zenkeriMaprounea membranaceaScyphocephalium ocochoaCentroplacus qlaucinas

7 468

103

1Xvlopia aethiopica 7Santiria trimera 9 3Coula edulis 4 5Auqouardia letestui 65Sindoropsis le-testui 6Strombosia zenkeri 8 1Cylicodiscus qabonensis 10Dialium sovauxii 9

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

Chimpanzee Food SpeciesPercentage

Gorilla Food SpeciesNumber Percentage

N° SpeciesHabitat-typeNumber

24.242 28.2 36Marantaceae ForestClosed Canopy ForestForestForest/savanna edgeSavannaWaterDisturbed areasRocky areas

14913.811.2 21152 17

50 55.658.990 5321.3 19 18.4103 22

2 3.03 4.5672 3.717.0953

11.817.6 46346 21.421.428 6

14023.4 20.7158TOTALyAVERAGE 676

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